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HARVARD    UNIVERSITY 

Library  of  the 

Museum  of 

Comparative  Zoology 


S^/^/^~S/an  Oi^^o] 


TRANSACTIONS 

OF  THE 

SAN  DIEGO  SOCIETY  OF  NATURAL  HISTORY 

MUS.  COMF.  ZOOL. 
LiBWARY 


AU6  1  9 1975 

HARVARD 


VOLUME  17 
1972-1975 


PRINTED  FROM  THE 
W.  W.  WHITNEY  PUBLICATION  ENDOWMENT 


CONTENTS 

1.  Sound  production  and  other  behavior  of  Southern  Right  Whales, 
Eubalena  glacalis.  By  William  C.  Cummings,  James  F.  Fish,  and  Paul  O. 
Thompson.  15  March  1972 1.14 

2.  Eastern  Pacific  Snake-Eels  of  the  genus  Callechelys  (Apodes:  Ophich- 
thidae).   By  John  E.   McCosker  and  Richard  H,   Rosenblatt.   25  April 

1972 15-24 

3.  Paleontology  and  paleoecology  of  the  San  Diego  Formation  in  northwestern 

Baja  California.  By  Robert  W.  Rowland.  9  June  1972 25-32 

4.  Seismic  risk  in  San  Diego.  By  Robert  B.  McEuen  and  Charles  J.  Pinckney. 

19  July  1972 33-62 

5.  The  feeding  techniques  of  Stilt  Sandpipers  and  Dowitchers.  By  P.  J.  K. 
Burton.  16  August  1972 63-68 

6.  Thoracic  Cirripedia  from  Guyots  of  the  Mid-Pacific  Mountains.  By  M.  V. 
Lakshmana  Rao  and  William  A.  Newman.  31  August  1972 69-94 

7.  A  new  Mitrid  from  the  western  Atlantic.  By  George  E.  Radwin  and  Loyal 

J.  Bibbey.  31  August  1972 95-100 

8.  Diagnoses  of  new  Cyprinid  fishes  of  isolated  waters  in  the  Great  Basin  of 
western  North  America.  By  Carl  L.  Hubbs  and  Robert  Rush  Miller.  29 
September  1972  101-106 

9.  Patterns  of  larval  development  in  Stenoglossan  Gastropods.  By  George  E. 

Radwin  and  J.  Lockwood  Chamberlain.  12  March  1973 107-118 

10.  A  marine  invertebrate  faunule  from  the  Lindavista  Formation,  San  Diego, 
California.  By  George  L.  Kennedy.  28  March  1973 1 19-128 

11.  Post-Batholithic  geology  of  the  Jacumba  area,  southeastern  San  Diego 
County,  California.  By  John  A.  Minch  and  Patrick  L.  Abbott,  10  April 

1973 129-136 

12.  Revision  of  the  coral-inhabiting  barnacles   (Cirripedia:   Balanidae).   By 

Arnold  Ross  and  William  A.  Newman.  20  April  1973 137-174 

13.  Biology,    geographical    distribution,    and    status    of   Atteva    exquisita 
(Lepidoptera:  Yponomeutidae).  By  Jerry  A.  Powell,  John  Adams  Comstock 

and  Charles  F.  Harbison.  14  May  1973 175-186 

14.  Life    history    of  the    Western    North    American    Goby,    Coryphopterus 
nicholsii  (Bean).  By  James  W.  Wiley.  30  October  1973 187-208 

15.  A    new    Platvdoris    (Gastropoda:    Nudibranchia)    from    the    Galapagos 
Islands.  By  David  K.  Mulliner  and  Gale  G.  Sphon.  12  April  1974 209-216 

16.  The  distribution  and  ecology  of  marine  birds  over  the  continental  shelf  of 
Argentina  in  winter.  By  Joseph  R.  Jehl,  Jr.  28  June  1974 217-234 

17.  Mexican  species  of  the  genus  Heterandria.  subgenus  Pseudoxiphophorus 

(Pisces:  Poeciliidae).  By  Robert  Rush  Miller.  28  June  1974 235-250 

18.  Lithostratigraphic    variations    in    the    Poway    Group    near    San    Diego, 
California.  By  Gary  L.  Peterson  and  Michael  P.  Kennedy.  6  December 

1974  251-258 

19.  The  autecology  of  Xantusia  henshawi  henshawi  (Sauria:  Xantusiidae).  By 

Julian  C.  Lee.  22  April  1975 259-278 

20.  A  catalogue  of  Muricacean  generic  taxa.   By  George  E.   Radwin  and 
Anthony  D'Attilio.  16  May  1975 279-292 

21.  Bloods  circulation  in  four  species  of  barnacles  (Lepas.    Conchoderma: 
Lepadidae).  By  Bryan  R.  Burnett.  20  June  1975  293-304 


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HARVARD 
UNIVERSITY 

SOUND  PRODUCTION  AND  OTHER  BEHAVIOR 

OF  SOUTHERN  RIGHT  WHALES,  EUBALENA  GLACIAUS 


WILLIAM  C.  CUM  MINGS,  JAMES  F.  FISH, 
AND  PAUL  O.  THOMPSON 


TRANSACTIONS 

OF  THE  SAN  DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  1  15  MARCH  1972 


SOUND  PRODUCTION  AND  OTHER  BEHAVIOR 

OF  SOUTHERN  RIGHT  WHALES,  EUBALENA  GLACIALIS 

WILLIAM  C.  CUMMINGS,  JAMES  F.  FISH,  AND  PAUL  O.  THOMPSON 


ABSTRACT.— In  late  June  and  early  July,  1971,  we  recorded  underwater  sounds  from  southern  right 
whales  in  Golfo  San  Jose,  Argentina.  The  most  common  sound,  a  belch-like  utterance,  averaged  1.4  sec  in 
duration,  with  most  of  its  energy  appearing  below  500  Hz.  Levels  of  this  strong  signal  ranged  from  172  to 
187  dB,  re  1  /^.N/nv  at  1  m.  The  whales  also  produced  two  kinds  of  low-frequency  moans.  Simple  moans 
had  a  narrow  band  of  frequencies  (centered  at  about  160  Hz)  without  appreciable  frequency  shifts.  Com- 
plex moans  exhibited  a  wider  band  width  (centered  at  about  235  Hz),  extensive  frequency  shifts,  and  over- 
tones. Other  sounds  were  categorized  as  pulses,  0.06-sec  bursts  extending  from  30  to  21(X)  Hz,  and  mis- 
cellaneous sounds,  comprising  numerous  phonations  below  1950  Hz  that  varied  in  length  from  0.3  to  1.3  sec. 
There  was  no  periodic  occurrence  of  sound  production  other  than  that  related  to  the  appearance  of  whales 
in  the  recording  area  at  low  tide. 

When  presented  underwater  playbacks  of  killer  whale  sounds,  a  right  whale  exhibited  a  behavior  pat- 
tern called  "spyhopping,"  but  there  was  no  obvious  avoidance.  An  attack  by  five  killer  whales  on  two  other 
right  whales  ended  after  25  min,  apparently  without  serious  harm  to  the  right  whales.  A  common  behavior 
of  these  southern  right  whales,  "headstanding,"  may  be  associated  with  bottom  feeding.  Patterns  of  breath- 
ing varied  considerably  depending  upon  associated  activities. 

Local  citizens  reported  that  right  whales  appear  in  Golfo  San  Jos6  and  nearby  Golfo  Nuevo  each  year 
in  late  June.  They  are  most  numerous  in  late  August  and  September,  and  they  disappear  in  November.  We 
saw  several  consorting  pairs  that  appeared  to  be  courting,  but  no  copulations  and  no  very  young  whales. 

This  report  describes  the  underwater  sounds  and  behavior  of  southern  right  whales 
in  Golfo  San  Jose,  Argentina,  in  June-July,  1971.  A  brief  summary  of  some  of  this  work 
was  presented  earlier  (Cummings  et  al.,  1971). 

The  right  whale  is  virtually  world-wide  in  distribution  (Hershkovitz,  1966).  Although 
three  subspecies  have  been  recognized  (australis:  Southern  Hemisphere;  glacialis:  North 
Atlantic;  sind  japonica:  North  Pacific),  their  validity  is  questionable  (Rice  and  Scheffer, 
1968).  In  this  report  we  call  this  cetacean  the  "southern  right  whale"  with  no  attempt  to 
evaluate  any  subspecific  rank. 

Right  whales,  so  named  by  whalers  because  these  animals  have  a  high  oil  content 
and  float  when  dead,  attain  a  maximum  length  of  18  m.  They  may  be  identified  at  sea  by 
a  characteristic  V-shaped  blow  (Fig.  lA),  light-colored  horny  protuberances  on  the  upper 
snout  which  include  the  bonnet  (see  Ridewood,  1901,  and  Matthews,  1938,  for  detailed 
descriptions),  and  by  the  lack  of  both  a  dorsal  fin  and  throat  grooves.  Compared  with 
other  species  of  great  whales,  right  whales  are  very  rotund  (Figs.  IB,  2).  Although  killing 
of  these  animals  has  been  prohibited  for  many  years,  the  population  remains  exceedingly 
small  in  many  regions  (Ohsumi  et  al.,  1971;  Doi  et  al.,  1971),  and  the  species  may  be  in 
danger  of  extinction  through  overharvesting. 

Southern  right  whales  have  long  been  known  to  breed  in  bays  and  other  sheltered 
waters  (Scammon,  1874),  and  recent  information  indicates  the  same  is  true  of  northern 
animals  (pers.  comm.  W.  E.  Schevill,  Woods  Hole  Oceanogr.  Inst.).  In  mid-July,  1969.  Gil- 
more  (1969)  located  20  to  25  right  whales  that  were  courting  and  presumably  mating  in 
Golfo  Nuevo,  Argentina.  On  his  advice,  our  attempts  to  record  the  vocalizations  of  this 
rare  cetacean  and  to  observe  its  behavior  were  concentrated  in  this  region. 

Little  is  known  about  vocalizations  of  the  right  whale.  In  discussing  this  whale's  be- 
havior after  being  harpooned,  Scammon  (1874)  wrote  that,  "after  going  a  short  distance, 
it  frequently  stops,  or  'brings  to,'  'sweeping'  as  it  is  said,  'from  eye  to  eye,'  and  at  the  same 
time  making  a  terrific  noise  called  'bellowing,'  this  sound  is  compared  to  that  of  a  mam- 
moth bull,  and  adds  much  to  the  excitement  of  the  chase  and  capture."  Schevill  and  Wat- 
kins  (1962)  presented  a  description  and  a  recording  of  low-frequency  moaning  sounds  of 
northern  right  whales.  Cummings  and  Phihppi  (1970)  reported  low-frequency  (20-174  Hz) 

SAN  DIEGO  see.  NAT.  HIST.,  TRANS.  17(1):  1-14.  15  MARCH  1972 


pulses  and  moans  tentatively  identified  as  being  from  northern  right  whales.  These 
sounds,  recorded  off  Newfoundland  in  December,  1965,  appeared  in  repetitive,  11-min  to 
14-min  stanzas  that  were  separated  by  8  to  10  min.  Each  stanza  was  composed  of  numer- 
ous signals  appearing  in  a  precise  sequence  that  was  repeated  in  the  next  stanza.  Payne 
and  McVay  (1971)  described  a  similar  repetitive  phenomenon,  "songs,"  from  the  hump- 
back whale,  Megaptera  novaeangliae. 

MATERIAL  AND  METHODS 

This  research  was  largely  carried  out  from  the  National  Science  Foundation's  re- 
search ship,  HERO,  a  38-m  vessel  that  is  managed  by  the  United  States  Antarctic  Re- 
search Program.  We  left  Punta  Arenas,  Chile,  on  11  June  and  proceeded  north  along  the 
coast  as  far  as  Bahi'a  Blanca,  Argentina  (39°N),  returning  to  Punta  Arenas  on  16  July.  Al- 
though other  marine  mammals  were  sighted  along  the  coast,  we  did  not  see  right  whales 
in  areas  other  than  Golfo  San  Jose,  an  enclosed  bay,  44  X  20  km,  on  the  north  side  of  the 
Valdes  Peninsula  (Fig.  3).  There,  from  21-24  June  and  from  1-8  July,  we  observed  about 
10  southern  right  whales,  this  estimate  based  on  searching  the  entire  Gulf  on  each  of  sev- 
eral days. 

The  deepest  area  of  the  Gulf  is  about  82  m,  but  most  of  the  whales  were  observed 
near  shore  in  less  than  37  m.  Because  the  Gulf  is  protected  and  shallow,  we  generally  ex- 
perienced moderate  or  calm  seas  of  State  2  or  less.  Air  temperatures  ranged  from  6.1  to 
8.3°C.  We  took  bathythermographs  in  several  areas  of  the  Gulf  and  found  isothermal 
conditions  averaging  9.4° C.  The  ship's  small  boats  occasionally  were  used  for  short  excur- 
sions close  to  the  whales,  although  we  generally  kept  our  distance  so  as  not  to  in- 
tentionally crowd  or  molest  them. 

Underwater  recordings  were  made  with  much  the  same  system  described  by  Calde- 
ron  and  Wenz  (1967).  Essentially,  our  recording  system  consisted  of  an  acceleration-bal- 
anced hydrophone,  flexible  spar  buoy,  floating  cable,  calibrating  device,  sound-level  me- 
ter, magnetic  tape  recorder,  and  monitoring  equipment.  Instruments  used  in  playing 
sounds  to  right  whales  included  a  tape  recorder  (Uher  4200),  preamplifier  (Bogen  BT- 
35A),  high-power  amplifier  (Optimation  PA  250  AC),  and  an  underwater  sound  projector 
specially  designed  by  Wesley  L.  Angeloff  of  the  Naval  Undersea  Research  and  Devel- 
opment Center.  The  frequency  response  of  the  playback  system  was  ±  5dB  from  650  to 
3100  Hz,  limited  by  the  projector.  Response  of  the  receiving  system  was  ±5  dB  from  25  to 
15000  Hz. 

To  obtain  good  recordings  of  low-frequency  mysticete  sounds,  we  have  found  it  best 
to  use  a  hydrophone  that  is  relatively  stationary  in  the  water  column  and  has  a  good  low- 
frequency  response,  but  not  all  the  way  down  to  0  Hz.  The  response  must  be  "roUed-off' 
in  the  low  frequencies  for  use  under  normal  sea  conditions  in  order  to  prevent  the  recep- 
tion of  low  frequency  noise.  Such  a  hydrophone  is  still  unsuitable,  however,  for  towing 
from  a  moving  vessel. 

A  hydrophone  is  designed  to  respond  to  changing  pressure.  Pressure  changes  caused 
by  towing  a  hydrophone,  or  those  resulting  from  the  vertical  excursions  so  characteristic 
of  moderate  or  greater  sea  states,  will  usually  produce  excessive  low-frequency  noise. 
Coupled  with  the  high  sound  pressure  of  low- frequency  ambient  noise  (Wenz,  1962),  ac- 
celeration and  flow  noise  of  this  type  will  easily  mask  a  low-frequency  mysticete  signal. 
Experience  has  taught  us  that  such  recordings  have  very  low  signal-to-noise  ratio.  More 
often,  they  are  rendered  useless  by  intermittent  or  even  continuous  blocking  of  the  hydro- 
phone's preamplifier.  The  dynamic  range  of  response  of  this  preamplifier  generally  will 
not  accommodate  that  of  the  electrical  energy  from  sound  pressures  imposed  on  a  hydro- 
phone under  the  above  circumstances.  The  blocking  occurs  when  the  preamplifier  is  over- 
driven by  an  input  signal  that  greatly  exceeds  the  maximum  input  level  designed  for  the 
amplifier. 

To  reduce  these  vertical  and  horizontal  movements,  we  use  an  inflatable  buoy  for 
flexibility  and  a  450-m  buoyant  cable  that  is  let  out  as  fast  as  the  ship  drifts.  The  cable 
floats  by  means  of  a  buoyant  sheath  that  is  molded  around  the  conductors.  In  com- 
bination with  a  good-quality  hydrophone  having  low-frequency  rolloff  and  acceleration 


Figure  1.  A,  V-shaped  blow  (exhalation)  of  a  southern  right  whale;  B,  quartering  view. 


balancing,  this  system  works  well  for  us  most  of  the  time.  Other  investigators  have  dealt 
with  the  problem  in  other  effective  ways  (Watkins,  1966). 

The  right  whale  sounds  were  recorded  under  quiet  conditions,  with  all  engines  shut 
off.  However,  the  ship's  generator  was  needed  during  playback  experiments.  Most  of  our 
recordings  were  made  while  the  ship  was  quietly  lying  to,  0.2  to  1.5  km  from  the  whales. 
Whenever  possible,  we  kept  an  account  of  the  whales'  behavior  during  the  recordings, 


both  in  a  written  log  and  as  verbal  comments  on  magnetic  tape.  Recording  times  of  con- 
tacts varied  from  10  to  120  min,  after  which  the  whales  either  moved  out  of  an  area  or  we 
simply  stopped  the  recording. 

PRONATIONS  AND  SOUND  PLAYBACK 

Southern  right  whales  made  several  different  types  of  powerful,  low-frequency 
sounds  resembling  belches,  moans,  and  pulses  (Fig.  4,  A-D).  They  also  produced  a  num- 
ber of  miscellaneous  low-frequency  sounds,  too  numerous  to  classify. 

By  far,  the  most  common  sound  was  a  belch-like  utterance  that  varied  from  0.9  to  2.2 
sec  in  duration  and  averaged  1.4  sec  with  principal  energy  centered  at  235  Hz.  Although 
the  frequency  ranged  from  30  Hz  to  about  2200  Hz,  the  major  portion  fell  below  500  Hz. 
Belch-like  sounds  often  ended  in  about  a  150-Hz  upward  frequency  shift  (Fig.  4A). 

The  first  portion  of  the  belch-like  sound  revealed  two  to  four  strong  overtones  with 
intervals  of  about  100  Hz.  Sound  pressure  levels  of  the  belch-like  sounds  were  172  to  187 
dB,  re  1  yuN/m^  (  =  72  to  87  dB,  re  1  /^bar)  at  1  m  from  the  source.  Source  levels  were  de- 
termined from  measurements  in  a  band  from  25  to  2500  Hz,  thus  including  all  fre- 
quencies oi  belch-like  sounds.  These  levels  were  derived  from  the  absolute  received  sound 
pressure  levels  at  the  hydrophone  (as  measured  in  the  laboratory  from  the  calibrated 
recordings)  and  the  estimated  distances  of  the  whales  from  the  hydrophone.  The  calcu- 
lations took  into  account  an  estimated  spreading  loss  of  6  dB  per  distance  doubled.  Atten- 
uation losses  were  regarded  as  negligible,  because  the  whales  were  so  close  and  their  calls 
so  low  in  frequency. 

Southern  right  whales  also  made  moaning  sounds  of  several  different  kinds.  Their 
moans  were  classified  into  two  basic  types,  simple  and  complex  (Fig.  4B).  Simple  moans 
had  sound  energy  that  was  confined  to  a  relatively  narrow  band  without  appreciable 
shifts  in  frequency.  Simple  moans  lasted  from  0.6  to  1.6  sec.  The  highest  frequency  noted 
was  320  Hz,  the  lowest  was  70  Hz,  and  the  region  of  principal  energy  was  about  160  Hz. 
Complex  moans  exhibited  a  wider  band  of  energy,  extensive  frequency  shifts,  overtones, 
and  a  longer  duration  compared  to  simple  moans.  The  highest  frequency  observed  among 
complex  moans  was  1250  Hz,  the  lowest  was  30  Hz,  and  the  region  of  principal  energy  was 
235  Hz.  The  duration  of  complex  moans  ranged  from  0.2  to  4.1  sec. 

The  third  most  common  of  the  major  types  of  right  whale  sounds  were  pulses  (Fig. 
4C).  These  sounds  extended  from  20  to  2100  Hz,  and  lasted  only  about  0.06  sec.  The  pul- 
ses frequently  occurred  in  conjunction  with  a  moan. 

The  remaining  sounds  consisted  of  numerous,  miscellaneous,  low-frequency  sounds 
that  varied  in  length  from  0.3  to  1.3  sec  (Fig.  4D).  All  of  these  were  below  1950  Hz. 

We  were  unable  to  associate  sound  production  with  any  specific  behavior.  The 
sounds  emanated  from  surfacing  as  well  as  from  diving  whales.  They  came  from  single 
whales  or  from  small  groups  of  two  to  three  individuals.  Although  some  of  the  right 
whales  may  have  been  feeding  and  others  presumably  were  courting  and  perhaps  mating, 
we  were  unable  to  associate  any  sounds  with  a  particular  activity.  In  one  instance  we  were 
in  a  small  rubber  boat,  close  to  a  surfacing  whale,  when  the  whale  produced  a  thun- 
derous, cavernous,  bellow  between  two  exhalations.  The  sound  was  clearly  audible  in  air 
and  may  have  been  the  same  type  of  sound  described  by  Scamrnon  (1874).  This  whale 
was  one  of  two  that  were  consorting  near  us,  in  very  shallow  water. 

Extensive  recordings  were  made  in  the  southeast  corner  of  Golfo  San  Jose  to  deter- 
mine if  there  was  any  diurnal  periodicity  in  sound  production.  These  recordings  were 
made  for  15  min  every  2  hrs,  beginning  at  1830  on  2  July  and  ending  at  1030  on  4  July. 
We  continued  to  listen  for  10  to  20  min  after  each  recording.  Most  of  the  right  whale 
sounds  on  these  recordings  occurred  close  to  the  three  low  tides  (Table  1),  a  phenomenon 
that  may  have  been  associated  with  the  appearance  of  right  whales  in  this  area  and  not 
necessarily  with  any  daily  rhythm  in  sound  production.  At  other  times  the  whales  moved 
along  shore,  either  toward  the  west  or  north.  There  was  no  indication  of  a  difference  in 
daytime  vs  nighttime  activity  in  sound  production.  In  this  general  vicinity  we  doubt  that 
whale  sounds  originating  more  than  about  2  km  away  would  have  been  detected,  because 


of  the  limited  propagation  which  could  be  expected  in  the  presence  of  the  coves,  shallow 
water,  and  disrupted  bottom. 

Table  1.  Occurrence  of  right  whale  sounds  on  nineteen  15-min  recordings. 


Time  of  Day 

Date 

1830 

2  July 

2030 

2  July 

2230 

2  July 

0030 

3  July 

0230 

3  July 

0430 

3  July 

0630 

3  July 

0830 

3  July 

1040 

3  July 

1245 

3  July 

1515 

3  July 

1945 

3  July 

2150 

3  July 

2345 

3  July 

0145 

4  July 

0345 

4  July 

0545 

4  July 

0830 

4  July 

1030 

4  July 

No.  of  Sounds 


Time  of  Low  Tide 


0 
0 
4 
3 

30 
0 
3 
1 
0 

28 
1 
4 
0 

10 

11 
1 
6 
0 

6' 


0112 


1348 


0200 


'Five  of  these  6  sounds  occurred  in  9  sec  and  appeared  to  be  a  series  of  sounds  from  a  single  right  whale. 

Our  earlier  experiments  had  shown  that  certain  marine  mammals  appeared  to  recog- 
nize underwater  sounds  of  killer  whales.  Migrating  gray  whales,  Eschrichtius  robustus,  off 
southern  California,  avoided  underwater  playbacks  of  killer  whale  "screams,"  (Cum- 
mings  and  Thompson,  1971);  and  playbacks  prevented  white  whales,  Delphinapterus 
leucas,  from  swimming  up  the  Kvichak  River  in  Alaska,  where  young  salmon  were  migra- 
ting to  the  open  sea  (Fish  and  Vania,  1971).  Thus,  playback  experiments  provide  a  new 
source  of  information  about  the  behavior  of  whales  in  their  natural  environment.  For  ex- 
ample, the  escape  reaction  of  the  gray  whale  to  killer  whale  sounds  can  be  used  to  test  the 
hearing  capabilities  of  this  species.  Also,  by  playing  tapes  of  certain  segments  of  killer 
whale  signals  it  may  be  possible  to  determine  which  parts  of  these  signals  induce  an  avoid- 
ance reaction. 

We  played  back  killer  whale  "screams"  to  a  group  of  three  southern  right  whales. 
HERO  was  1.8  km  from  the  beach,  in  12  m  of  water,  and  all  of  the  whales  at  first  were 
less  than  2  km  away.  The  underwater  sound  was  monitored  with  a  hydrophone.  We  ob- 
served the  whales  for  15  min  before  playback.  The  first  transmission  consisted  of  5  min  of 
random  noise,  as  a  control.  After  2  min  of  silence  we  transmitted  a  5-min  tone  (described 
by  Cummings  and  Thompson,  1971),  also  as  a  control.  After  3  min  of  silence  we  played 
back  5  min  of  prerecorded  killer  whale  sounds.  Sound  pressure  levels  of  the  playbacks, 
measured  in  situ  with  a  calibrated  system,  varied  from  159  to  163  dB,  re  1  /tN/m-  at  1  m 
from  the  source.  Generator  noise  from  the  ship  measured  134  dB,  over  the  effective  band- 
width of  the  recording  system,  at  the  same  location  and  time  of  the  playback  experiments. 

Only  one  whale  could  be  seen  well  enough  to  observe  its  reaction.  For  15  min  before, 
it  had  been  moving  slowly  back  and  forth  along  the  beach,  blowing  at  irregular  intervals. 
There  was  no  obvious  change  in  its  behavior  when  confronted  with  either  the  random 
noise  or  the  tone.  When  the  killer  whale  "screams"  were  played  back,  the  right  whale  ap- 
peared to  behave  as  before,  except  that  it  frequently  raised  its  head  out  of  the  water  in  a 
"spyhopping"  posture.  This  behavior  consisted  of  raising  the  head  vertically  out  of  the 
water  with  the  eyes  above  the  surface.  In  earlier  experiments,  Cummings  and  Thompson 
(1971)  found  that  gray  whales  fled  a  sound  source  for  an  appreciable  distance  and  then 
"spyhopped."  They  concluded,  as  did  Gilmore  (1958),  that  "spyhopping"  was  an  in- 
vestigative behavior. 


Figure  2.  Dorsal  view  of  a  southern  right  whale  heading  away  from  the  ship.  Note  the  widely  separated  blow 
holes  and  the  rotund  form  of  this  species. 


OTHER  BEHAVIOR 

On  4  July,  in  order  to  make  detailed  photographs,  we  accompanied  a  pair  of  right 
whales  that  apparently  had  been  courting.  We  had  just  finished  our  work  at  dusk,  and 
they  had  resumed  their  rolling  antics  near  the  surface,  when  a  group  of  killer  whales  ap- 
peared off  hero's  stern.  The  killer  whales  were  heading  away  from  the  right  whales; 
then,  suddenly,  they  whirled  around  and  swam  straight  towards  the  two  right  whales 
which  were  separated  by  about  45  m. 

When  the  killer  whales  were  about  70  m  away,  the  right  whales  came  together  so 
closely  that  they  appeared  to  be  touching  one  another.  As  soon  as  the  killer  whales  had 
reached  them,  the  right  whales  started  slashing  the  water's  surface  with  their  flukes  and 
flippers.  The  right  whales  were  then  blowing  every  10  to  20  sec,  twisting  and  turning  in 


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Figure  3.  Chart  showing  the  Valdes  Peninsula  of  Argentina. 

the  water  as  the  killer  whales  swam  around  them. 

We  were  too  far  away  to  see  whether  or  not  the  killer  whales  were  actually  biting  the 
right  whales.  However,  in  other  respects  it  appeared  to  be  a  full-fledged  attack.  On  at 
least  three  occasions,  one  or  the  other  right  whale  was  on  its  back,  thrashing  the  water's 
surface  with  its  flippers  and  flukes  at  the  same  time.  At  one  point,  the  right  whales  were 
completely  encircled  by  the  killer  whales.  The  most  impressive  of  the  large  whales'  defen- 
sive maneuvers  was  the  way  they  kept  together— rolling,  turning,  and  slashing  within  such 
close  quarters.  In  at  least  two  instances,  the  attacked  and  the  attackers  all  were  below  the 
surface  with  nothing  showing  but  a  slick  of  whirhng  water. 

We  moved  to  within  0.5  km  to  record  any  underwater  sounds  at  short  range.  How- 
ever, when  the  ship  had  stopped,  and  we  had  only  been  recording  for  a  short  time,  the 
killer  whales  left  their  prey  and  swam  toward  the  ship.  We  counted  five  killer  whales,  in- 
cluding one  very  young  animal  and  apparently  four  females.  The  attack  had  lasted  25 
min,  and  occurred  in  30  m  of  water. 

The  right  whales  then  moved  into  very  shallow  water  (7-11  m)  where  they  rolled  at 
the  surface,  more  slowly  than  before  the  attack,  exhibiting  a  notable  decrease  in  activity. 
There  were  no  signs  of  blood  or  other  evidence  of  physical  harm  in  the  vicinity  of  the  at- 
tack. 

We  recorded  for  about  3  min  before  the  attack  ended  and  for  15  min  afterward,  but 
obtained  no  underwater  sounds  from  either  species,  even  though  both  were  well  within 
range  of  the  hydrophone.  Gray  whales  and  white  whales  became  significantly  quieter 
when  confronted  with  killer  whale  sounds  (Cummings  and  Thompson,  1971;  Fish  and 
Vania,  1971).  Nevertheless,  even  in  a  recording  as  short  as  ours,  we  expected  some  phona- 
tions  from  the  loquacious  killer  whales  or  the  right  whales.  However,  under  these  circum- 


1.5- 


1.0- 


TIME,  sec 

Figure  4.  Sonagrams  of  sounds  from  southern  right  whales  recorded  in  Golfo  San  Jose,  Argentina.  Row  A,  two 
belch-like  sounds;  Row  B,  one  simple  and  two  complex  moans;  Row  C,  pulses  associated  with  simple  moans; 
Row  D,  two  examples  of  miscellaneous  sounds.  The  effective  bandwidth  of  the  analyzing  filter  for  these  sona- 
grams was  10  Hz. 


2  - 


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UJ 

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CO 


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6 

5  - 


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Figure  5.  Histogram  showing  the  time  du- 
ration between  consecutive  ventilations  of 
the  southern  right  whale.  Conditions  A, 
B.  and  C  explained  in  text. 


3  - 


2  - 


111 


CONSECUTIVE  VENTILATIONS 


Stances  neither  may  have  profited  from  being  noisy  (see  Schevill,  1964). 

We  saw  killer  whales  on  two  other  occasions  in  Golfo  San  Jose— a  group  of  three  on 
1  July,  in  the  mid  Gulf,  and  another  group,  of  six,  on  8  July,  near  the  east  end.  Evidently, 
these  animals  are  common  near  Valdes  Peninsula  as  several  observers  reported  seeing 
them  inside  and  outside  of  the  two  gulfs. 

Two  mussel  fishermen,  Jorge  Enrique  Ramirez  and  Jorge  Raul  Terenzi,  related  to  us 
that  their  Captains,  Roque  Godio  and  Calixto  Gerez,  had  also  witnessed  attacks  of  killer 
whales  on  right  whales  near  the  Valdes  Peninsula.  In  one  such  attack  they  related  that  five 
killer  whales  "hammered"  away  at  the  head  region  until  the  right  whale  opened  its 
mouth.  The  killer  whales  then  started  tearing  away  at  the  tongue.  The  area  was  colored 
with  blood,  the  killer  whales  left,  and  the  right  whale  was  lying  motionless  at  the  surface, 
apparently  dead. 

Apparent  pairs  of  right  whales  were  seen  on  several  occasions.  They  spent  much  time 
rolling  at  the  water's  surface,  exposing  bellies,  backs,  flukes  and  flippers,  and  occasionally 
"spyhopping."  Members  of  a  pair  were  often  very  close  to  each  other,  and  at  times  they 
appeared  to  be  in  physical  contact.  Some  of  this  behavior  may  have  been  associated  with 
courtship,  but  we  obtained  no  evidence  of  actual  mating.  Since  we  could  not  recognize  in- 
dividual whales,  we  could  not  determine  if  the  association  was  prolonged  for  more  than 
about  half  a  day. 

The  "headstanding"  posture  consisted  of  holding  the  flukes  upright  and  out  of  the 
water  for  periods  up  to  2  min.  During  this  time,  the  flukes  slowly  rocked  back  and  forth  or 
from  side  to  side,  occasionally  arching  downward  toward  the  water's  surface.  "Head- 
stands"  only  occurred  in  very  shallow  depths,  and  the  behavior  was  more  frequent  among 
single  whales. 

Right  whales  appeared  to  have  little  fear  of  the  ship  or  the  rubber  boats.  We  once 
ventured  as  close  as  9  m  with  a  rubber  boat,  and  neither  one  of  a  consorting  pair  showed 
any  apprehension.  Other  observers  have  had  the  same  experience  (Matthews,  1938). 
Moreover,  the  two  fishermen  who  were  interviewed  reported  that  they  once  were  awak- 
ened at  night  by  a  right  whale  rubbing  its  head  on  the  side  of  their  boat. 


10 


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Occasionally,  kelp  gulls,  Larus  dominicamis,  and  brown-headed  gulls,  L.  macuU- 
pennis,  landed  and  rode  on  whales  at  the  surface.  One  gull  that  landed  near  the  blowhole 
was  blown  off  by  the  next  expiration.  The  gulls  actually  pecked  at  the  whales,  and  possi- 
bly were  feeding  on  parasites. 

The  whales  we  studied  were  rather  sluggish.  Many  swam  very  slowly  at  the  surface 
for  long  distances— in  one  case  about  4  km  without  diving.  Apparently,  the  whales  were 
incapable  of  speeds  greater  than  about  14.8  km/hr  (8  knots).  HERO's  top  speed  of  18.5 
km/hr  (10  knots)  easily  enabled  us  to  overtake  two  whales,  whereas  at  14.8  km/hr  the 
ship  could  only  keep  up  with  them.  The  two  that  we  accompanied  tired  more  quickly  than 


II 

other  mysticete  whales  we  have  studied. 

The  whales'  respiratory  rate  depended  upon  their  activity.  Whales  swimming  slowly 
at  the  surface  ventilated  about  once  a  minute,  or  once  every  2  to  3  min  when  performing 
"headstands"  in  the  interim  (Fig.  5A).  A  whale  encountered  in  deeper  water  as  it  swam 
fairly  rapidly  and  directly  towards  shore  breathed  irregularly,  diving  for  periods  that  var- 
ied between  0.5  and  4.3  min  (Fig.  5B).  A  whale  we  had  been  following  stayed  down  for 
6.2  min  (Fig.  5C),  and  at  another  time  its  longest  dive  was  8  min. 

Transiting  right  whales  (those  that  moved  along  on  a  direct  course  without  spending 
much  time  at  the  surface)  usually  displayed  their  flukes  just  before  an  extensive  dive  (Fig. 
6).  However,  the  next  to  last  surfacing  was  sometimes  accompanied  by  "false  fluking,"  the 
flukes  being  drawn  up  close  to  but  not  above  the  surface. 

Our  brief  stay  in  Golfo  San  Jose  did  not  permit  long-term  observations  of  southern 
right  whales,  but  the  following  information  was  obtained  from  local  observers,  particu- 
larly Santiago  Ortega,  Perez  Macchi,  and  Carlos  Oscar  Garci'a: 

1.  Southern  right  whales  occur  in  Golfo  San  Jose  and  nearby  Golfo  Nuevo  each 
year.  The  whales  come  from  the  south  and  begin  to  show  up  at  Golfo  San  Jose  in  late 
June,  before  they  appear  in  Golfo  Nuevo.  They  are  most  numerous  in  late  August  and 
September.  Fewer  are  present  in  October,  and  all  disappear  during  November. 

2.  The  total  number  of  right  whales  in  the  Valdes  region  is  unknown,  mainly  because 
there  is  replacement,  with  some  arriving  and  some  departing  throughout  the  season.  Up 
to  30  may  be  seen  at  one  time  in  Golfo  San  Jose  during  the  peak  of  the  season,  and  others 
can  be  found  out  to  sea,  just  east  of  the  peninsula.  The  reporters  knew  of  no  other  place 
along  the  Argentine  coast  where  this  species  enters  bays  each  year. 

3.  All  observers  thought  that  right  whales  court  and  copulate  in  the  two  gulfs,  but 
none  thought  that  the  young  were  born  there.  Small  whales,  seen  in  these  gulfs,  were 
judged  too  large  to  have  been  born  there  during  a  current  "whale  season." 

4.  Right  whales  reportedly  have  become  more  numerous  in  this  area  in  recent  years, 
as  is  true  of  elephant  seals  and  sea  lions  of  the  region,  possibly  because  the  Valdes  Penin- 
sula has  become  a  wildlife  refuge  for  land  and  sea  animals.  It  is  illegal  to  kill  marine 
mammals  in  either  gulf  and  the  area  is  heavily  patrolled. 

DISCUSSION 

We  have  recorded  underwater  sounds  from  six  of  the  ten  species  of  mysticete  whales, 
and  all  the  sounds  have  been  low-frequency  utterances  below  3000  Hz.  Although  there 
are  a  few  reports  of  high-frequency  phonations  in  the  presence  of  mysticetes  (Perkins, 
1966;  Poulter,  1968;  Beamish  and  Mitchell,  1971),  baleen  and  toothed  whales  are 
markedly  diff"erent  in  that  low  frequency  is  more  typical  of  the  former  and  high  frequency 
of  the  latter.  A  good  single  hydrophone  can  yield  an  acceptable  recording  of  these  strong 
low-frequency  signals,  when  used  in  calm  seas  and  not  towed  through  the  water  . 

(The  single,  omnidirectional  hydrophone  may  eventually  be  replaced  by  an  eff"ective 
line  array  of  several  transducers  that  could  be  towed  from  a  drifting  or  sailing  ship.  In 
theory,  the  advantage  of  a  line  array  comes  from  increased  signal-to-noise  ratio  accom- 
plished through  directivity  toward  the  sound  source.  The  means  for  attaining  such  direc- 
tivity generally  involves  proper  spacing  of  the  individual  hydrophones  to  phase  out  the 
input  from  directions  other  than  those  that  are  normal  to  the  array's  conformation.  The 
extent  of  spacing  is  related  to  the  wavelength  involved  and  thus  would  be  considerable  m 
the  case  of  low-frequency  mysticete  sounds  where  wavelengths  are  as  long  as  120  m  or 
more.  A  5-hydrophone  array  for  this  frequency  would  approximate  300  m  in  length.  Un- 
fortunately, present  systems  such  as  this  are  burdensome  to  use,  and  they  involve  complex 
signal  processing.  Moreover,  the  line  array  is  not  within  the  budget  of  most  scientific  in- 
vestigations, nor  is  one  yet  available  that  is  relatively  noise-free  with  a  high  enough 
response  across  the  entire  frequency  range  of  mysticete  sounds.) 

The  belch-like  sounds  and  complex  moans  of  southern  right  whales  reported  here  re- 
semble some  of  the  sounds  of  northern  right  whales  presented  by  Schevill  and  Watkms 
(1962).  However,  we  also  found  dissimilarities.  Diff"erences  in  behavior,  m  addition  to  pos- 


12 

sible  regional  differences  resulting  from  evolutionary  divergence,  could  account  for  such  a 
disparity.  Furthermore,  our  recordings  had  no  evidence  of  repetitive  stanzas  that  Cum- 
mings  and  Philippi  (1970)  suggested  were  from  northern  right  whales.  Their  recordings 
were  made  in  the  open,  deep  sea,  far  from  land  in  an  area  of  the  North  Atlantic  where 
breeding  of  northern  right  whales  is  unknown.  The  present  recordings  were  made  near 
shore  in  a  protected,  shallow  embayment,  where  whales  were  courting. 

Our  earlier  sound  playback  experiments  were  so  successful  in  producing  avoidances 
by  gray  whales  and  white  whales  that  we  expected  the  right  whales  to  behave  similarly, 
especially  as  they  occur  in  an  area  where  killer  whales  are  known  to  attack  them.  Perhaps 
other  playback  attempts  would  produce  a  more  decisive  reaction.  The  source  level  of  the 
sounds  played  was  less  than  that  used  in  experiments  with  gray  whales  and  white  whales. 
Also,  the  right  whales  were  an  appreciable  distance  from  the  ship,  in  very  shallow  water 
where  we  would  not  expect  good  propagation.  Possibly  the  playback  was  not  very  audible 
to  the  whales  in  this  location. 

Right  whales  exhibited  the  "headstanding"  posture  mostly  in  the  southeast  comer  of 
Golfo  San  Jose,  in  an  area  reported  by  fishermen  to  have  dense  populations  of  mussels. 
Local  fishermen  believe  that  this  behavior  is  associated  with  feeding  on  mussels,  and  we 
support  this  idea.  We  observed  the  whales  in  water  that  was  shallow  enough  to  allow  their 
heads  to  touch  bottom  and  their  tails  to  be  exposed.  In  two  instances  of  "headstanding," 
at  a  location  where  the  water  was  only  6  to  8  m  deep,  the  whales  either  had  to  be  in  con- 
tact with  the  bottom  or  had  to  arch  their  whole  bodies.  It  is  doubtful  that  they  could  have 
held  their  tails  out  of  the  water  for  such  a  long  time,  with  the  remainder  of  the  body  bent 
away  from  the  bottom.  Moreover,  we  did  not  observe  "headstanding"  in  places  known  to 
be  deeper  than  the  estimated  length  of  the  whales.  Right  whales  are  positively  buoyant, 
but  holding  their  tails  above  the  surface  may  make  them  just  heavy  enough  in  water  to 
keep  their  heads  against  the  bottom. 

The  idea  of  mysticetes  feeding  on  mussels  is  not  without  precedent.  Scammon  (1874) 
reported  that  gray  whales  fed  on  mussels,  based  upon  his  having  observed  the  mussels  in 
the  whales'  "maws."  In  fact,  he  included  the  term  "Mussel-digger"  in  his  list  of  vernacular 
names  for  the  gray  whale. 

The  Valdes  Peninsula  area  is  well  suited  for  studies  of  marine  mammals.  In  addition 
to  southern  right  whales  and  killer  whales  in  Golfo  San  Jose,  we  observed  southern  sea 
lions  (Otaria  flavescens),  elephant  seals  (Mirounga  leonina),  bottlenose  porpoises  {Tur- 
siops  truncatus),  "white-sided"  porpoises  {Lagenorhynchus  sp.),  and  common  porpoises 
{Delphinus  sp.).  To  the  south,  large  numbers  of  sea  lions  haul  out  at  points  Leon,  Loma, 
Piramida,  and  Delgada  (Fig.  3).  There  were  about  800  sea  lions  at  Punta  Piramida  at  the 
time  of  our  visit,  and  we  were  told  this  number  would  increase  to  3000  in  December.  Nat- 
ural parks  with  facilities  for  visitors  have  been  established  at  Punta  Loma  and  Punta  Pira- 
mida. Some  elephant  seals  occurred  with  the  sea  Uons  at  Punta  Leon.  From  our  inter- 
views, we  learned  that  elephant  seals  are  most  numerous  from  September  to  December, 
especially  at  Punta  Norte.  It  was  in  this  area,  in  1969,  that  Dr.  Raymond  M.  Gilmore  saw 
150-200  elephant  seals  as  early  as  13  July  (pers.  comm.).  The  seals  were  in  three  scattered 
groups,  some  of  them  occurring  with  sea  lions. 

ACKNOWLEDGMENTS 

This  work  was  supported  by  the  National  Science  Foundation,  U.S.  Antarctic  Research  Program,  Grant 
AG-261;  the  Naval  Undersea  Research  and  Development  Center,  Independent  Research  Projects;  and  by  the 
Office  of  Naval  Research,  Oceanic  Biology  Program.  Dr.  George  A.  Llano  (NSF)  was  very  helpful  in  making  ar- 
rangements for  the  cruise.  We  are  grateful  to  Captain  Franklin  P.  Liberty  and  the  entire  crew  of  HERO  for  their 
splendid  support  and  seamanship.  We  thank  Dr.  Joseph  R.  Jehl,  Jr.,  for  his  help  in  spotting  marine  mammals 
and  identifying  the  birds  for  us;  CDR  Alfredo  A.  Yung,  geophysicist  with  the  Argentine  Navy,  and  Angel  Fer- 
rante,  marine  technician  at  the  Argentine  Institute  of  Oceanography,  for  their  valuable  advice  and  assistance  on 
board;  Carlos  Bassi,  Perez  Macchi,  Santiago  Ortega,  Carlos  O.  Garci'a,  Jorge  E.  Ramirez,  Jorge  R.  Terenzi,  and 
others,  for  their  very  useful  information;  Charlotte  L.  Meinert  and  Alan  R.  Hamel,  for  assistance  in  preparing 
the  manuscript;  and  William  E.  Schevill,  William  A.  Watkins,  and  R.  S.  Gales,  for  their  helpful  comments. 


13 

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Beamish,  P.,  and  E.  Mitchell 

1971.   Ultrasonic  sounds  recorded  in  the  presence  of  a  blue  whale,  Balaenoptera  musculus  Deep-sea  Res 

18:  803-809. 

Calderon,  M.  A.,  and  G.  M.  Wenz 

1967.  A  portable  general-purpose  underwater  sound  measuring  system.  Naval  Undersea  Warfare  Ctr., 
Tech.  Paper  25.  46  p. 

Cummings,  W.  C,  J.  F.  Fish,  P.  O.  Thompson,  and  J.  R.  Jehl,  Jr. 

Bioacoustics  of  marine  mammals  off  Argentina,  R/V  HERO,  Cruise  71-3.  Antarctic  Jour  U  S  VI(6)- 
266-268. 

Cummings,  W.  C,  and  L.  A.  Philippi 

1970.  Whale  phonations  in  repetitive  stanzas.  Naval  Undersea  Res.  Develop.  Center,  Tech.  Publ.  196.  8  p. 
Cummings,  W.  C,  and  P.  O.  Thompson 

1971.  Gray  whales,  Eschrichtius  robustus,  avoid  the  underwater  sounds  of  killer  whales,  Orcinus  orca.  Fish. 
Bull.  69(3):  525-530. 

Doi,  T.,  S.  Ohsumi,  and  Y.  Shimadzu 

1971.  Status  of  stocks  of  baleen  whales  in  the  Antarctic,  1970/71.  Internatl.  Comm.  Whahng,  21st  Rept.- 
90-99. 

Fish,  J.  F.,  and  J.  S.  Vania 

1971.  Killer  whale,  Orcinus  orca,  sounds  repel  white  whales,  Delphinapterus  leucas.  Fish.  Bull.  69  (3):  531- 
535. 

Gilmore,  R.  M. 

1969.  Populations,  distribution,  and  behavior  of  whales  in  the  western  South  Atlantic:  Cruise  69-3  of  R/V 
HERO.  Antarctic  Jour.  U.S.,  IV  (6):  307-308. 

Hershkovitz,  P. 

1966.  Catalog  of  living  whales.  U.S.  Natl.  Mus.  Bull.  246. 

Matthews,  L.  H. 

1938.   Notes  on  the  southern  right  whale,  Eubalaena  australis.  Discovery  Rept.  XVll:  169-182. 

Ohsumi,  S.,  Y.  Shimadzu,  and  T.  Doi 

1971.  The  seventh  memorandum  on  the  results  of  Japanese  stock  assessment  of  whales  in  the  North  Paci- 
fic. Internatl.  Comm.  Whaling,  21st  Rept.:  76-89. 

Payne,  R.  S.,  and  S.  McVay 

1971.  Songs  of  humpback  whales.  Science  173  (3997):  585-597. 

Perkins,  P.  J. 

1966.  Communication  sounds  of  finback  whales.  Norsk  Hvalfangst-Tidende  (Norwegian  Whaling  Gazette) 
55:  199-200. 

Poulter,  T.  C. 

1968.  Marine  mammals,  p.  405-465.  /«  T.  A.  Sebeok  [ed.].  Animal  communication,  techniques  of  study  and 
results  of  research.  Indiana  Univ.  Press,  Bloomington,  Ind. 

Rice,  D.  W.,  and  V.  B.  SchefTer 

1968.  A  list  of  the  marine  mammals  of  the  world.  U.S.  Fish  WildHfe  Ser.,  Spec.  Sci.  Rept.-Fish.  No.  579. 

Ridewood,  W.  G. 

1901.  On  the  structure  of  the  horny  excrescence,  known  as  the  bonnett,  of  the  southern  right  whale  {Ba- 
laena  australis).  Proc.  Zool.  Soc.  Lond.  1901:  AA-Al. 

Scammon,  C.  M. 

1874.  The  marine  mammals  of  the  north-western  coast  of  North  America,  and  the  American  whale  fishery. 
Facsimile  edition,  Manessier  Publishing  Co.,  Riverside,  Calif.  319  p. 

Schevill,  W.  E. 

1964.  Underwater  sounds  of  cetaceans,  p.  307-316.  In  W.  N.  Tavloga  [ed.].  Marine  bio-acoustics,  proceed- 
ings of  a  symposium  held  at  the  Lerner  Marine  Laboratory,  Bimini,  Bahamas,  April  11  to  13,  1963. 
Pergamon  Press,  N.Y. 

Schevill,  W.  E.,  and  W.  A.  Watkins 

1962.  Whale  and  porpoise  voices.  Woods  Hole  Oceanogr.  Inst,  (with  a  phonographic  record)  24  p. 

Watkins,  W.  A. 

1966.  Listening  to  cetaceans,  p.  471-476.  In  K.  S.  Norris  [ed.].  Whales,  Dolphins,  and  Porpoises.  Univ.  Cali- 
fornia Press,  Berkeley  and  Los  Angeles. 

Wenz,  G.  M. 

1962.  Acoustic  ambient  noise  in  the  ocean:  spectra  and  sources.  J.  Acoust.  Soc.  Amer.  34  (12):  1936-1956. 

Naval  Undersea  Research  and  Development  Center,  Applied  Bioacoustics  Branch,  San 
Diego,  California  92132 


MUS.  COMI>.  ZOOU 

LIBRARY 

DEC    71976 

EASTERN  PACIFIC  SNAKE-EELS  u^tvERSr^f 

OF  THE  GENUS  CALLECHELYS  (APODES:  OPHICHTHIDAE) 


JOHN  E.  McCOSKER  AND  RICHARD  H.  ROSENBLATT 


TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  2  25  APRIL  1972 


EASTERN  PACIFIC  SNAKE-EELS 
OF  THE  GENUS  CALLECHELYS 
(APODES:  OPHICHTHIDAE) 

JOHN  E.  McCOSKER  AND  RICHARD  H.  ROSENBLATT 

ABSTRACT.-Three  species  of  Callechelys  are  recognized  from  the  eastern  tropical  Pacific.  Two,  C.  eris- 
tigmus  and  C.  galapagensis.  are  described  as  new.  Callechelys  cliffi  Bohlke  and  Briggs  is  redescribed  from 
adults.  The  species  differ  in  coloration,  body  proportions  and  vertebral  number.  Callechelys  cliffi  and 
C.  eristigmus  n.  sp.  range  from  Panama  to  the  Gulf  of  California,  and  C.  galapagensis  is  known  only  from 
the  Galapagos  Islands.  Vertebral  number  and  proportional  tail  length  of  the  15  species  of  Callechelys  are 
given.  Lineages  within  the  genus  are  indicated  by  the  presence  or  absence  of  a  scapula  and  the  condition  of 
the  urohyal. 

Callechelys  is  one  of  the  larger  genera  of  the  Ophichthidae  with  fifteen  species, 
mainly  Hmited  to  the  tropics.  The  species  are  distinguished  on  the  basis  of  coloration,  ver- 
tebral number,  and  certain  body  proportions,  mainly  body  depth  and  preanal  distance. 
Like  most  snake-eels,  members  of  the  genus  are  sand  dwelling  and  restricted  to  continen- 
tal shelf  depths.  Some  of  the  species  attain  lengths  of  one  meter.  It  is  not  known  whether 
they  occupy  burrows  or  wander  extensively  through  the  sand.  Despite  the  sand-dwelling 
habit,  many  of  the  species  are  boldly  marked.  It  is  possible  that  they  leave  the  sand  at 
night,  and  the  color  pattern  may  have  significance  at  these  times.  Occasional  specimens 
have  indeed  been  taken  at  the  surface  under  lights  at  night. 

When  Storey  (1939)  published  her  revision  of  Callechelys,  a  single  specimen,  doubt- 
fully referred  to  C.  marmoratus  or  C.  luteus,  was  known  from  the  eastern  tropical  Pacific. 
Subsequently  (Bohlke  and  Briggs,  1954)  another  specimen  was  taken  and  made  the  holo- 
type  of  a  new  species.  The  collections  at  the  Scripps  Institution  of  Oceanography,  the  De- 
partment of  Zoology,  University  of  California,  Los  Angeles,  and  the  University  of  Costa 
Rica  now  contain  55  eastern  Pacific  specimens  of  Callechelys.  The  recent  collections  of 
these  eels  are  attributable  to  the  development  of  synergized  emulsified  rotenone  products. 
The  use  of  these  products  has  resulted  in  rich  collections  of  ophichthid  eels  and  other 
sand-dwelling  fishes  not  obtainable  with  powdered  derris  root.  Even  with  the  use  of  pow- 
erful ichthyocides,  the  collection  of  these  eels  is  not  easy.  Either  because  of  a  resistance  to 
rotenone  or  the  time  involved  in  transport  of  rotenone  down  into  the  sand,  they  emerge 
long  after  most  fishes  are  dead.  Ophichthids  may  begin  to  appear  after  other  fishes  have 
been  picked  up  and  the  station  apparently  completed. 

Our  material  can  be  separated  into  three  species,  only  one  of  which  has  been  de- 
scribed. Callechelys  cliffi  Bohlke  and  Briggs,  heretofore  known  only  from  the  just-trans- 
formed holotype,  can  now  be  described  on  the  basis  of  adult  characters. 

MATERIALS  AND  METHODS 

Material  used  in  this  study  is  housed  in  the  following  institutions:  University  of  Cali- 
fornia at  Los  Angeles,  Department  of  Zoology  (UCLA);  National  Museum  of  Natural 
History  (USNM);  California  Academy  of  Sciences,  material  previously  at  Stanford  Uni- 
versity, (SU);  Universidad  de  Costa  Rica,  Museo  de  Zoologia  (UCR);  and  Scripps  In- 
stitution of  Oceanography  (SIO).  Paratypes  of  Callechelys  eristigmus  will  be  deposited  at 
the  Academy  of  Natural  Sciences  of  Philadelphia  and  the  USNM. 

All  measurements  are  straight-line  measurements,  made  either  with  a  300  mm  ruler 
with  0.5  mm  gradations  (for  standard  length,  trunk  length,  and  tail  length)  and  recorded 
to  the  nearest  0.5  mm,  or  with  dial  calipers  (all  other  measurements)  and  recorded  to  the 
nearest  0. 1  mm.  Head  length  is  measured  from  the  snout  tip  to  the  posterodorsal  margin 
of  the  gill  opening;  trunk  length  is  taken  from  the  end  of  the  head  to  mid-anus;  body 

SAN  DIEGO  see.  NAT.  HIST,  TRANS.  17  (2):  15-24,  25  APRIL  1972 


16 


depth  does  not  include  the  fin.  Counts  and  proportions  in  Tables  2-4  include  the  mean, 
range,  and  95%  confidence  limits  of  the  mean.  Fin  ray  and  vertebral  counts  (which  in- 
clude the  hypural)  were  made  using  radiographs  or  cleared  and  stained  specimens. 

Callechelys  Kaup,  1856 

The  genus  Callechelvs  may  be  distinguished  from  all  other  ophichthid  genera  on  the 
basis  of  the  following  combination  of  characters:  tip  of  tail  a  hard  point;  pectorals  absent; 
anal  fin  present;  dorsal  fin  originating  on  head;  head  and  body  laterally  compressed;  an- 
terior nostrils  tubular;  a  median  groove  on  underside  of  snout;  gill  openings  low-lateral 
and  converging  forward,  the  isthmus  much  narrower  than  the  gill  opening  length;  inter- 
maxillary and  vomerine  teeth  present,  canine  teeth  absent. 

Key  to  the  Eastern  Pacific  Species  of  Callechelys 

la.  Tail  considerably  shorter  than  head  and  trunk,  3.25-3.75  in  total  length.  Greatest 
body  depth  of  adults  3.3-3.5  times  in  head  length.  Color  pattern  of  strongly  con- 
trasted round  dark  spots  about  as  long  as  snout. 
Vertebrae  154-163 Callechelys  ehstigmus  n.  sp. 

lb.  Tail  almost  equal  to  head  and  trunk,  2.2-2.4  in  total  length.  Greatest  body  depth  of 
adults  1.7-2.8  times  in  head  length.  Color  pattern  either  solely  of  numerous  small 
dark  spots,  or  with  larger  dark  oblongs  as  well,  in  which  case  the  total  vertebrae  are 
170-174 2 

2a.  Color  pattern  of  numerous  fine  spots,  not  much  larger  than  eye  diameter,  fins  with  a 
distinct  white  edge.  Vertebrate  149-158 Callechelys  clijfi  Bohlke  and  Briggs 

2b.  Color  pattern  with  oblong  blotches  varying  in  size  between  an  eye  diameter  and  a 
snout  length  along  the  major  axis,  fins  without  a  white  edging. 
Vertebrae  170-174 Callechelys  galapagensis  n.  sp. 


B 


Figure  la.  Callechelys  eristigmus  n.  sp.,  holotype,  SIO  65-263,  503.5  mm  total  length,  b.  Head  region  of  holotype 
of  Callechelys  eristigmus  n.  sp.  Arrow  indicates  true  dorsal  fin  origin. 

Callechelys  eristigmus  n.  sp. 

Figs.  1,  2a,  5;  Tables  1,  2,  5 
Description  of  holotype. —Counts  and  proportions  of  the  holotype  are  given  in  Table 
1.  Proportions  of  the  holotype  and  29  paratypes  are  given  in  Table  2. 

Body  laterally  compressed  throughout  its  length,  tapering  posteriorly  to  a  hard  fin- 


17 


Table  1.    Counts,  and  proportions  in  thousandths,  of  the  holotypes  of  the  eastern  Pacific  species  of 
Callechelys. 


C.  cliffi 

C.  eristigmus 

C.  galapagensis 

Total  length  (mm) 

93.5 

503.5 

818.0 

Total  vertebrae 

155 

159 

172 

Preanal  vertebrae 

105 

92 

(thousandths 

of  total  length) 

Head 

101 

72 

74 

Trunk 

467 

628 

483 

TaU 

433 

300 

444 

(thousandths 

of  head  length) 

Dorsal  fin  origin 

606 

318 

312 

Snout 

170 

142 

133 

Upper  jaw 

340 

285 

262 

Eye 

74 

41 

46 

Interorbital 

132 

108 

Isthmus 

74 

55 

119 

Depth  behind  gill  opening 

351 

345 

464 

Width  behind  gill  opening 

249 

265 

Depth  at  anus 

330 

258 

365 

Width  at  anus 

195 

249 

Gill  opening  length 

145 

202 

less  point.  Depth  behind  gill  openings  40  times  and  at  anus  51  times  in  total  length;  width 
behind  gill  openings  55  and  at  anus  71  in  total  length.  Head  and  trunk  1.4.  head  13.8  in 
total  length.  Snout  acute,  rounded  at  tip.  Lower  jaw  included,  its  tip  slightly  before  eye 
and  midway  between  anterior  and  posterior  nostrils.  Eye  small,  about  as  long  as  tube  of 
anterior  nostril.  Posterior  nostrils  open  into  mouth  although  their  distal  edges  are  open  to 
the  outer  edge  of  lip,  visible  externally  as  a  slit.  Surface  of  head,  trunk  and  tail  markedly 
wrinkled  (except  top  and  sides  of  anterior  portion  of  head  smooth),  with  approximately 
30  longitudinal  grooves  on  each  side  of  body.  Tongue  adnate.  Branchial  basket  expanded, 
supported  by  31  pairs  of  branchiostegals  and  jugostegalia  which  broadly  overlap  along 
ventral  midline.  Urohyal  simple,  a  single  slender  filament  posteriorly.  Tip  of  lower  jaw 
and  lateral  skin  folds  of  upper  jaw  covered  with  numerous  papillae  (Fig.  2a). 

Teeth  small  and  pointed  (Fig.  2a),  uniserial  in  jaws.  An  anterior  intermaxillary  tooth- 
pair  covered  by  skin  folds  for  most  of  their  length,  followed  by  seven  vomerine  teeth  that 
become  biserial  posteriorly. 

Preoperculomandibular,  temporal,  postorbital,  suborbital,  and  supraorbital  series  of 
pores  present  (Fig.  lb).  Lateral  line  beginning  on  head  with  10  pores  before  gill  opening 
(lateral  line  canal  and  pores  difficult  to  discern  due  to  skin  folds  and  waxy  precipitate 
which  forms  on  preserved  specimens).  Total  right  lateral  line  pores  of  the  cleared  and 
stained  specimen  140,  92  before  anus.  Last  pore  ca.  0.15  head  lengths  before  tail  tip.  Dor- 
sal fin  origin  on  head,  above  and  slightly  behind  rictus;  median  fins  end  about  a  snout 
length  before  tail  tip.  Fin  rays  in  dorsal  498,  anal  143  (counted  from  the  cleared  and 
stained  paratype). 

Gill  arches  and  hyoid  apparatus  of  two  paratypes  removed  and  stained.  First  basi- 
branchial  ossified,  second  cartilaginous,  third  and  fourth  absent.  Hypobranchials  1-2  os- 
sified, third  cartilaginous.  Ceratobranchials  1-4  ossified,  fifth  absent.  Infrapharyngobran- 
chials  2-3  ossified.  Lower  pharyngeal  teeth  in  elongate  patches  on  fourth  ceratobranchial 
and  extend  onto  hypobranchial;  upper  pharyngeal  tooth  plate  smaller,  attached  to  distal 
ends  of  epibranchials  2-4  and  second  infrapharyngobranchial  (I3  of  Nelson,  1966). 

Color  in  alcohol  mostly  cream,  overlain  with  numerous  dark  spots  that  extend  onto 
the  dorsal  fin  membrane.  Chin,  throat,  and  venter  often  spotted,  but  always  less  so  than 
dorsum.  Spots  on  nape  and  snout  smaller  (nearly  as  large  as  eye). 

Etymology.— From  the  Greek  epi  (eri),  intensive  participle,  and  a-TLyna  (stigma),  spot, 
in  reference  to  the  distinctive  coloration;  regarded  as  an  adjective. 


18 


Remarks.— Gulf  of  California  specimens  are  inseparable  on  the  basis  of  coloration 
and  morphology  from  those  from  southern  localities  (Cocos  Island,  Costa  Rica,  and  Pan- 
ama). The  mean  vertebral  number  of  specimens  from  these  localities  (158.3  for  29  Mexi- 
can specimens,  156.1  for  8  specimens  from  the  south)  are  significantly  different  (P  =  .05  by 
t  test);  however,  the  degree  of  joint  non-overlap  is  not  sufficient  to  warrant  taxonomic  rec- 
ognition. 

Table  2.    Callechelys  eristigmus  n.  sp.,  counts,  and  proportions  in  thousandths,  of  holotype  and  29  para- 
types;  mean,  95%  confidence  limits  of  the  mean,  and  range. 


X 

95%  C.  L. 

range 

Total  length  (mm 

) 

284-1 126mm 

Vertebrae  (37  specimens) 

157.9 

157.2-158.5 

154-163 

(thousandths  of  total  length) 

Head 

76 

74-77 

67-83 

Trunk 

628 

624-632 

610-662 

Tail 

295 

291-299 
(thousandths  of  head  length) 

268-309 

Dorsal  fin  origin 

327 

316-337 

250-379 

Snout 

152 

149-155 

135-167 

Upper  jaw 

266 

260-272 

228-289 

Eye 

46 

44-48 

34-56 

Interorbital 

113 

109-117 

98-140 

Left  gill  opening  1 

ength 

142 

136-147 

111-168 

Isthmus 

58 

54-61 

46-77 

Depth  behind  gill 

opening 

295 

286-304 

254-345 

Depth  at  anus 

250 

237-262 

172-338 

Width  behind  giU 

opening 

194 

185-203 

144-272 

Width  at  anus 

182 

174-190 

140-234 

Material  examined.— WoXoiy^Q:  SIO  65-263,  a  503.5  mm  adult  from  Isla  San  Jose, 
Gulf  of  Cahfornia,  Baja  California  Sur  (24°52'15"N,  1 10°37'00"W).  Taken  with  rotenone 
and  SCUBA  in  depths  of  20-25  m  on  a  sand  and  boulder  bottom  by  R.  H.  Rosen- 
blatt and  party  on  7  July  1965.  Paratypes:  all  collected  using  rotenone  ichthyocides  in  rel- 
atively shallow  water  (5-25  m),  generally  over  a  sand  and  rock  bottom.  Panama-Islas 
Secas,  Isla  Cavada,  SIO  70-136, 2(359-465);  Islas  Secas,  Isla  Seca,  70-140, 2(342-450).  Costa 
Rica-Isla  del  Coco,  UCLA  58-378,  1(361);  Isla  del  Caiio,  UCR  423-126,  5(216-491).  Gulf 
of  California,  Baja  California  Sur-Isla  Carmen,  UCLA  65-77,  3(325-420),  SIO  65-299, 
2(328-357);  Isla  Santa  Catalina,  SIO  65-337,  2(411-469);  Isla  Santa  Cruz,  SIO  65-342, 
2(408-420),  SIO  65-354,  2(431-498,  the  smaller  specimen  cleared  and  stained);  Punta 
Nopolo,  SIO  65-270,  1(564);  Isla  San  Jose,  SIO  65-263,  1(323,  collected  with  the  holotype), 
SIO  65-260,  1(372);  Isla  Espiritu  Santo,  SIO  61-277,  1(1 126);  Bahia  de  los  Lobos,  SIO  61- 
279,  1(418);  Isla  Ceralbo,  SIO  61-256,  1(494),  SIO  61-259,  2(399-493);  Bahia  de  Palmas, 
UCLA  59-249,  1(285),  UCLA  59-251,  2(284-538);  Punta  Pescadero,  SIO  61-252,  2(317- 
557);  Punta  Los  Frailes,  SIO  61-239,  1(425);  El  Tule  Ranch,  east  of  Cabo  San  Lucas,  SIO 
65-185,3(363-552). 

Callechelys  galapagensis,  n.  sp. 
Figs.  2b,  3,  5;  Tables  1,  3,  5 
Callechelys  marmoratus,  nee  Bleeker,  Fowler,  1932:  3.  Fowler,  1938:  251. 
Callechelys  luteus,  nee  Synder,  Storey,  1939:  69. 

Description  of  holotype.— Connie  and  proportions  of  the  holotype  are  given  in  Table 
1.  Proportions  of  the  holotype  and  3  paratypes  are  given  in  Table  3. 

Body  laterally  compressed  throughout  its  length,  tapering  posteriorly  to  a  hard  fin- 
less  point.  Depth  behind  gill  openings  29  times  and  at  anus  37  in  total  length;  width  be- 
hind gill  openings  51  and  at  anus  54.5  in  total  length.  Head  and  trunk  1.8,  head  13.5  in 
total  length.  Snout  acute,  rounded  at  tip.  Lower  jaw  included,  its  tip  closer  to  base  of  ante- 
rior nostrils  than  to  a  vertical  from  anterior  margin  of  eye.  Eye  small,  about  equal  in 
length  to  tube  of  anterior  nostril.  Posterior  nostrils  open  into  mouth,  visible  externally  as 
a  slit.  Surface  of  head  and  trunk  markedly  wrinkled  (except  top  and  sides  of  anterior  por- 


19 


Figure  2a.  Dentition  of  holotype  of  Callechelvs  eristigmus.  b.  Callechelys  galapagensis.  n.  sp..  a  paratype,  UCLA 
67-33.  c.  Callechelys  cliffi.  SIO  62-42. 

tion  of  head  smooth)  as  in  C.  eristigmus,  but  becoming  smoother  posteriorly.  Tongue  ad- 
nate.  Branchial  basket  expanded,  supported  by  26  pairs  of  branchiostegals  and  jugoste- 
galia  which  broadly  overlap  along  ventral  midline.  Urohyal  split  into  two  slender 
filaments  for  about  90%  of  its  length.  Tip  of  lower  jaw  and  lateral  skin  folds  of  upper  jaw 
papillose,  as  in  C.  eristigmus. 

Teeth  small  and  pointed  (Fig.  2b).  Intermaxillary  teeth  comprising  two  or  three  pairs 
partially  covered  by  skin  folds,  largest  anteriorly,  followed  by  from  four  to  six  pairs  of  bi- 
serial  vomerine  teeth.  Lower  jaw  teeth  uniserial  and  small,  about  10  to  15  on  each  side. 

Preoperculomandibular,  temporal,  postorbital,  suborbital,  and  supraorbital  series  of 
head  pores  present,  not  unlike  those  of  C.  eristigmus  (Fig.  lb)  in  number  and  position. 
Lateral  line  (of  the  left  side  of  holotype)  beginning  on  head,  with  10  pores  before  gill 
opening,  87  to  anus,  and  157  total  pores  ending  0.2  head  lengths  from  tail  tip.  Dorsal  fin 
origin  on  head,  above  and  slightly  behind  rictus;  median  fins  ending  less  than  a  snout 


Table  3.    Callechelys  galapagensis  n.  sp.,  counts,  and  proportions  in  thousandths,  of  holotype  and  3  para- 
types;  mean,  and  95%  confidence  limits  of  the  mean,  and  range. 


X 

95%  C.  L. 

range 

Total  length  (mm) 

248-818  mm 

Vertebrae 

172 

169.4-174.6 

170-174 

(thousandths  of  total  length) 

Head 

76 

67-85 

69-82 

Trunk 

478 

456-499 

463-494 

TaU 

444 

(thousandths 

432-456 
;  of  head  length) 

437-455 

Dorsal  fin  origin 

325 

293-357 

312-355 

Snout 

132 

127-138 

128-137 

Upper  jaw 

248 

194-302 

197-269 

Eye 

55 

42-68 

46-66 

Interorbital 

103 

80-126 

81-112 

Left  gill  opening  length 

178 

115-240 

127-214 

Isthmus 

109 

71-148 

76-133 

Depth  behind  gill  opening 

418 

293-544 

349-506 

Depth  at  anus 

352 

226-479 

289-460 

Width  behind  gill  opening 

252 

170-334 

203-319 

Width  at  anus 

226 

158-293 

178-272 

20 


length  before  tail  tip.  Fin  rays  in  dorsal  520,  anal  230  (counted  from  a  radiograph  of  holo- 
type). 

Gill  arches  and  hyoid  apparatus  of  the  largest  paratype  (UCLA  64-40)  removed  and 
stained.  Configuration  and  condition  of  the  gill  arch  members  like  that  of  C.  eristigmus 
except  that  the  upper  and  lower  pharyngeal  tooth  plates  are  nearly  equal  in  length,  and 
oblong  rather  than  elongate  (the  lower  pharyngeal  plate  of  C.  eristigmus  is  larger  and 
more  slender  than  the  upper). 


B 


Figure  3a.  Callechelvs  galapagensis  n.  sp.,  paratype,  UCLA  64-40,  767  mm  total  length,  b.  Callechehs  galapa- 
gensis  n.  sp.,  a  darkly  colored  paratype,  UCLA  67-33,  248  mm  total  length. 

Color  in  alcohol  mostly  cream,  overlain  with  numerous  dark  oblong  markings  that 
vary  in  length  from  the  size  of  the  eye  to  the  length  of  the  upper  jaw.  These  spots  extend 
onto  median  fins  and  become  densely  aggregated  on  chin  and  top  and  sides  of  head.  Ven- 
tral and  dorsal  surfaces  more  spotted  than  flanks,  which  have  a  row  of  small  spots  une- 
venly distributed  along  midline.  The  smallest  paratype  (Fig.  3b)  differs  from  the  other 
types  in  having  a  chocolate  brown  background  coloration,  although  the  spotting  is  sim- 
ilar. 

f/vmo/ogy.— Named  galapagensis,  for  the  locality  at  which  all  known  specimens  were 
collected. 

Material  examined.— \\o\o{y\>Q:  SIO  72-1,  formerly  UCLA  64-39,  an  818  mm  adult 
from  the  Galapagos  Islands,  Isla  Santa  Cruz,  north  shore,  off  small  cove.  Taken  with 
Chemfish  and  SCUBA  over  a  sand,  rock,  and  sparse  coral  bottom  in  ten  meters  by  B.  W. 
Walker  and  E.  S.  Hobson  on  24  February  1964.  Paratypes:  all  from  the  Galapagos  Is- 
lands. USNM  89728,  1(312),  Isla  Santa  Maria,  Black  Beach  Anchorage.  UCLA  64-40, 
1(767),  Isla  Santa  Cruz,  North  Coast.  UCLA  67-33,  1(248),  Isla  San  Salvador,  James  Bay. 

Callechelvs  cliffi  Bohlke  and  Briggs 
Figs.'2c,  4,"5;Tables  1,4,5 
Callechelys  cliffi  Bohlke  and  Briggs,  1954:  275.  Fraile  Bay  (Los  Frailes),  Gulf  of  Califor- 
nia. 

Description.— Counts  and  proportions  of  the  holotype  are  given  in  Table  1.  Propor- 
tions of  several  juvenile  and  adult  specimens  are  given  in  Table  4.  The  following  descrip- 
tion is  based  on  the  adult  specimens. 


21 

Body  laterally  compressed  throughout  its  length,  tapering  posteriorly  to  a  point. 
Depth  behind  gill  openings  23  times  and  at  anus  27  in  total  length;  width  behind  gill 
openings  43.5  and  at  anus  45  in  total  length.  Head  and  trunk  1.7,  head  10.7  in  total 
length.  Snout  acute,  rounded  at  tip.  Lower  jaw  included,  tip  reaches  level  of  anterior  nos- 
trils. Eye  small,  about  as  long  as  anterior  nostril  base.  Posterior  nostrils  open  into  mouth, 
visible  externally  as  a  slit.  Surface  of  head,  trunk,  and  tail  wrinkled  (except  top  and  sides 
of  anterior  portion  of  head  smooth),  becoming  smoother  on  flanks  posteriorly,  as  in  C.  ga- 
lapagemis.  Tongue  adnate.  Branchial  basket  expanded,  supported  by  26  pairs  of  bran- 
chiostegals  and  jugostegalia  which  broadly  overlap  along  ventral  midline.  Urohyal  split 
into  two  slender  filaments  for  ca.  80%  of  its  length.  Tip  of  lower  jaw  and  anterolateral  skin 
folds  of  upper  jaw  hghtly  papillose  (Fig.  2c). 


t'-.<i»itr'6to.>ftiwi».JtAt.M^;lri»itLyyji>s».^ 


Figure  4.  Adult  of  Callechelys  cliffi,  SIO  62-42,  455  mm  total  length. 

Teeth  small  and  pointed  (Fig.  2c)  uniserial  in  jaws.  Two  anterior  pairs  of  inter- 
maxillary teeth  hidden  by  skin  folds  of  underside  of  snout,  followed  by  a  patch  of  six  to 
seven  intermaxillary  teeth  joined  posteriorly  to  uniserial  vomerine  row. 

Preoperculomandibular,  temporal,  postorbital,  and  supraorbital  pore  series  present. 
Lateral  line  beginning  on  head  with  ten  pores  before  gill  opening,  86  to  anus,  and  151  to- 
tal pores,  ending  0.3  head  lengths  from  tail  tip  (SIO  62-42).  Dorsal  fin  origin  on  head, 
above  and  slightly  behind  rictus;  median  fins  end  about  a  snout  length  before  tail  tip.  Fin 
rays  in  dorsal  457,  anal  205  (from  radiograph  of  UCLA  63-45). 

Gill  arches  and  hyoid  apparatus  of  two  specimens  (SIO  61-247,  65-281)  removed  and 
stained.  The  configuration  and  condition  of  the  gill  arch  members  is  similar  to  that  of  C 
galapagensis  (see  description)  in  that  the  upper  and  lower  pharyngeal  tooth  plates  are 
nearly  equal  in  length  and  oblong,  rather  than  elongate  and  unequal  as  in  C  ehstigmus. 

Color  in  alcohol  tan,  overlain  with  numerous  fine  brown  spots  on  body  and  fins.  Me- 
dian fins  margined  in  white.  Tips  of  snout,  lower  jaw,  and  tail  cream-colored. 

Table  4.    Callechelys  cliffi  Bohlke  and  Briggs,  counts,  and  proportions  in  thousandths,  mean,  95%  confidence 
limits  of  the  mean,  and  range. 


X 

95%  C.  L. 

range 

n 

Total  length  (mm) 

97.5-455  mm 

8 

154-455  mm 

5 

Vertebrae                                                    154.9 

153.4-156.5 

149-158 

14 

(thousandths  of  total  length) 

Head                                                             93 

84-101 

77-108 

8 

Trunk                                                            474 

459-488 

441-496 

8 

Tail                                                             434 

426-442 

418-450 

8 

(thousandths  of  head  length) 

Dorsal  fin  origin                                           4 1 3 

351-474 

363-480 

5 

Snout                                                             142 

123-161 

121-170 

5 

Upper  jaw                                                   311 

294-329 

294-340 

5 

Eye                                                               61 

54-68 

53-74 

5 

Isthmus                                                           97 

79-115 

74-115 

5 

Depth  behind  gjU  opening                           494 

416-573 

351-568 

5 

Depth  at  anus                                               429 

370-488 

330-502 

5 

22 

Remarks.— Out  material  includes  a  single  collection  (SIO  67-40)  which  contains  a 
series  of  individuals  from  newly  settled  juveniles  to  adults.  This  series  displays  the  juve- 
nile to  adult  color  transformation  and  was  compared  with  the  holotype  of  C  cliffi. 
Smaller  specimens  in  the  series  were  identical  with  the  type  in  coloration,  pore  pattern, 
and  morphometry. 

Material  examined— Mexico,  Baja  California  Sur,  Golfo  de  California— Bahia  Los 
Frailes,  SU  47521,  1(93.5  mm),  the  holotype.  Punta  Pulmo,  SIO  61-247,  1(218).  Punta  San 
Telmo,  SIO  65-281,  1(298).  Buena  Vista,  UCLA  63-45,  1(382).  Mexico,  Nayarit,  Bahia  de 
Banderas,  SIO  62-42,  1(455).  Panama,  Archipielago  de  las  Perlas,  Isla  Saboga,  SIO  67-40, 
9(80-154). 


DISCUSSION 

We  recognize  15  tropical  and  subtropical  species  in  the  genus  Callechelys.  C.  guiche- 
noti  Kaup,  the  generic  type,  is  considered  by  us  to  be  a  junior  synonym  of  C  marmoratus 
(Bleeker,  1853).  Kaup's  (1856)  description  and  Pellegrin's  (1912)  redescription  of  the  type 
of  C  guichenoti,  a  475  mm  specimen  from  Tahiti,  do  not  separate  it  from  adults  of  C. 
marmoratus.  Furthermore,  recent  extensive  collecting  efforts  in  Tahiti  and  the  Southern 
Caroline  Archipelago  (by  the  Vanderbilt  Foundation,  J.  E.  Randall,  and  others)  using  im- 
proved ichthyocides  have  obtained  numerous  specimens  of  C.  marmoratus  and  C.  melano- 
taenius  Bleeker.  It  is  highly  unlikely  that  C.  guichenoti,  if  indeed  distinct,  would  not  have 
been  taken  in  the  various  habitats  sampled.  Smith  (1957:  838;  1962)  also  suspected  C 
guichenoti  to  be  a  synonym  of  C.  marmoratus,  but  was  incorrect  in  considering  C.  hiteus 
Snyder  conspecific  with  C.  marmoratus. 

Table  5.    Vertebral  number  and  tail  length  of  the  species  of  Callechelys. 


Tail/SL 

Vertebrae 

Location 

Source 

C.  bilinearis  Kanazawa 

.3642 

162 

West  Atlantic 

Kanazawa,  1952;  this  study 

C.  bitaeniatus  (Peters) 

.385 

E.  Africa,  Mombasa 

Storey,  1939 

C.  cliffi  Bohlke  &  Briggs 

.434 

155 

Eastern  Pacific 

this  study 

C.  eristigmus  sp.  nov. 

.295 

158 

Eastern  Pacific 

this  study 

C.  galapagensis  sp.  nov. 

.444 

172 

Galapagos  Is. 

this  study 

C.  holochromus  (Ginsburg) 

.333 

Gulf  of  Mexico 

Ginsburg,  1951 

C.  leucopterus  (Cadenat) 

.431-.475  164 

Eastern  Atlantic 

Blache  and  Cadenat,  1971 

C.  luteus  Snyder 

.415 
•  345; 
.282; 
.385^ 

213 

Hawaii 

Gosline,  1951 

C.  marmoratus  (Bleeker) 

180 

West  Pacific 

Storey,  1939;  this  study 

C.  melanotaenius  Bleeker 

203 

West  Pacific 

Storey,  1939;  this  study 

C.  muraena  Jordan  &  Evermann 

141^ 

West  Atlantic 

Storey,  1939;  this  study 

C.  nebulosus  Smith 

.408 

178^ 

Red  Sea 

this  study 

C.  perryae  Storey 

.328 

Gulf  of  Mexico 

Storey,  1939;  Blache  and 

Cadenat,  1971 

C.  perryae  Storey 

.310 

179 
170^ 

Eastern  Atlantic 

Blache  and  Cadenat,  1971 

C.  springeri  (Ginsburg) 

.350 

Gulf  of  Mexico 

Ginsburg  1951 

C.  striatus  Smith 

.304 

192 

Red  Sea 

this  study 

Rounded  mean  value 
Type  specimen 

Characters  currently  used  for  species  separation  in  this  genus  include  the  coloration, 
body  depth,  preanal  length,  and  vertebral  number  (Table  5).  The  angle  of  the  gill  open- 
ing, sometimes  used  as  a  character  (Storey  1939),  is  of  little  use,  because  of  variability. 
The  species  most  closely  related  to  the  eastern  Pacific  species  C  cliffi  and  C.  eristigmus  ap- 
pear to  be  C.  muraena  Jordan  and  Evermann  and  C  perryae  Storey,  respectively.  The  re- 
markable similarity  of  each  species  pair  is  evidenced  in  the  body  depth  and  taper,  color- 
ation, preanal  length,  and  certain  osteological  characters.  The  members  of  each  pair  are, 
however,  separable  by  vertebral  number.  A  preliminary  osteological  study  of  several  spe- 
cies of  Callechelys  and  closely  related  genera  has  revealed  trenchant  differences  in  the 
urohyal  and  pectoral  girdle.  The  urohyal  is  either  a  simple  slender  filament  (in  C  eris- 
tigmus, C.  marmoratus,  and  C.  melanotaenius)  or  is  split  posteriorly  into  two  slender  diver- 


23 


g  =  C.  galapagensis 
e=  C.  eristigmus 


c  =  C.  cliffi 


Figure  5.  Distribution  of  the  eastern  Pacific  species  of  Callechelvs. 

gent  rays  (in  C.  clijfi,  C.  galapagensis,  and  C.  muraena).  The  pectoral  girdle,  as  in  most 
ophichthids  that  lack  pectoral  fins,  is  quite  reduced,  consisting  of  a  slender  cleithrum, 
supracleithrum,  and  small  rodlike  coracoid  and  scapula  (?).  Certain  species  of  Callechelvs 
(C.  eristigmus,  C.  marmoratus,  and  C  melanotaenius)  however,  have  lost  the  scapula, 
whereas  others  (C  hi  linearis,  C.  cliffi,  C.  galapagensis,  C.  luteus,  C.  muraena,  and  C.  nebu- 
losus)  have  retained  it.  The  retention  of  the  scapula,  along  with  the  simple  urohyal,  may 
represent  the  generalized  condition  in  Callechelys.  The  similarity  of  the  New  World 
forms,  as  well  as  their  dissimilarity  to  other  Indo-west  Pacific  species,  strongly  suggests  a 
common  ancestry  prior  to  the  closure  of  the  middle  American  seaway.  C.  galapagensis  ap- 
pears most  similar  to  the  central  Pacific  C.  luteus  Snyder,  but  differs  in  having  fewer  ver- 
tebrae, a  deeper  body,  and  a  spotted  anal  fin.  (Snyder  (1904:  517)  described  the  type  as 
having  "fins  colored  like  the  body";  our  specimen,  SIO  68-497,  1038  mm,  has  an  un- 
spotted anal  fin.)  None  of  the  three  Atlantic  species  shows  a  close  resemblance  to  C  ga- 
lapagensis. On  the  basis  of  present  evidence  we  therefore  suggest  that  the  three  eastern 


24 

Pacific  species  of  Callechelys  have  had  two  separate  histories,  with  one  species  arising 
from  a  Pacific  ancestor  and  the  other  two  with  a  common  New  World  ancestry. 

ACKNOWLEDGMENTS 

Carl  L.  Hubbs  read  the  manuscript  critically  and  Elizabeth  Parker  prepared  the  illustrations.  For  per- 
mission to  utilize  specimens  in  their  care,  thanks  are  due:  William  A.  Bussing,  Universidad  de  Costa  Rica;  Wil- 
liam N.  Eschmeyer  and  Warren  C.  Freihofer,  California  Academy  of  Sciences:  Lev  Fishelson,  Hebrew  Univer- 
sity; C.  Richard  Robins,  University  of  Miami  Marine  Laboratory;  and  Boyd  W.  Walker,  University  of 
California  at  Los  Angeles. 

This  is  a  contribution  from  the  Scripps  Institution  of  Oceanography. 

LITERATURE  CITED 

Blache,  J.  and  J.  Cadenat 

1971.  Contribution  a  la  connaissance  des  Poissons  anguilliformes  de  la  cote  occidentale  d'Afrique.  Dix- 
ieme  note:  les  genre  Mvrichihvs.  Bascanichthvs  et  Callechelvs  (Fam.  des  Ophichthidae).  Bull,  de 
IT.F.A.N.,  ser.  A,  33(1):  158-201. 
Bohlke,  J.  E.  and  J.  C.  Briggs 

1954.  Callechelvs  cliffi.  a  new  ophichthid  eel  from  the  Gulf  of  California.  Stanford  Ichthyol.  Bull.  4(4): 
275-278.  ■ 

Fowler,  H.  W. 

1932.  The  fishes  obtained  by  the  Pinchot  South  Seas  Expedition  of  1929,  with  descriptions  of  one  new 
genus  and  three  new  species.  Proc.  U.S.N.M.,  80(6):  1-16. 

1938.  The  fishes  of  the  George  Vanderbilt  South  Pacific  Expedition,  1937.  Zool.  results,  part  3.  Monog. 
Acad.  Nat.  Sci.  Philadelphia,  no.  2.  349  p. 

Ginsburg,  I. 

1951.  The  eels  of  the  northern  Gulf  coast  of  the  United  States  and  some  related  species.  Texas  J.  Sci.  3(3): 
431-485. 

Gosline,  W.  A. 

1951.  The  osteology  and  classification  of  the  ophichthid  eels  of  the  Hawaiian  Islands.  Pacific  Sci.  5(4):  298- 
320. 

Kanazawa,  R.  H. 

1952.  More  new  species  and  new  records  of  fishes  from  Bermuda.  Fieldiana,  Zool.  34(7):  71-100. 
Kaup,  J.  J. 

1856.  Catalogue  of  apodal  fish,  in  .  .  .  the  British  Museum.  London,  163  p. 
Nelson,  G.  J. 

1966.  Gill  arches  of  teleostean  fishes  of  the  order  Anguilliformes.  Pacific  Sci.,  20(4):  391-408. 
Pellegrin,  J. 

1912.  Sur  une  collection  de  poissons  des  Nouvelles-Hebrides  du  Dr.  Cailliot.  Bull.  Mus.  Hist.  Nat.,  Paris, 
18(4):  205-207. 

Smith,  J.  L.  B. 

1957.  The  fishes  of  Aldabra.  Part  IX.  (with  a  new  eel  from  East  Africa).  Ann.  Mag.  Nat.  Hist.,  ser.  12, 
10:833-842. 

1962.  Sand-dwelling  eels  of  the  Western  Indian  Ocean  and  the  Red  Sea.  Rhodes  Univ.  Ichthyol.  Bull.  24: 
447-466. 

Snyder,  J.  O. 

1904.  Catalogue  of  the  shore  fishes  collected  by  the  Steamer  Albatross  about  the  Hawaiian  Islands  in  1902. 
Bull.  U.S.  Fish  Comm.,  Vol.  22  for  1902:  513-538. 
Storey,  M.  H. 

1939.  Contributions  toward  a  revision  of  the  ophichthyid  eels.  I.  The  genera  Callechelvs  and  Bascanichthys, 
with  descriptions  of  new  species  and  notes  on  Mvrichihvs.  Stanford  Ichthyol.  Bull.,  1(3):  61-84. 


Scripps  Institution  of  Oceanography,  University  of  California,  San  Diego,  La  Jolla, 
California  92037 


PALEONTOLOGY  AND  PALEOECOLOGY 

OF  THE  SAN  DIEGO  FORMATION 

IN  NORTHWESTERN  BAJA  CALIFORNIA 


ROBERT  W.  ROWLAND 


TRANSACTIONS 

OF  THE   SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  3  9  JUNE  1972 


PALEONTOLOGY  AND  PALEOECOLOGY 

OF  THE  SAN  DIEGO  FORMATION 

IN  NORTHWESTERN  BAJA  CALIFORNIA 

ROBERT  W.  ROWLAND 


ABSTRACT-More  than  100  species  of  invertebrate  fossils  were  collected  from  gray  sandstones  of  the  San 
Diego  Formation  (Pliocene)  exposed  along  the  coast  of  northwestern  Baja  California.  Paleoecologic  eval- 
uation of  the  extant  species  and  associated  sediments  suggests  that  the  fauna  accumulated  at  shallow  sub- 
tidal  depths  near  a  submarine  bank  or  low  rocky  headland.  This  feature,  which  probably  formed  the  south- 
ern flank  of  the  Pliocene  San  Diego  embayment,  was  composed  of  a  Miocene  volcanic  formation  which 
underlies  the  fossiliferous  strata. 

The  light-colored  sandstone  underlying  the  southwestern  portion  of  San  Diego 
County,  California,  is  known  as  the  San  Diego  Formation  (Arnold  and  Arnold,  1902).  Ex- 
tinct species  indicate  a  Pliocene  age  for  the  formation  and  recent  workers  (ZuUo,  1969;  Wi- 
cander,  1970)  regard  it  as  late  Pliocene.  The  formation  extends  into  Baja  California,  where 
highway  construction  exposed  a  number  of  fossiliferous  sites  (Fig.  1).  This  paper  describes 
the  fauna  of  these  beds  and  presents  a  paleoenvironmental  reconstruction  of  the  area  using 
geological  and  paleontological  evidence. 

The  geology  of  the  Tijuana-Rosarito  Beach  area  was  mapped  and  described  by  Minch 
(1967).  In  this  area  the  San  Diego  Formation  is  underlain  by  a  thick  sequence  of  basalt,  tuff, 
sandstone  and  breccia.  Minch  named  these  rocks  the  Rosarito  Beach  Formation;  their  age 
is  middle  Miocene  (Hawkins,  1970;  Minch,  et  al.,  1970).  Minch  ( 1967)  recognized  two  mem- 
bers in  the  San  Diego  Formation.  The  lower  unit  is  composed  of  light  gray  to  light  brown 
sandstone  containing  several  discontinuous  lenses  of  conglomeratic  sand.  The  upper  mem- 
ber is  a  yellowish  brown  coarse  sandstone  containing  beds  of  cobble  conglomerate.  Over- 
lying the  San  Diego  Formation  and  capping  the  higher  mesas  is  a  reddish  brown  sandstone 
which  Minch  referred  to  the  Lindavista  Formation.  Fossiliferous  sand  of  Late  Pleistocene 
age  occurs  on  the  low  terraces  along  the  coast  (Valentine  and  Rowland,  1969). 

Structurally  the  area  consists  of  a  series  of  elongate  fault  blocks  which  parallel  the  Paci- 
fic coast.  These  blocks  are  separated  by  high  angle  normal  faults.  Movement  on  the  major 
faults  has  produced  a  west-tipped  stepped  structure  and  has  separated  the  fossiliferous 
strata  of  the  San  Diego  Formation  into  two  belts.  One  lies  within  %  mile  of  the  ocean,  and 
the  other,  less  clearly  defined,  more  than  1  Va  miles  inland.  The  relationship  between  the  two 
belts  is  not  clear,  for  no  stratum  can  be  correlated  in  both  deposits. 

Three  collections  of  fossils  (A-239,  A-241,  A-249)  were  made  from  the  inland  belt.  Lo- 
cality A-252  is  at  the  north  end  of  the  coastal  belt,  in  the  terrace  which  forms  a  base  for  the 
Pleistocene  deposits  of  the  Tijuana  Playas  area.  The  remaining  localities  are  exposed  in 
coastal  bluffs  and  canyons  from  the  International  Border  south  to  Rosarito  Beach.  Local- 
ities A-240,  A-241,  A-246,  A-248,  A-250  and  A-251  are  in  basal  conglomerates  less  than  20 
feet  above  the  base  of  the  formation.  Locality  A-245  is  104  feet  above  locality  A-240.  The 
remaining  localities  are  not  in  local  superposition  and  their  relative  ages  remain  uncertain. 
Complete  descriptions  of  the  fossil  localities  have  been  presented  elsewhere  (Rowland. 
1968). 

METHODS 

Thirteen  localities  were  sampled  semi-quantitatively.  At  each  locality  approximately 
1.5  cubic  feet  of  fossiliferous  matrix  was  collected.  The  number  of  specimens  of  each  species 
in  this  sample  is  recorded  in  Table  1 .  The  matrix  was  used  for  grain  size  analysis.  The  weight 
of  sediment  remaining  on  each  sieve  (from  -3<}>  to  -1-4  l/2<^)  was  calculated  as  a  percent  of 

SAN  DIEGO  see.  NAT.  HIST..  TRANS.  17(3):  25-32.  9  JUNE  197: 


26 


^ 


Figure  1.  Map  showing  location  of  fossil  localities  sampled. 

the  total  sample  weight,  then  plotted  against  sieve  size  to  produce  size  frequency  histograms 
(Fig.  2).  The  collected  material  is  deposited  in  the  Geology  Department,  University  of  Cali- 
fornia, Davis. 

Terminology  for  environmental  parameters  of  wave  exposure  and  water  depth  follows 
that  ofValentine(  1961:  Fig.  2).  The  littoral  zone  is  delineated  by  the  high  and  low  tide  lines 
and  the  inner  sublittoral  zone  by  the  low  tide  line  and  the  25  fathom  contour  Une.  Depth 
range,  geographic  range,  and  substrate  requirements  for  the  extant  species  were  compiled 
from  Grant  and  Gale  (1931),  Keen  (1971),  Morris  (1966),  Ricketts  and  Calvin  ( 1968)  and 
Valentine  (1958). 


RESULTS  AND  DISCUSSION 

Megafossils  collected  are  listed  in  Table  1.  Extinct  species  form  a  significant  portion  of 
the  fauna  and  are  important  in  confirming  the  age  of  the  strata.  Common  extinct  species 
are:  Dendraster  ashlevi  forma  ynezensis,  Acanthina  eniersoni,  Nassariiis gramniafus,  Terehra 
martini.  Ostrea  erici,  Anadara  irilineata,  Padnopecten  dilleri,  Patinopecten  healyi.  and 
Chlamys  parmeleei.  These  species  are  characteristic  of  Pliocene  strata  in  the  San  Diego, 
Ventura,  and  Santa  Maria  areas  of  California.  The  exact  age  of  the  San  Diego  Formation, 
to  which  the  deposits  in  northwestern  Baja  California  are  assigned,  is  not  known.  The  un- 
certainty arises  partly  from  problems  associated  with  establishing  the  Pliocene-Pleistocene 
boundary  in  the  marine  strata  of  California  (see  Bandy  and  Wilcoxon,  1970)  and  from  com- 


27 


0 

-1 

o 
a 

3 


V 


c 


H(^  i-Ztp         +3<t>         +4(^       PAN 


Figure  2.  Size-frequency  histograms  of  the  sedimentary  matrix  at  each  fossil  locality. 

plexities  within  the  formation  itself.  All  the  marine  Pliocene  deposits  of  western  San  Diego 
County  are  referred  to  the  San  Diego  Formation,  though  the  relationships  between  the 
strata  of  difterent  areas  are  presently  undeterminable.  For  example.  Mission  Valley  and  the 
Rose  Canyon  fault  separate  the  San  Diego  Mesa  with  its  Pliocene  faunas  from  the  Pliocene 
strata  at  Pacific  Beach.  These  deposits  may  represent  different  times  of  deposition  as  discussed 
by  Woodring  and  Bramlette  (1950:  104-107),  and  surely  they  represent  different  dep- 
ositional  environments.  The  Pacific  Beach  fauna  is  indicative  of  a  sand-cobble,  open  coast 
environment,  whereas  quiet  water  faunas  characterize  the  San  Diego  Mesa  deposits.  The 
collections  at  hand  contain  faunas  whose  components  show  aflfiliation  to  both  of  these 
biostratigraphic  zones. 

The  fauna  contains  species  representative  of  three  environments.  Calliostoma.  Oce- 
nehra,  Acanthina,  Tegula,  Thais,  Ostrea,  Hinnites,  andArchitecfonia  indicate  a  litloral-sub- 
littoral  rocky  coast  area.  Representatives  of  this  group  occur  at  1 1  localities  (see  Tables  1 
and  2).  Tivela,  Dentalium,  Cadulus,  Dendraster,  and  certain  lucinid  bivalves  suggest  a  sub- 
littoral  open  coast  environment  with  a  sand  substrate.  This  element  is  less  abundant  and 
occurs  at  four  localities.  Species  of  Nuculana,  Panope,  Spisula,  Tresus,  and  Nassarius  are 
predominant  elements  of  a  large  fauna  indicative  of  a  fine  mud  or  mud-sand  substrate  at 


28 


Tabic  1.   FosmK  Ironi  llii'  S.m  Dicgu  rornialion  in  northucsliTn  H.i|a  Calit'ornu 


Spo 


Antllo/oa. 

Astrangia  \:\.  A.  insignificata 

Nomland.  1915 
Balanophvllia  clfgans  Vernll. 

1846 
(iaNtropoda 

Haliotis  tulgens  Pluhppi,  1845 
Diodora  a  IT.  D.  iiuivquatts 

(Sowcrby.  18351 
CatUostonin  costalum  (Martyn. 

17841 
Calliostotna  gemmuLitum 

rarpcnicr.  1864 
Calltosrorna  kern  Arnold.  1 910 
Tegiila  funebralis  (A.  Adams, 

18541 
Teguta  gallinn  (!  orbes,  1850) 
AstToca  inncqunlis  (Martyn, 

18741 
Astraea  undosa  (Wood,  1828) 
Elilimn  alT  E.  rulila  Carpenter, 

1864 
Epitonium  hellastriatum 

(C'arpcnlcr.  1864) 
O/'alui  \'anco%tatum  Stearns, 

1875 
Tachvrhvmiiits  erosus  torma 

imior  Dall,  1919 
Tt/rrittila  iOopcri  Carpenter. 

1864 
Tumtctta  f^onostoina  torni.i 

hcmphilU  Merrlam.  1941 
Architecloitiva  nohtis  forma 

discus  drant  and  Gale.  1931 
Cerilhtdea  catifornka 

(Haldeman.  18401 
Calvptraea  irtnmtlbrts  Bmdenp. 

1834 
Trochito  radians  Lamark.  1822 
Crcpidula  onvx  .Sowerby.  1824 
l.unatia  Icwisii  ((iould.  1847) 
Polinices  rccluzianus  ( Deshayes, 

18391 
Trivin  sutt^anea  (Sowerby. 

18321 
h'usttriton  aregonensis 

(Redlield.  18461 
Bursa  californica  Hinds,  1843 
'  Ccralustonia  foliata  (Ctmelin, 

18451 
Jalun  j estiva  (Hinds,  1844) 
Ocenebra  aft.  O.  fraseri 

lOldroyd.  19201 
Acanthina  emersoni  Hertlein 

and   Mlison.  1959 
Tliais  enmrf^nata  (Deshayes, 

18391 
Thais  lamellosa  ic;melin.  17901 
Cantharus  aff  C  ringens 

(Reeve.  18461 
Kcllelia  all.  A.  kellelii   (1  orbes, 

18501 
Ainphissa  el.  A   reticulata  Dall. 

1916 
%Utrella  ^ausapata  (tioiild. 

18541 
Sassarius  californianus 

(Conrad.  18561 
Xassarius Kraninialus  (Dall. 

19171 
Nassariits  mendieus  forma 

mdisputabllis  (01dr..yd.  19271 
Nassariits  e  t .  .V  pcrpinguis 

(Hinds.  18441 
Barharojusiis  barbarcnsis  (Trask. 

18551 
Psephaea  oregonensis  I  Dall. 

19071 
(Hivelta  btplteata  (Sowerby. 

1825) 
Olnelta  pedrtmna  (Conrad.  1855) 
Canceltaria  fugleri  (Arnold.  19071 
Caneellaria  tritonidea   Clabb. 

1866 
Caneellaria  rapa  (Nomland.  1919 
Conns  eatifornieus  (Hinds,  I  844 1 
Conus  aff.  C  reeurvus 

Broderip.  1833 
Conus  sp. 

Tercbra  niarltnt    1  nj.'lish.  1914 
Terebra  sp, 
Clavus  ef.  C  empvrosia  (Dall. 

18991 


I  nlversin    ol  (  ahlnrnia,  Dans        I  ossil  1  oialil  les        A  series 

239     240     241      243     244     245     246     247     24K     249     25(1     251      252 


10 


50 


1 

15 
1 


I 
30 


10 


25 


20 


2 
15 


1 

5 
10 

cf7 


12      21 


48 


ef2 
1 


25 
1 


15 


25      4 
5 


4      2 
1 


sublittoral  depths  or  in  a  semi-protected  area;  this  fauna  occurs  at  11  localities.  Because 
Cerithidea  califoniica.  Bulla  gouldiana,  and  Cryptomya  californica  are  rare  or  absent  in 
these  collections  but  are  dominant  species  in  modern  coastal  lagoons  (Warme,  1971),  I  be- 
lieve this  last  complex  represents  a  sublittoral,  open  coast  environment  rather  than  a  pro- 
tected lagoon. 

The  hydrographic  regime  indicated  by  the  fossil  faunas  is  paradoxical.  Commonly  a 
locality  contains  representatives  of  extant  species  whose  present  geographic  ranges  along 


29 


Tjblo  1  (Lunliruii.\l  i 


SpC.K- 


Titrridac  sp. 

Ategasurcula  carihiucruind 

iGahh,  lSh5l 
Mc^asurcula  irvoniana  Klabb. 

1866) 
Volvolclla  cylinJrua  ((  urpL-ntcr. 

1S64) 
Acteocim  a(t.  .-1.  inculla 

((;ou!d  and  Carpcnlcr.  1857) 
Bivalvia 

Acib  castrcnsis  (Hindv  1834) 
Sachi-m  taphria  (Dall.  18')6) 
Afiadara  tritmeata  (Conrad. 

1856) 
CAycymeris  cf.  C  subobsoleta 

(CariK-nlLT.  1864) 
Myliltis  c(.  coalingcusis 

Arnold,  WIO 
Myfilus  sp. 

Ostrea  erici   McrlU-m.  1929 
Ostrea  vesperliiia  (Conrad. 

1854) 
Ostrta  sp. 
Chlamys  (Argopectcn)  ctrcularis 

(Sowcrby.  1835) 
Chlamys  ha slata  forma  hcruia 

(Gould.  1850) 
Chlamvs  parmcleci  { Dall. 

189S) 
Chlamys  sp. 
I.vropccten  ccrroscnsis  ((iahb. 

1866) 
Pccicn  helliis  torma  hemphilli 

Dall.  1879 
Pctlcnslcarnsi  Dall.  1879 
Pcclen  (Palinopectcnl  Jillcri 

Dall,  1901 
Pcctcn  (Palinopectcnl  hcaiyi 

Arnold.  191)6 
Hinnites  sp. 

Pododesmtis  ccpio  ((ira\.  1850) 
Cvclocardia  californua  (Dall. 

1903) 
l.ucinonia  annulata  (Rcc-vc.  1850) 
l.ucina  cxcavata  Carpcntfr. 

1857 
Lucinisca  nuiiallii  ((  onrad. 

1837) 
Parvihicina  tcnuisculpta 

CarpL-nlcr.  1864 
Thyasira  hisccta  ((  onrad.  1849) 
Ijicvicardiuni  quadra^cnarium 

iConrad.  1837) 
Thcia  <iiiiltoriim  (Maui.'.  1823) 
Dosinia  ponderosa    (iray. 

1  838  nc-u  torni 
Chionc  lI.  elcsmerensis 

1  n^hsh.  1914 
Chionc  fcrnandocnsis  il  nulisli. 

1914 
Prntofhaca  lencrrima 

(C  arpi-n(cr.  1856) 
Spisula  hemphilli  (Dall.  1SM4) 
Madra  sp.  ^1 
Mai  tra  sp.  *2 
\factra  sp.  #3 
Trcsus  nutlallii  ((  onrad. 

1837) 
Garicdentula  (Clabb.  1868-1869) 
Semclf  ruhropuia  Dall.  1871 
Siliqua  hnida  ((onrad.  1837) 
Corbtita  ^hhtjormis 

(Sowcrby.  1833) 
Panopc abrupta  (Conrad.  1849) 
Scaphopoda; 

Dcntalium  ncohcxagoniim 

Sbarpand  Cilsbry.  1897 
Cadiilus  fusijormis    I'lKbry  and 

Sharp.  1897 
Arthropodj 

Balamn  (BalamtsI  sp. 
Balanus  (Balamis)  f^rcgaritis 

(Conrad.  1856) 
I  ».hinodi.Tmaia: 

Dendraster  ashlvyi  lornia 

ynczcnsis    Ki-w.  1919 
I  Lhint)id  spines 
hucidaris  Ihourasii  ( VaUni_icnncs. 

1846) 
Vfrtcbrata; 

Oircharodon  arnoldi    lordan. 

1909 


UnivcrsiCy  of  Calilorniu.  Davis  -   I  oiiil  Localilics       A  scries 
239  240  241  24?  244  245  246 


249  250 


4 

21 


15 

•t 

5 

1 

25 

3 

20 

5 

2 

1 

1 
II 

5 

3 

26 

13 


14 
1 


13 

13 
19 

5 

25 


25 

37 

5 


19 

5 


8 
I  I 


12 


1 
15 


10 


I 

25 


75 
1 


20 


the  western  coast  of  North  America  do  not  overlap.  For  example.  A-246  contains  elements 
of  all  habitats  as  well  as  three  Trivia  sanguinea.  which  is  living  only  in  the  Gulf  of  California 
and  southward  and  a  variety  oi^  Dosinia  ponderosa.  At  present  Dosina  is  not  found  north  of 
Scammon's  Lagoon,  Baja  California.  The  same  locality  also  yielded  CaUiostoma  cosiaium, 
Parvihicina  tenuiscidpta  and  Chlamys  hastata  forma  hericia,  which  range  from  southern 
California  to  Alaska,  as  well  as  Fusitriton  oregonensis.  a  submergent.  stenothermal,  frigiphi- 
lic  species  which  lives  intertidally  north  of  Oregon  and  is  not  found  at  depths  less  than  80 


30 


Table  2:   Correlation  between  faunal  elements  and  preferred  sediment  type. 
F  ~  Faunal  element  present,  S  -  Sediment  peak  developed 


Locality 
Number 


Sediment  coarser 
tlian-2.0</) 

Rocky  substrate. 
Open  coast  fauna 


Sediment  between 
-2.0^  and +2.0  <^ 

Sand  substrate. 
Open  coast  fauna 


Sediment  finer  than  +2.0  </> 


Fine  sand  or  mud  substrate. 
Offshore  fauna 


A-239 


A-240 


A-241 


A-243 


A-244 


A-245 


S 
F 


S 
F 

S 
F 


F  (2  species) 

S 
F 


S 
F 

S 
F 

S 
F 


S 
F 

S 
F 


A-246 

S 

F 

A-247 

S 

F 

A-248 

S 

F 

A-249 

S 

F 

A-250 

A-251 

S 

F 

A-252 

S 

F 

s 

F 


S 
F 


S 
F 

S 
F 

S 
F 

S 
F 


F  (2  species) 

S 
F 


S 
F 


fm.  off  southern  California  (Smith,  1970:  493).  Depths  of  this  magnitude  are  incompatible 
with  environments  indicated  by  the  other  species  collected  at  this  locality. 

The  concurrence  of  these  northern  and  southern  elements  and  the  presence  of  Fusitri- 
ton  is  not  readily  explicable.  Presumably  nearshore  upwelling  of  cool  waters,  perhaps  into 
coastal  waters  warmer  than  presently  found  off  southern  California,  was  important  in  al- 
lowing these  thermally  anomalous  species  to  coexist  (Valentine  and  Emerson,  1961:  617- 
618). 

Data  from  the  sedirnent  analysis  can  be  used  to  clarify  the  paleoecological  inter- 
pretations. Sediment  samples  from  the  center  of  the  study  area  are  poorly  sorted  (Fig.  2). 
The  distinct  size  concentrations  of  coarse  and  fine  material  suggest  that  these  sediment  sizes 
were  not  transported  together  (see,  for  example,  the  curve  for  locality  A-246).  The  coarse 
fraction  of  the  sediment  may  represent  either  lag  gravels  of  underlying  volcanic  rocks  or 
cobbles  transported  by  storm  waves.  Presumably  the  sand-  and  silt-size  material  infiltrated 
the  coarser  material.  Samples  with  the  best  sorting  (A-249,  A-250,  and  A-252)  are  found  on 


31 

the  margins  of  the  area.  The  unimodal  sand  of  A-249  impHes  sorting  by  beach  transport.  At 
most  locaUties  the  sediment  size  indicated  by  the  size-frequency-histograms  is  compatible 
with  the  substrate  on  which  the  extant  species  are  commonly  found.  For  example,  the  fauna 
of  locality  A-246  contains  species  indicative  of  the  three  environments  described  above. 
The  size-frequency  distribution  curve  for  this  locality  has  peaks  (-3</>,  +  \i>  and  -t-3.5</>). 
which  correspond  to  the  substrate  of  each  of  these  environments.  Interrelationships  be- 
tween fauna  and  sediment  are  shown  in  Table  2.  There  is  significant  accord  between  the 
fauna  and  the  sediment  in  23  of  the  29  cases.  The  heterogeneous  nature  of  the  fauna  and  the 
poor  sorting  of  the  sediments  suggests  that  the  environments  that  contributed  to  the  fossil 
beds  were  in  close  proximity  to  the  site  of  burial. 

In  summary,  the  environment  of  deposition  of  the  Pliocene  strata  of  northwestern  Baja 
California  can  be  reconstructed  as  follows:  the  Rosarito  Beach  Formation  does  not  extend 
northward  into  California;  presumably  it  formed  the  southern  flank  of  the  San  Diego  em- 
bayment  as  a  bank  or  headland.  The  faunas  studied  accumulated  on  this  feature  at  subti- 
dal.  inner  sublittoral  depths  in  areas  where  patches  of  fine  sand  and  mud  were  interposed 
between  cobble  beds  and  rocky  exposures. 

ACKNOWLEDGEMENTS 

J.  A.  Minch.  University  of  California.  Riverside,  provided  me  with  an  excellent  field  orientation.  J.  W.  Valen- 
tine gave  generously  of  his  time  and  information  throughout  the  study.  Figures  were  drafted  by  L.  Valentine  and 
R.  Darden. 

LITERATURE  CITED 

Arnold,  D.,  and  R.  Arnold 

1902.  Stratigraphy  of  southern  California.  J.  Geol.  10:  117-138. 

Bandy,  O.  L.,  and  J.  A.  Wilcoxon 

l'970.  The  Pliocene- Pleistocene  boundary,  Italy  and  California.  Geol.  Soc.  Amer.  Bull.  81:  2939-2948. 

Grant,  U.  S.,  IV,  and  H.  R.  Gale 

1931.  Catalogue  of  the  marine  Pliocene  and  Pleistocene  molluscs  of  California  and  adjacent  regions.  San 
Diego  Soc.  Nat.  Hist.  Memoir  1 
Hawkins,  J.  W. 

1970.  Petrology  and  possible  tectonic  significance  of  Late  Cenozoic  volcanic  rocks,  southern  California  and 
Baja  California.  Geol.  Soc.  Amer.  Bull.  81:  3323-3338. 

Keen,  A.  M. 

1 97 1.  Sea  shells  of  tropical  west  America.  Stanford,  Stanford  Univ.  Press. 

Minch,  J.  A. 

1967.  Stratigraphy  and  structure  of  the  Tijuana-Rosarito  Beach  area,  northwestern  Baja  California.  Mexico. 
Geol.  Soc.  Amer.  Bull.  78:  1155-1178. 

Minch,  J.  A.,  K.  C.  Schulte,  and  G.  Hofman 

1970.  A  middle  Miocene  age  for  the  Rosarito  Beach  Formation  in  northwestern  Baja  California,  Mexico. 
Geol.  Soc.  Amer.  Bull.  81:  3149-3154. 

Morris,  P.  A. 

1966.  A  field  guide  to  the  shells  of  the  Pacific  coast  and  Hawaii.  Boston,  Houghton-Mifflin. 

Ricketts,  E.  F.,  and  J.  Calvin 

1968.  Between  Pacific  tides.  Revised  by  J.  W.  Hedgepeth.  Stanford,  Stanford  Univ.  Press. 
Rowland,  R.  W. 

1968.  Paleontology  of  the  San  Diego  Formation  in  northwestern  Baja  California,  Mexico.  M.S.  thesis.  Uni- 
versity of  California.  Davis.  60  p. 

Smith.  J.  T. 

1970.  Taxonomy,  distribution  and  phylogeny  of  the  Cymatiid  gastropods  Argobuccinum.  Fusitriton,  Me- 
diargo.  and  Priene.  Bull.  Amer.  Paleo.  56:  445-573. 

Valentine,  J.  W 

1958.  Paleoecologic  molluscan  geography  of  the  Californian  Pleistocene.  Ph.D.  dissertation.  University  of 

California,  Los  Angeles.  458  p. 

Valentine,  J.  W. 

1961.  Paleoecologic  mollu-scan  geography  of  the  Californian  Pleistocene.  Univ.  California  Publ.  Geol.  Sci. 
34:  309-442. 

Valentine,  J.  W..  and  W.  K.  Emerson 

1961.  Environmental  interpretation  of  Pleistocene  marine  species:  a  discussion.  J.  Geol.  69:  616-618. 

Valentine.  J.  W.,  and  R.  W.  Rowland 

1969.  Pleistocene  invertebrates  from  northwestern  Baja  California  Del  Norte,  Mexico.  California  Acad.  Sci. 


32 


Proc.  36:  511-530. 

Warme,  J.  E. 

1971.  Paleoecological  aspects  of  a  modern  coastal  lagoon.  Univ.  California  Publ.  Geol.  Sci.  87:  1-131. 

Wicander,  E.  R. 

1970.  Planktonic  foraminifera  of  the  San  Diego  Formation,  p.  105-117.  In,  Pacific  slope  geology  of  northern 
Baja  California  and  adjacent  Alta  California.  Amer.  Assoc.  Petrol.  Geol.  (Pacific  Section)  Fall  Field 
Trip  Guidebook. 

Woodring,  W.  P.,  and  M.  N.  Bramlette 

1950.  Geology  and  paleontology  of  the  Santa  Maria  district,  California.  U.  S.  Geol.  Survey  Prof.  Paper  222: 
1-185(1951). 
Zullo,  V.  A. 

1969.  Thoracic  Cirripedia  of  the  San  Diego  Formation,  San  Diego  County,  California.  Los  Angeles  Co.  Mus. 
Contr.  Sci.  159:  1-25. 


Department  of  Geology,  University  of  California,  Davis,  California,  95616 


5»a/ 


Lip  V  .^'^ 
HARVARQ 


SEISMIC  RISK  IN  SAN  DIEGO 


ROBERT  B.  McEUEN  AND  CHARLES  J.  PINCKNEY 


TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  4  19  JULY  1972 


SEISMIC  RISK  IN  SAN  DIEGO 


ROBERT  B.  McEUEN  AND  CHARLES  J.  PINCKNEY 

ABSTRACT.— Data  from  artificial  earth  satellites  suggest  that  the  regional  shear  stress  responsible  for  earth- 
quakes is  relatively  high  in  the  San  Diego  region.  The  direction  of  this  stress  is  related  to  the  orientation  of  the 
East  Pacific  Rise  and  the  Gorda  Rise.  The  apparent  nonequivalence  of  strain  accumulation  to  release  along 
active  faults  northeast  of  San  Diego  also  suggests  that  shear  stress  is  accumulating  over  a  wide  zone  and  that 
seismic  activity  may  occur  on  adjoining  but  presently  less  active  faults. 

Maximum  credible  earthquake  magnitudes  for  San  Diego  proper  and  for  the  San  Clemente  Island  fault 
zone  are  6.8  and  7.7.  respectively.  These  events  are  highly  unlikely  but  should  be  considered  in  designing 
critical  structures  such  as  nuclear  power  plants.  Regarding  probable  earthquakes,  San  Diego  falls  in  the  sec- 
ond most  active  of  six  regional  zones  in  the  United  States  in  terms  of  strain  release  and  in  the  most  active  of 
three  zones  in  terms  of  regional  seismic  risk.  In  the  recent  past,  how  ever,  most  of  the  regional  strain  energy  has 
been  released  outside  the  immediate  San  Diego  area.  Our  analysis  of  the  mechanics  of  strain  release  indicates 
that  San  Diego  proper  will  experience  an  acceleration  of  0.2  g  on  worse  ground  approximately  every  60  years. 
The  area  ofwor.se  ground  is  around  the  bays,  where  a  two-fold  amplification  of  bedrock  acceleration  could 
occur.  The  most  likely  source  for  this  acceleration  is  the  Elsinore  fault. 

A  magnitude  6.3  or  greater  earthquake  within  the  Continental  Borderland  could  produce  seismic  sea 
waves  capable  of  inundating  San  Diego's  low  lying  coastal  areas,  but  the  occurrence  of  such  an  event  is  un- 
likely. 

The  purpose  of  this  paper  is  to  define  seismic  risk  in  San  Diego,  CaUfornia.  We  have 
attempted  to  strike  a  balance  between  an  "it  couldn't  happen  here"  attitude  and  one  that 
states  "it's  only  a  matter  of  time."  Most  of  the  data  on  which  our  conclusions  are  based  are 
available  in  the  literature.  Dr.  Charles  F.  Richter's  (1959)  comment  sets  the  stage  for  the 
discussion  which  follows: 

"There  has  been  a  general  impression  that  earthquake  risk  does  not  exist  at  San  Diego,  historical  records  to 

the  contrary  being  forgotten  or  ignored.  Older  structures  were  erected  with  no  close  attention  to  soundness. 
During  and  since  World  War  II,  population  has  increased  enormously,  and  the  city  area  has  expanded  at  a 
pace  hardly  consistent  with  careful  construction  and  inspection.  Fortunately  most  of  this  expansion  has  been 
over  the  higher  ground..." 

SEISMOTECTONICS 

The  seismotectonic  framework  of  the  San  Diego  area  can  be  best  understood  by  ana- 
lyzing seismotectonics  on  a  global  scale  and  by  then  studying  in  more  detail  those  parts  that 
have  direct  bearing  on  regional  and  local  seismicity. 

GLOBAL  SEISMOTECTONICS 

The  seismic  stresses  responsible  for  earthquakes  are  thought  to  originate  in  the  up- 
welling  of  hot  light  material  of  the  earth's  mantle.  As  this  material  convects  it  causes  hori- 
zontal stresses  in  the  earth's  crust.  The  presence  of  such  convection  cells  below  the  crust  is 
indicated  by  slight  changes  in  density  of  the  material  undergoing  convection.  These  density 
variations  in  the  earth  cause  deviations  in  the  orbits  of  artificial  satellites.  Figure  1.  shows 
satellite-deduced  density  variations  for  regions  of  high  seismic  activity  (Schwiderski.  1968). 

The  Western  Pacific  (Fig.  la)  represents  one  of  the  most  seismically  active  regions  in 
the  world.  Here,  zones  of  maximum  seismic  activity  fall  between  low  density  areas  presum- 
ably indicative  of  upwelling  and  high  density  areas  indicative  of  convection  downturn. 
Along  the  western  coast  of  North  and  South  America  (Fig.  lb)  the  zone  of  seismic  activity 
also  lies  between  an  area  of  convective  upwelling  and  downturn.  This  eff"ect  is  pronounced 
along  the  western  coast  of  South  America  and  we  believe  the  western  coast  of  North  Amer- 
ica is  being  influenced  by  similar  convective  stress.  But,  the  magnitude  of  this  stress  appears 
to  be  less  than  in  the  high  seismicity  zones  of  the  Western  Pacific  or  along  the  western  coast 
of  South  America.  Note,  particularly,  that  the  zones  of  seismic  activity,  almost  without  ex- 
ception, occur  where  the  rate  of  change  of  density  at  the  mantle  surface  is  a  maximum,  that 

SAN  DIEGO  SOC.  NAT.  HIST..  TRANS.  17(4):  33-62,  19  JULY  1972 


34 


30' 


0  - 


30- 


L  EGE  ND 
X         SHAL  LOW     DEPTH 
O         INTERMEDIATE     DE  PTH 
•        GREAT    DEPTH 


Figure  1.  Contour  map  of  density  anomaly  p'  (in  g/(dm)"^)  at  the  surface  of  the  mantle.  Contour  lines 
change  by  8p' =  1.2.  Earthquakes  after  Gutenberg  and  Richter  (1954).  a.  Western  Pacific  showing 
earthquake  zones  along  unstable  strips  between  low-density  sources  and  high-density  sinks,  b.  Eastern 
Pacific  showing  earthquake  zones  along  unstable  strip  between  low-density  sources  and  high-density 
sinks. 

is,  where  the  contour  lines  in  Figure  1  are  most  closely  spaced.  San  Diego  occurs  in  such  an 
area. 

REGIONAL  SEISMOTECTONICS 

Figure  2  shows  regional  tectonic  features  thought  to  have  direct  bearing  on  the  seismi- 
city  of  San  Diego  and  adjoining  areas.  Figure  3  shows  locations  of  earthquake  epicenters  for 
that  portion  of  the  area  lying  south  of  34°N  latitude.  This  area  can  be  located  on  Figure  lb 
by  noting  where  the  120°W  longitude  line  intersects  the  western  coast  of  North  America. 

Between  the  zone  marked  East  Pacific  Rise  at  the  bottom  of  Figure  2  and  the  zone 
marked  Gorda  Rise  at  the  top,  there  exists  a  series  of  en  echelon  faults  extending  from  the 
Tamayo  fracture  zone  to  the  San  Andreas  Fault  Zone.  All  these  faults  have  a  right-lateral 
sense  of  displacement,  with  the  eastern  side  moving  southeast  relative  to  the  western  side. 

Ridge  crests,  such  as  the  East  Pacific  Rise  and  the  Gorda  Rise,  are  areas  in  which  new 
oceanic  crust  is  being  created.  Molten  material  presumably  wells  up  within  these  zones  and 
in  so  doing  shoulders  aside  older  material.  As  this  process  is  repeated  the  area  on  both  sides 
of  the  ridge  crest  moves  laterally.  Records  of  seismic  activity  indicate  that  zones  of  shallow 
earthquakes  are  associated  with  ridge  crests;  these  zones  also  tend  to  align  between  areas  of 
mantle  upwelling  and  downturning  as  delineated  by  upper-mantle  density  anomalies 
(Schwiderski.  1968).  The  apparent  extension  of  the  East  Pacific  Rise  into  the  Basin  and 
Range  Province  (Fig.  2)  follows  the  zone  of  maximum  upper-mantle  density  change  (Fig. 
lb). 

The  large  arrows  perpendicular  to  the  strike  of  the  East  Pacific  Rise  and  the  Gorda 


35 


Figure  2.  Fractures  and  spreading  centers  along 
part  of  the  Pacific  Coast  of  North  America.  S.D. 
marks  the  location  of  San  Diego.  Oceanic  fracture 
zones  (FZ)  and  continental  faults  (F)  are  solid 
black  lines,  dashed  where  uncertain.  S.A.F.— San 
Andreas  Fault;  D.V.F.— Death  Valley-Furnace 
Creek  Wash  Fault;  O.V.F.-Owens  Valley  Fault; 
G.F.— Garlock  Fault;  E.F.— Elsinore  Fault; 
S.J.F.— San  Jose  Fault;  A.B.F.— Agua  Blanca 
Fault;  S.R.F.— Santa  Rosalia  Fault;  S.C.F.— San 
Clemente  Fault;  R.F.— Rampart  Fault.  Post- 
ulated spreading  centers  along  the  crest  of  the 
East  Pacific  Rise  and  its  possible  continuation  into 
the  continent  shown  in  gray.  Representation  of 
spreading  centers  in  the  Basin  and  Ranges  is  sym- 
bolic, indicating  a  region  of  crustal  extension 
(Modified  from  Elders,  et  al.,  1970). 


Rise  indicate  the  relative  direction  in  which  stress  is  appHed  due  to  forces  produced  along 
these  spreading  centers  (Fig.  2).  The  right-lateral  movement  of  the  series  of  faults  is  accom- 


36 


modating  this  stress,  and  one  is  tempted  to  consider  the  whole  zone  of  fauhing  as  a  shear  set. 
Gastil  et  al.  (in  press)  note,  however,  this  simphstic  view  is  not  consistent  with  the  right- 
stepping  nature  of  the  en  echelon  fault  breaks. 

In  the  San  Diego  region,  the  zone  of  possible  stress  accumulation  reaches  its  widest 
extent  in  an  area  bounded  on  the  east  by  the  San  Andreas  fault  and  on  the  west  by  the 
Rampart  fault,  which  marks  the  western  hmit  of  the  Cahfornia  Continental  Borderland. 


«a 

J 

.  ^' 

\   ■••••    • 

i 

• 

••• 

• 

•      • 

• 

1)6 

\ 

X120*W30*N 

/ 

(      ■■'.:. 

V  •  •   •; 

) 
< 

•      / 

•  •    fJ  * 

•        f 

1     • 
•     /   •  • 

1     *  * 

>x  v. 

...     '-N 

*      •             • 

• 

• 

\ 

\ 
\ 

\ 

/ 

f        •  \ 

Y 

\ 
\ 

• 

\ 

•  1 

.\ 

) 

\ 

'^ 

' 

< 

* 
• 

T*'    * 

J 

\ 
\ 
\ 
\ 

\ 

^4 

• 

\ 

\      i 

.  *  1 

• 

112* 

W30*Nj( 

X  114*W  26*N 

.            V 

( 

12*W26Nr' 

*                *                                      ^ 

*                        \ 

• 

L 

^ 

•                          r 

• 

• 

L 

Figure  3.  Earthquake  epicenters  with 
magnitudes  of  4  or  greater  between 
1958  and  1968  (from  Gastil,  Allison, 
and  Phillips,  in  press). 


Earthquakes  occur  in  areas  where  differential  stress  exceeds  the  shearing  resistance  of 
the  material  in  the  area,  and  rupture  occurs.  If  diff"erential  stress  is  applied  across  the  broad 
zone  of  en  echelon  faults  strain  should  be  occurring  throughout  the  zone.  A  necessary  con- 
straint to  a  surface  rupture  of  considerable  length  is  that  the  rate  of  surface  strain  accumula- 
tion must  be  a  maximum  in  the  immediate  zone  of  faulting.  If  not,  and  large  surficial  plates 
are  moving  in  opposite  directions  tangential  to  the  fault  (Fig.  2),  the  faulting  must  spread 
through  time  over  a  wide  lateral  area.  With  these  thoughts  in  mind,  let  us  examine  the 
strain-accumulation  and  strain-release  history  of  the  onshore  regions  adjoining  San  Diego. 

Onshore  strain  accumulation.    To  determine  strain  accumulation,  geodetic  surveys 


37 


should  be  carried  out  in  intervals  between  earthquakes.  Following  the  magnitude  7. 1  Impe- 
rial Valley  earthquake  of  1940,  the  Imperial  Valley  triangulation  network  was  resurveyed;  a 
second  resurvey  was  made  in  1954.  and  a  third  in  1967.  Scholz  and  Fitch  ( 1969).  summa- 
rized the  data  obtained  in  the  area  of  the  Imperial  fault,  which  ruptured  along  40  miles  of  its 
length  in  1940.  and  therefore  represents  a  zone  along  which  strain  accumulaton  should  at- 
tain a  local  maximum.  They  concluded  that  slip  occurred  along  the  Imperial  fault  between 
surveys,  releasing  about  15  per  cent  of  the  accumulated  strain.  Their  slip-corrected  data 
(Fig.  4)  verifies  that  shear  strain  is  accumulating  at  a  higher  rate  near  the  fault.  The  data  for 
the  1 94 1  - 1 954  period  are  shown  in  vector  form  in  Figure  5a.  The  fact  that  the  shear  strain  is 
appreciable  at  considerable  distance  from  the  fault  zone  led  Scholz  and  Fitch  ( 1969)  to  state 
that  "the  Elsinore.  San  Jacinto,  and  Mission  Banning  Creek  faults... may,  in  fact,  testify  to  a 
'spreading'  of  the  fault  zone  due  to  a  nonequivalence  of  strain  accumulation  and  release..." 

Mi  les 


10 


20 


30 


40 


E 


c 

9) 

E 

u 
O 


50 


30   — 


10 


1 

1 

1 

\. 

-^o 

•    East 

.°^o 

0  West 

^»    o 

_      'o^ 

•^ 

b^to 

• 

•  •:? 

.o 

0 

\* 

\ 

- 

• 

• 

^ 

--.      o 

1 

•      lo 

—  20 


—  10 


u 

c 


20 


40 


60 


Distance  —  k 


m 


Figure  4.  Strain  accumulation  parallel  to  the  Imperial  Fault  versus  distance  from  the  fault  for  the  period  1941- 
1954.  The  data  has  been  corrected  for  the  observed  slip  and  superimposed  on  one  side  of  the  fault  (Scholz  and 
Fitch.  1969). 

Savage  (1970)  shows  that  the  strain  that  accumulated  between  1941  and  1967  is  ap- 
proximately twice  that  accumulated  between  1941  and  1954  and  concludes  that  for  the  Im- 
perial Valley  triangulation  network  the  strain  rate  is  about  0.4-1-0.1  /x  strain/yr  over  a  zone 
100  km  (63  miles)  wide. 

Onshore  strain  release. —Strain  release  can  be  obtained  bv  direct  measurement  if  good 
geodetic  control  is  available  prior  to  the  release  of  strain  by  an  earthquake.  Figure  5b  shows 
retriangulation  results  obtained  following  the  1940  Imperial  Valley  earthquake.  Such  good 
geodetic  control  in  the  past  has  been  the  exception  rather  than  the  rule,  and  seismologists 
have  had  to  rely  on  empirical  relationships  to  establish  probable  strain  release  associated 
with  earthquakes. 

Figure  6  shows  a  strain-release  map  derived  from  a  two  year  study  of  28.000  earth- 
quakes in  the  conterminous  United  States,  including  16,000  in  California.  The  map  is  useful 
in  that  it  shows  the  relative  rate  of  seismotectonic  activity  in  various  areas.  Note  that  a  zone 
of  maximum  seismotectonic  activity  lies  just  east  of  San  Diego  and  that  the  San  Diego  area 
falls  in  the  second  most  active  of  zones. 

Offshore  faiilting.-Moore  (1969)  published  a  structural  map  of  the  California  Conti- 
nental Borderland  based  largely  upon  the  interpretation  of  seismic  reflection  profiles.  These 
profiles  commonly  show  folded  structure  within  strata  and  either  directly  or  indirectly  the 
location  of  faults.  He  concluded  that  the  primary  offshore  structural  pattern  comprises  two 


38 


^Coltpotr  ia 

\ 

Westmoreland  \ 

\@Brawley 

\ 


(S)Hollville 


El  Ceniro 


1941-1954 


MILES 
10  20 


30 


VECTORS 
0       1       2       3      4       S 


in  feet 


•^ 


1939-1941 


® 


Colipatr  ia 


^  ^ 


MILES 

0  20  30  40 

I 1 ■ 

VECTORS • 
0  2  4 


Westmoreland 


N 


\ 


Figure  5.  a,  Retriangulation  strain-accumulation  results,  U.S.  Coast  and  Geodetic  Survey,  Imperial  Valley,  1941- 
1954.  b,  Retriangulation  strain-release  results,  U.S.  Coast  and  Geodetic  Survey,  Imperial  Valley,  1939-1941  [after 
Whitten  by  C.  R.  Allen]  (Ritcher,  1958). 

sets  of  faults,  a  set  trending  northwest  and  another  east-northeast.  The  northwest  trend  so 
predominant  onshore  clearly  carries  through  to  most  of  the  Borderland  as  the  principal 
set  and,  where  topographic  offsets  are  well  enough  developed  to  be  significant,  movement 
appears  to  be  right-lateral  (Fig.  7).  This  conclusion  is  consistent  with  first-motion  studies 
of  earthquakes  which,  with  one  important  exception,  indicate  right-lateral  movement  on 
northwest  trending  faults  (Gutenberg,  1941;  Allen,  1960).  The  exception  is  the  5.9  magni- 
tude earthquake  that  occurred  off  the  southeast  tip  of  San  Clemente  Island  in  1951.  First 
motion  for  this  earthquake  can  only  be  explained  by  assigning  it  a  large  component  of  dip- 
shp  movement  (Allen,  1960). 

The  dominant  fault  of  the  inner  zone  of  major  northwest  trending  faults  is  the  San  Cle- 
mente Island  fault  extending  from  the  eastern  side  of  San  Clemente  Island  to  the  Cabo  Col- 
nett  area  of  Baja  California,  Mexico.  Another  major  fault  forming  the  eastern  face  of  Santa 
Catalina  Island  may  be  continuous  with  a  fault  along  the  western  boundary  of  the  San 
Diego  Trough.  Moore  (1969)  suggested,  based  on  the  sedimentation  and  structure  of  the 
Borderland  and  records  of  modern  earthquake  epicenters,  that  about  1  million  years  ago 
formation  or  rejuvenation  of  the  Agua  Blanca  fault  initiated  strike-slip  faulting  in  the  inner 


39 


zone  of  the  Borderland,  formed  new  inshore  basins,  and  realigned  drainage  systems,  form- 
ing centers  of  deposition  for  Pleistocene  turbidity  currents.  This  latest  movement  continues 
today,  as  shown  by  the  modern  seismic  activity  of  the  inner  zone,  but  perhaps  with  abated 
intensity,  inasmuch  as  deformation  of  Pleistocene  sediment  in  the  inner  basis  is  uncommon. 
Wiegand  ( 1970)  suggested  that  the  Newport-Inglewood  fault  has  an  offshore  extension 
of  major  proportion.  Although  there  are  faults  observable  offshore  as  far  south  as  Encinitas 
that  can  be  extrapolated  northward  in  such  a  way  as  to  merge  with  the  Newport-Inglewood 
trend,  there  is  insufficient  evidence  that  these  are  continuous  or  that  they  involved  offsets  of 
the  same  order  of  magnitude  as  the  major  faults  farther  offshore. 


■ 

1024+    'l^ 

1 

256-1024 

1 

64-256 

16-64 

4-16 

1-4 

'            V.^l. 

V            i 

•m^ 


^ 

% 


Figure  6.  Strain  release  in  the  United  States,  1900  to  1965,  expressed  as  the  equivalent  number  of  magnitude  4 
earthquakes/ 10,000  km766  years  (from  Algermissen,  1969). 

LOCAL  SEISMOTECTONICS 

Strain  /-e/ea^e— Earthquake  epicenters  recorded  in  the  San  Diego  area  are  plotted  on  Figure 
8.  The  data  (Fig.  6)  indicate  that  the  energy  associated  with  strain  release  in  the  San  Diego 
area  during  the  past  66  years  should  be  equivalent  to  that  produced  by  approximately  100 
magnitude  4  earthquakes  per  1,000  km-  of  surface  area.  Figure  9  gives  a  generalized  rela- 
tionship between  the  Richter  Magnitude  and  Modified-Mercalli  Intensity.  Within  a  1,000 
km-  circle  centered  on  downtown  San  Diego,  only  12  earthquakes  have  been  reported  dur- 
ing the  past  66  years.  The  magnitudes  of  these  average  less  than  4  and  their  intensities  aver- 
age about  IV.  We  conclude  that  in  the  recent  past  most  of  the  regional  strain  energy  has 
been  released  outside  the  immediate  San  Diego  area.  Algermissen  (1969)  gives  the  average 
strain  release  per  1,000  km-  for  the  entire  Pacific  Coast  (west  of  1 14°  W  longitude)  as  being 
equivalent  to  12  magnitude  4  earthquakes  per  66  year  interval. 

Surficial  faulring.-The  principal  surficial  faults  in  the  San  Diego  area  are  shown  on 
Figure  10.  Many  of  these  appear  to  be  associated  with  past  tectonic  forces,  but  some  of  the 
northwest  trending  and  north  trending  faults  offset  Holocene  (less  than  1 1 ,000  years  old),  as 


40 


30 

I        I L 


NAUTICAL    MILES 

-  VERIFIED    LENGTH    OF    F 

-  DEPTH    IN     FATHOMS 


Figure  7.  Structure  of  the  Continental  Borderland  and  adjacent  regions  (modified  from  Moore.  1969). 

well  as  late  Pleistocene  (less  than  400,000  years  old)  sediments.  The  north  trending  parallel 
series  of  faults  on  Point  Loma,  as  well  as  faults  in  the  Rose  Canyon  area,  offset  Pleistocene 
sediments.  These  faults  can  typically  be  mapped  for  at  least  a  few  miles  (see  Buffington, 
1964;  Kennedy,  1969).  There  is  a  series  of  normal  faults  east  of  San  Diego  Bay  that  offset 
Pleistocene  sediments.  One  of  these,  the  La  Nacion  fault,  offsets  Pliocene  sediments  by  over 
230  ft.  Pleistocene  sediments  by  over  200  ft,  and  Holocene  materials  a  few  feet. 

Faulting  in  basement  rock.—\r\  areas  covered  by  a  veneer  of  sedimentary  rock,  changes 
in  surface  topography  caused  by  faulting  quickly  become  obscured  by  erosion  or  by  depo- 
sition. For  such  areas,  faulting  is  best  delineated  by  changes  in  elevation  of  basement  rock. 
In  the  parts  of  San  Diego  where  batholithic  rocks  are  absent,  basement  is  represented  by 
dense  metavolcanic  rocks  of  Jurassic  age.  Gravity  data  provide  an  excellent  basis  tor  pre- 
dicting the  elevation  of  this  basement  unit  due  to  its  high  density  relative  to  the  overlying 
sediments  (Elliott,  1970). 

Location  of  probable  basement  faulting  by  analysis  of  gravity  data  is  limited,  since 
steep  slopes  on  the  ba.sement  surface  produce  gravity  anomalies  which  differ  only  slightly 
from  those  produced  by  near-vertical  faulting.  The  inferred  regional  stress  pattern  dis- 
cussed earlier  strongly  suggests  that  primary  basement  faulting  is  responsible  for  the  north- 
west trends  evident  in  the  gravity  data  in  the  San  Diego  Bay,  Mission  Bay,  and  Mount  Sole- 
dad  area. 

Figure  1 1  shows  probable  basement  faulting  superimposed  on  the  gravity  data  re- 
cently published  by  Elliott.  The  two  primary  and  presumably  right-lateral  faults,  indicated 


4i 


117"30' 

I     2  -54 

IV     M.M. 

_qJ     J7-H4I 

33*00' 


-J 


32*  30' 

I117-30' 


|117*00' 


Ovil    M.M 


3  3' 00' 


IV    MM 


N 


S- 13     ttOl 


117'00' 


32    30 


Figure  8.  Epicenter  map  of  the  San  Diego  area  giving  date  (e.g.  11-4-25),  Modified-Mercalli  Intensity  (e.g.  IV 
M.M.),  and  Richter  Magnitude  (e.g.  3.4)  when  known.  Circle  shown  has  area  of  1,000  square  kilometers. 

by  the  dashed  Unes,  form  the  edges  of  a  zone  which  narrows  to  the  northwest.  Within  this 
zone,  secondary  fauUing,  indicated  by  the  dotted  hnes,  may  have  occurred. 

The  right-lateral  sense  of  the  primary  faults  is  suggested  by  the  manner  in  which  they 
offset  the  La  JoUa  Submarine  Canyon  (Buffmgton,  1964).  The  westernmost  primary  fault 
has  some  surface  expression  which  has  been  mapped  by  Milow  (see  Buffmgton,  1964)  and 
by  Kennedy  (1969).  The  right-lateral  sense  of  this  fault  may  be  further  indicated  by  offset 
topography  along  the  eastern  margin  of  Point  Loma.  Other  geophysical  data  suggest  that 
this  fault  continues  to  the  south,  where  it  parallels  the  western  coast  of  Baja  California 
(Allen  et  al.,  1965).  The  easternmost  of  the  primary  faults  has  little  surface  expression.  Rose 
Canyon  notwithstanding.  It  can  possibly  be  traced  for  a  limited  distance  in  the  area  north- 
west of  Rose  Canyon,  and  it  may  be  correlated  with  faulting  exposed  east  of  Mission  Bay. 
That  motion  along  the  primary  fault  set  is  not  exclusively  right-lateral  is  attested  to  by  the 
pronounced  gravity  low  in  the  region  of  San  Diego  Bay.  This  view  is  corroborated  by  the 
depths  of  basement  rock  in  the  area  (Elliott,  1964).  Vertical  offset  of  basement  rock  in  excess 
of  a  thousand  feet  is  certainly  consistent  with  geophysical  and  well  data  for  the  South  Bay 
area. 

Due  to  the  convergence  of  the  primary  fault  set,  a  zone  of  transitional  stress  occurs 
north  of  San  Diego  Bay.  Compressional  stress  to  the  north  may  account  for  the  anomalous 
elevation  of  Mount  Soledad.  It  has  been  suggested  that  Mount  Soledad  represents  the  up- 
tilted  edge  of  a  block  which  contains  Mission  Bay  as  the  downtilted  counterpart  (Peterson. 
1970).  The  secondary  faults,  indicated  by  dotted  lines  on  Figure  II,  allow  for  this  possi- 
bility. The  northernmost  secondary  fault  is  provided  in  order  to  allow  for  the  possibility  that 
Mount  Soledad  and  Mission  Bay  represent  a  horst-graben  set.  These  secondary  faults  are 
consistent  with  the  gravity  data  and  are  placed  along  the  strike  of  "positive"  faults,  as 
mapped  by  Milow  (see  Buffington.  1964).  These  "positive"  faults  have  a  direction  of  dip  or 
relative  vertical  separation  which  is  consistent  with  the  gravity  interpretation  presented 
herein. 


42 


Wiegand  (1970)  analyzed  selected  geologic  and  geophysical  data  covering  the  area  be- 
tween the  two  suggested  primary  basement  faults  shown  on  Figure  11.  He  concluded  that 
primary  faulting  has  occurred  along  the  axis  of  San  Diego  Bay  and  proposes  this  fault  may 
be  an  extension  of  the  Newport-Inglewood  fault  zone  to  the  north  and  the  San  Miguel  fault 
to  the  south.  The  gravity  data  of  Figure  1 1  do  not  support  such  faulting  through  San  Diego 
Bay. 


Not  felt  except  by  very  lew  under  especially  favoruble  conditions 


Felt  only  by  a  few  persons  at  rest,  especially  on  upper  Hoors  of  buildings.  Delicately  suspended  objects  may 
swing. 


Felt  quite  noticeably  indoors,  especially  on  upper  floors  of  buildings,  but  many  people  do  not  recognize  it  as  an 
earthquake.   Standing  motor  cars  may  rt)ck  alightly.   Vibration  like  passing  of  truck.   Duration  estimated. 


During  the  day  felt  indoors  by  many,  iiuldoors  by  few.  At  night  some  awakened.   Dishes,  windows,  doors  dis- 
turbed; walls  make  creaking  sound.   Sensation  like  heavy  truck  striking  building.  Standing  motor  cars  rock 
noticeably. 


Fell  by  nearly  everyone;  many  awakened.  Some  dishes,  windows,  etc.,  broken;  a  few  instances  of  cracked  plaster; 
unstable  objects  overturned.   Disturbance  of  trees,  poles  and  other  tall  objects  sometime  noticed.  Pendulum 
clocks  may  stop. 


Ul 

(1 

< 

z 

*A 

K 

z 

UJ 

n 

I.n— 

*- 

u 

ae 

u/ 

Ui 

UJ 

U 

O 

<J 

3 

< 

H- 

O 

Z 

z 

0 

< 

0 

VI 


Felt  by  all;  many  frightened  and  run  outdoors.  Some  ] 
or  damaged  chimneys.   Damage  slight. 


;avy  furniture  moved;  a  few  instances  of  fallen  plaster 


Vli 


Everybody  runs  outdoors.   Damage  negligible  in  buildings  of  good  design  and  construction;  slight  to  moderate  in 
well-built  ordinary  structures;  considerable  in  poorly  built  or  badly  designed  structures;  some  chimneys  broken 
noticed  by  persons  driving  motor  cars. 


Vli 


Damage  sliglit  in  specially  designed  structures;  considerable  in  ordinary  substantial  buildings  with  partial  collapse; 
great  in  poorly  built  structures.  Panel  walls  thrown  out  of  frame  structures.   Fall  of  chimneys,  factory  stacks, 
columns,  monuments,  walls.   Heavy  furniture  overturned.   Sand  and  mud  ejected  in  small  amounts.  Changes  in 
well  water.  Persons  driving  motor  cars  disturbed. 


Damage  considerable  in  specially  designed  structures;  well  designed  frame  structures  thrown  out  of  plumb;  great 
in  substantial  buildings,  with  partial  collapse.  Buildings  shifted  off  foundations.  Ground  cracked  conspicuously. 
Underground  pipes  broken. 


Some  well-built  wooden  structures  destroyed;  most  masonry  and  frame  structures  destroyed  with  foundations, 
ground  badly  cracked.    Rails  bent.    Landslides  considerable  from  river  banks  and  steep  slopes.   Shifted  sand  and 
mud.   Water  splashed  (slopped)  over  banks. 


J 


Figure  9.  Modified  Mercalli  Intensity  scale  showing  approximate  relationship  with  ground  acceleration  and  mag- 
nitude of  shallow  local  earthquakes  (from  Linehan  1970). 


43 


SEISMIC  RISK 

The  determination  of  seismic  risk  is  fraught  with  uncertainties.  To  quantify  these  un- 
certainties is  beyond  the  scope  of  this  paper,  but  we  have  attempted  to  estimate  the  max- 
imum probable  risk  presented  to  the  San  Diego  area  by  earthquake  energy.  An  in- 
troduction to  the  probabiHstic  approach  has  been  recently  presented  by  Esteva  (1970).  Dr. 
Clarence  Allen  (1964)  in  a  discussion  of  the  engineering  implications  of  seismic  geology 

made  the  following  comments: 

"Seismic  zoning  maps  for  engineering  purposes  have  usually  been  constructed  on  the  basis  of  the 
earthquake  history  of  a  region,  sometimes  in  combination  with  the  locations  of  so-called  "active"  faults  and 
related  seismo-tectonic  features.  Indeed,  these  are  normally  the  only  items  of  pertinent  information  avail- 
able—however inadequate.  It  should  be  emphasized,  however,  that  these  data  may  be  even  far  more  in- 
adequate than  most  people  realize.  The  difficulties  and  dangers  in  interpreting  a  relatively  short  recorded 
earthquake  historv,  as  well  as  the  problems  in  attempting  to  differentiate  between  active  and  inactive  faults, 
have  already  been  pointed  out,  together  with  the  very  widespread  distribution  of  earthquake-induced  ef- 
fects during  a  great  shock.  In  addition,  major  after-shocks  of  a  great  earthquake  are  distributed  over  a  far 
wider  area  than  has  generally  been  appreciated,  and  they  constitute  a  hazard  that  may  seemingly  be  quite 
unrelated  to  the  local  fault  pattern.  Potentially  damaging  after-shocks  of  the  1960  Chilean  earthquake,  for 
example,  blanketed  an  area  almost  the  size  of  California.  It  is  significant  that  those  countries  with  the  long- 
est and  most  complete  recorded  earthquake  histories  are  generally  those  in  which  the  mapped  zones  of 
potential  high  seismic  hazard  are  the  broadest,  and  this  lesson  should  be  kept  in  mind  by  those  persons 
attempting  to  construct  new  zoning  maps  or  by  engineers  who  are  facing  the  same  problems  in  regard  to 
specific  sites." 

These  comments  apply  to  much  of  what  follows. 


FORMATIONS 


SAN    DIEGO 


OTAY (NEW) 


SWEE  T  WATER 
(NEW) 


POWAY 


□ 


3r      LA     JOIL A 
O 


ROSARIO 


o-    metamorphic  JJ; 

^      AND  GRANITIC    - 
ROCKS  " 


INFERRED 

CONCEALED 

ANTICLINE 

FAULT 

SYNCLI  NE 

CONTACT 


Figure  10.  Principal  surficial  faults  and  related  structures  (from  Artim  and  Pinckney,  m  press). 


DEFINITIONS 

It  is  of  Utmost  importance  to  define  the  terminology  used  in  ascertaining  seismic  risk. 

Maximum  credible  earl hquake.— This  is  the  maximum  earthquake  that  in  our  judg- 
ment appears  capable  of  occurring.  It  is  the  maximum  rational  and  believable  event  con- 
sistent with  the  known  facts.  While  it  is  highly  unlikely,  it  is  still  a  believable  event  that 
could  occur  within  the  present  geologic  framework  and  present  geologic  epoch.  No  state- 


44 


O    Dry  Oil   Well  35 

(-2138)     Top   of    Basement 


0(-5236)     \         -15 


Figure  11.  Probable  basement  faulting,  Bouguer  gravity  map  after  Elliott  (1970). 
Faulting; Secondary  Basement  Faulting. 


Primary  Basement 


ment  can  be  made  with  regard  to  its  probability  of  occurrence,  other  than  that  it  is  finite 
(modified  from  Cluflfet  al.,  1969). 

Maximum  probable  earthquake.— This  is  the  maximum  earthquake  that  might  occur 
with  a  fairly  high  probability.  The  tectonic  forces  which  cause  it  are  reasonably  well  under- 
stood. Statistical  data  allow  the  prediction  of  a  recurrence  interval  for  this  earthquake.  For 
all  but  the  most  critical  considerations,  it  is  the  maximum  "design"  earthquake. 

Active  fault.— An  active  fault  is  one  that  has  moved  in  historic  time  or  along  which  off'- 
set  of  Holocene  materials  can  be  demonstrated.  If  Holocene  materials  are  not  offset,  but 
numerous  epicenters  have  been  recorded  in  or  in  close  proximity  to  the  fault,  a  classifica- 
tion of  active  may  be  used. 

Potentially  active  fault.— A  potentially  active  fault  is  one  that  offsets  Pleistocene  ma- 
terials, but  for  which  offset  of  Holocene  materials  is  lacking  and  for  which  seismic  activity 
is  nominal  or  absent. 

REGIONAL  RISK 

Algermissen's  regional  risk  map  is  reproduced  in  Figure  12.  On  this  map  San  Diego  is 
shown  in  the  zone  where  "major  destructive  earthquakes  may  occur."  This  map  is  based  on 
the  following:  the  distribution  of  M.M.  (Modified-Mercalli)  intensities  associated  with  the 
known  seismic  history  of  the  United  States;  strain  release  in  the  United  States  since  1900; 
and  the  association  of  strain  release  patterns  with  large  scale  geologic  features  believed  to 
be  related  to  recent  seismic  activity.  Since  this  map  is  based  partly  on  maximum  observed 
intensities,  it  is  biased  towards  conditions  expected  on  worse  ground.  The  probable  fre- 
quency of  occurrence  of  damaging  earthquakes  in  each  zone  was  not  considered  in  assign- 
ing ratings  to  the  zones. 


45 


Figure  12.  Seismic  Risk  (after  Algermissen,  1969).  Zone  0-No  damage.  Zone  1-Minor  damage;  distant  earth- 
quakes may  cause  damage  to  structures  with  fundamental  periods  greater  than  1.0  seconds,  corresponds  to  in- 
tensities V  and  VI  of  the  M.M.  Scale.  Zone  2-Moderate  damage;  corresponds  to  intensity  VII  of  the  M.M.  Scale. 
Zone  3— Major  damage;  corresponds  to  intensity  VIII  and  higher  of  the  M.M.  Scale. 


LOCAL  RISK 

Zone  3,  which  includes  the  San  Diego  area,  corresponds  to  intensity  VIII  and  higher  on 
the  M.M.  scale.  Assuming  that  San  Diego  falls  in  the  intensity  VIII  portion  of  the  zone  leads 
to  the  conclusion  that  it  will  experience  an  acceleration  on  worse  ground  of  approximately 
0.2  g  (Fig.  9). 

An  acceleration  of  0.2  g  exceeds  by  only  a  factor  of  four  the  acceleration  which  San 
Diego  in  all  probability  repeatedly  experiences;  the  isoseismal  maps  shown  in  Figure  13 
indicate  probable  ground  accelerations  in  San  Diego  of  0.05  g.  Events  of  this  size  (i.e.  mag- 
nitude 6.3)  originating  in  the  same  general  10,000  km-  area  as  the  earthquakes  plotted  on 
Figure  13  can  be  expected  to  occur  approximately  once  every  15  years.  Note  that  if  for  the 
earthquakes  shown  on  Figure  13  one  calculates  the  expected  acceleration  directly  from 
magnitude  and  distance  considerations  using  the  most  recent  empirical  relationships,  con- 
siderably lower  probable  acceleration  is  obtained  (Esteva,  1970;  Seed,  Idriss,  and  Kiefer, 
1969).  This  deviation  may  be  explained  in  terms  of  the  shallow  depth  at  which  events  in 
Southern  Cahfornia  occur  and  local  geology. 

Seismic-input  estimates. -The  most  difficult  aspect  of  determining  seismic  risk  is  esti- 
mating the  maximum  energy  which  can  be  expected.  Correlations  between  geological  and 
seismological  data  must  be  used  where  statistically  significant  seismic  data  are  lacking.  Fig- 
ure 14  shows  an  idealized  relationship  between  the  length  of  the  surface  fault  breaks  occur- 
ring at  the  time  of  earthquake  (determined  geologically)  and  earthquake  magnitude  (deter- 
mined seismologically).  The  portion  of  the  curve  above  magnitude  6.5  is  based  on  a  study  of 
historic  surface  faulting  in  the  continental  United  States  and  adjacent  parts  of  Mexico  car- 
ried out  by  Bonilla  (1967).  We  have  plotted  on  this  figure  the  reported  surface  breaks  and 
measured  magnitudes  of  some  important  earthquakes.  The  size  of  the  earthquakes  plotted 
range  from  the  great,  magnitude  8.6,  Chilean  earthquake  of  1960  where  surface  breakage 
occurred  over  a  distance  of  600  miles  to  the  recent  6.6  magnitude  San  Fernando  earthquake 
which  had  surface  breakage  of  over  9  miles.  This  plot  confirms  that  the  stress  drop  across 
the  slipped  fault  is  the  same  for  all  large  earthquakes;  and  therefore,  that  for  a  constant 


46 


Morch    19,  I9S4 

limilt   of  F«ll   Area 

CALIFORNIA 


ARIZONA 


J<f- 


Aprll    t,  196t  ,  It:  2t:  5(9    P.S.T 
Mognitvd*      6.5 
CAIIFORNIA 


-3^ 


Figure  13.  Borrego  area  earthquakes  (Murphy  and  Cloud,  1956:  Von  Hake  and  Cloud,  1966).  Stations  reporting 
anomalously  high  intensity  are  indicated  by  black  dots. 

depth  of  faulting  the  total  strain  energy  released  is  proportional  to  the  length  of  surface 
breakage. 

Maximum  probable  input  from  San  Andreas  system.— The  work  of  Bonilla  ( 1967)  and 
Housner  (1969)  allows  us  to  predict  maximum  probable  earthquake  magnitude  for  a  given 
fault  system  by  assuming  a  maximum  probable  length  of  surface  breakage.  For  example,  if 
one  assumes  for  the  Elsinore  fault  zone  that  surface  breakage  could  occur  along  a  zone  50  to 
70  miles  long,  then  the  resultant  magnitude  would  be  7.3.  Such  a  surface  break  might  be 
expected  to  have  its  southern  limit  near  Vallecito  Valley  and  its  northern  limit  somewhere 
between  Temecula  and  Lake  Elsinore.  Similarly,  if  one  assumes  for  the  San  Jacinto  fault 
zone  that  surface  breakage  could  occur  along  a  zone  150  to  190  miles  long,  then  the  result- 
ant magnitude  would  be  7.8.  Such  a  surface  break  might  be  expected  to  have  its  northern 
limit  between  Riverside  and  San  Bernardino  and  its  southern  limit  between  Imperial  and 
the  known  southern  limit  of  the  Imperial  fault  in  Mexico.  The  above  discussed  approx- 
imations are  plotted  on  Figure  14  and  shown  in  map  view  on  Figure  15. 


47 


Figure  14.  Idealized  relation  between  length  of 
surface  breakage  and  magnitude  of  earthquake 
used  for  determination  of  maximum  probable 
events  on  Elsinore  and  San  Jacinto  Faults. 
C  =  Chile  (1960),  A  =  Alaska  (1964),  S  =  San 
Francisco  (1906),  E  =  E1  Centre  (1940), 
S.M.  =  San  Miguel  ( 1956),  S.Fer.  =  San  Fernando 
(1971)  (Modified  from  Housner,  1969). 


1000 
500 


100 
50 

10 


I 


1.0 


0.5 


0.1 


1 

1 

T 

T 

1 

r 

/•c- 

- 

1  =  2.25X10'* 

e^M 

--^. 

/a  . 

- 

SAN       JACINTO 

/ 

I 

/ 

, 

-        ELSINORE 

/ 

/ 

- 

EV 
/SM 

- 

'- 

IP 
.*S.FER. 
/l 

; 

1-- 

=  1.82X10''e'*^ 

/ 

- 

1 

/ 

/ 

-_ 

1 

1 

1 

1 

2  3  4  5  6  7 

Magn  itude 


8  9 


•a^ 


SAN   DIEGO 


20 

1 


SCALE    OF     MILES 


Eorthquoke     Epicenteri 

a     6   0-69 

A    55-59 

A    5.0-54 

•     4.5-4.9 

•  (1931-1968) 


Figure  15.  Ma.ximum  probably  surface  breakage-Epicenters  and  faults  from  California  Dept.  Water  Res.  Bull. 
1 16-2.  Cross-hatched  zone  =  10,000  km'. 


48 


There  is  presently  considerable  debate  as  to  the  nature  of  the  San  Andreas  fault  in  the 
area  immediately  south  of  the  Salton  Sea  (see  Fig.  16).  Because  the  position  of  faults  in  that 
portion  of  the  Imperial  Valley  is  uncertain,  estimates  of  maximum  probable  surface  break- 
age along  this  part  of  the  San  Andreas  fault  are  not  presently  feasible. 


Figure  16.  Faulting  south  of  Salton  Sea  (from  Elders  et  al.,  1970). 

Figure  17.  which  relates  ground  acceleration  to  epicentral  distance,  indicates  that  a 
magnitude  7.3  event  on  the  Elsinore  fault  produces  greater  ground  acceleration  in  San 
Diego  than  a  magnitude  7.8  event  on  the  San  Jacinto  fault.  However,  the  duration  of  shak- 
ing for  the  magnitude  7.8  event  can  be  expected  to  be  20  per  cent  greater  than  that  pro- 
duced by  the  magnitude  7.3  event  (Steinburgge,  1966).  These  relative  comparisons  are 
valid,  but  the  value  of  ground  acceleration  for  a  given  site  can  range  widely  from  the  "inter- 
mediate ground"  values  given  in  Figure  17.  An  order  of  magnitude  variation  in  acceleration 
is  feasible  (Esteva,  1970).  At  San  Diego,  the  slightly  longer  duration  of  expected  shaking 
produced  by  the  larger  San  Jacinto  fault  event  does  not  result  in  total  dehvered  energy 
greater  than  that  produced  by  the  smaller  event  on  the  Elsinore  fault. 

The  frequency  of  earthquakes  in  Imperial  Valley  has  been  graphed  by  Evernden 
(1970)  in  a  form  selected  to  reflect  the  repeat  interval  for  earthquakes  in  a  10,000  km'  area 
(Fig.  18).  For  comparative  purposes  Figure  15  shows  a  10,000  km-  zone  35  miles  in  width 
which  extends  southward  from  the  town  of  San  Jacinto  to  an  area  just  west  of  the  Laguna 
Salada  in  Baja  California.  From  1934-1971  this  zone  experienced  five  earthquakes  having 
magnitudes  greater  than  6.  This  rate  is  consistent  with  the  data  of  Figure  18.  which  can 
therefore  be  used  to  estimate  repeat  intervals  for  maximum  probable  events  on  the  Elsinore 
and  San  Jacinto  faults.  Linear  extrapolation  to  high  magnitude,  justified  below  magnitude 
8,  yields  a  repeat  interval  of  approximately  60  years  for  the  7.3  event  on  the  Elsinore  and 
approximately  170  years  for  the  7.8  event  on  the  San  Jacinto. 

Note  that  if  the  surface  break  on  the  Elsinore  fault  shown  on  Figure  15  is  logical,  and  if 
the  seismicity  of  the  Imperial  Valley  can  be  applied  to  this  area,  then  the  Elsinore  fault  rep- 
resents the  source  of  maximum  risk  to  the  San  Diego  area.  The  maximum  probable  event 
for  this  fault  occurs  more  often  and  delivers  more  energy  to  the  San  Diego  area  than  does 


49 


)0 


a 

i    6 

o 


Figure  17.  Acceleration  on  intermediate 
ground  as  a  function  of  epicentral  distance 
and  magnitude  (after  Esteva  et  al.,  1964). 


9=  981   cmAec^ 

R=    Epicentrol   Distance 

in   Miles 

/ 

^ 

/ 

7  8    ON     SAN    JACINTO 

/ 

/ 

^ / 

/ 

/ 

/' 

/ 

'  / 

/ 

.<k 

/ 

/ 

/ 

73    ON    EISINORE/ 

A 

'A 

'\ 

^-] 

^/  ^ 

^ 

y 

/         oX 

/^ 

/ 

K 

/ 

^)^ 

/ 

y\ 

o 

1 

0    20 
0  15 


0   10 


0  05 


0  01 


z 
o 


< 

Q 

Z 

o 


10 


20  30  40  60  80        100 

EPICENTRAL     DISTANCE    R,   MILES 


the  maximum  probable  event  for  the  San  Jacinto  fauU.  The  geologically  short  seismic  his- 
tory for  this  fault  does  not,  so  far,  corroborate  this  view.  The  San  Jacinto  fault  poses  an 
equivalent  threat  in  cases  where  the  induced  failure  is  extremely  sensitive  to  the  duration  of 
shaking. 

Maximum  credible  input  from  ojfshore— On  Figure  7,  the  maximum  verified  lengths  of 
offshore  faults  are  plotted.  In  order  to  be  considered  verified,  two  conditions  must  be  met: 
the  fault  must  have  been  positively  identified  on  seismic  profiles;  and  its  extension  between 
profiles  must  be  consistent  with  submarine  topography. 

The  largest  fault  within  the  California  Borderland  is  the  San  Clemente  Island  fault, 
with  a  verified  length  of  approximately  110  statute  miles.  The  maximum  credible  earth- 
quake for  the  Borderland,  produced  by  breakage  of  this  fault  over  its  entire  verified  length, 
would  have  a  magnitude  of  7.7.  Since  this  fault  is  approximately  the  same  distance  from 
San  Diego  as  the  Elsinore  fault,  we  conclude  that  the  maximum  credible  off'shore  event  will 
produce  approximately  50  per  cent  greater  acceleration  in  San  Diego  then  the  maximum 


Figure  18.  Regional  seismicity.  Imperial 
Valley,  California,  1934-1963  (Everden. 
1970). 


MAGNfTUOE 


50 


I        1-6 
t       1.2 

u 
_0 

0)       0.8 
> 

"5 


Q. 

1/1 


0.4 
0 


1.6 
1.2 

0.8 

0.4 

0 

1.6 
1.2 

0.8 

0.4 

0 

^ 

1.6 
1.2 

0.8 

0.4 

0 

i\ 

fi 

\ 

\ 

S    . 

fi 

\ 

J 

^ 

'^'>_ 

A 

. 

^„.^ 

r- 

/ 

A 

/^ 

/ 

^>« 

t 

0     0.5     1.0      1.5        0     0.5     1.0     1.5 
Period— Sec.  Period— Sec. 


0      0,5     1.0    1.5 
Period  —Sec. 


0      0.5     1.0     1.5 
Period  —  Sec. 


MAX.     ACCN.=  0.05g 


Figure  19.  Computed  response  spectra  for  sand  and  gravel  deposit  (from  Seed,  1969).  Predominate  period  of  bed- 
rock motion  =  0.35  seconds;  equivalent  damping  =  5%. 

probable  event  on  the  Elsinore.  It  would  also  cause  a  greater  duration  of  shaking. 

Tsunami  (seismic  sea  wave)  risk.— ]oy  ( 1968)  discussed  tsunamis  and  their  occurrence 
along  the  San  Diego  County  coast.  He  pointed  out  that  the  relatively  wide  Continental  Bor- 
derland off  the  coast  has  historically  acted  as  an  effective  diffuser  and  reflector  of  the  energy 
which  arrives  from  remotely  generated  tsunamis.  Damage  associated  with  remotely  gener- 
ated tsunamis,  therefore,  will  be  most  likely  confined  to  small  craft  in  the  harbor,  although 
some  waterfront  structures  may  also  be  affected. 

Locally  generated  tsunamis  risk  is  difficult  to  assess.  Tsunami  generation  requires  a  rapid 
dislocation  of  the  sea  surface  or  bed  over  a  very  large  area  (thousands  of  square  miles).  This 
size  dislocation  is  not  likely  to  occur  if  the  source  earthquake  has  magnitude  less  than  6.3 
(lida,  1970).  Most  major  dislocations  of  the  sea  surface  or  bed  are  thought  to  be  associated 
with  fault  movement  of  the  "dip-slip"  type.  This  type  motion  seems  to  have  been  associated 
with  the  5.9  magnitude  earthquake  that  occurred  near  San  Clemente  in  1951.  Generation 
times  for  tsunami  producing  dislocations  can  be  as  large  as  several  minutes.  An  earthquake 
could  therefore  initiate  a  large  submarine  landslide,  which  could  then  become  a  tsunami 
source. 

Seven  per  cent  of  Southern  Cahfornia  earthquakes  have  submarine  epicenters  (Cle- 
ments and  Emery,  1947),  and  yet  only  two  or  three  locally  generated  tsunamis  are  known  to 
have  occurred  off  Southern  California  since  1800,  none  in  the  San  Diego  area.  Joy  (1968) 
points  out  that  if  the  San  Clemente  Island  fault  and  the  Agua  Blanca  fault  in  Mexico  (Fig. 
2)  "actually  constitute  a  single  larger  feature, ...  it  could  represent  a  standing  threat  to  San 
Diego  County."  He  concluded,  however,  that  "it  is  entirely  speculative  to  suggest  at  this 
time  that  any  significant  threat  exists.  Certainly  the  nonoccurrence  of  tsunamis  generated 
nearby,  even  if  for  the  geologically  short  period  of  170  years,  cannot  be  ignored." 

Whalin  et  al.  (1970)  have  modeled  the  wave  run-up  which  would  occur  at  San  Diego  if 
a  tsunami  were  to  be  generated  locally.  The  period  of  the  waves  studied  were  intermediate, 
falling  between  values  associated  with  the  wind-wave  and  the  typical  tsunami-wave 
spectra;  the  longest  studied  was  186  seconds.  Waves  of  this  period  are  generated  near  tsu- 
nami sources,  but  become  attenuated  at  great  distance.  They  concluded  that  "the  narrow, 
low-lying  Silver  Strand,  the  City  of  Coronado,  California,  and  portions  of  North  Island 
were  completely  inundated  for  most  conditions  tested.  Wave  heights  in  the  restricted  har- 
bor entrance  approached  26  feet  .  .  .  bores  were  observed  in  the  smaller  partially  enclosed 
basins,  bays,  and  creeks  surrounding  the  harbor." 

LOCAL  RISK  REGIONALIZATION 

Subdivision  of  earthquake  risk  into  local  regions  depends  on  knowledge  of  local  faults 
and  on  the  response  of  the  surficial  geology  to  shaking. 

Risk  due  to  surficial  and  basement  faulting.— Sur^cml  faults  in  the  San  Diego  area  pre- 


51 


sent  zones  of  increased  risk  due  to  the  fact  that  the  effects  of  shaking  may  be  different  on 
opposite  sides  of  a  fault.  Where  the  fault  zone  itself  is  of  considerable  width,  a  third  zone, 
possibly  having  a  still  different  response,  must  be  considered.  Construction  in  zones  of  such 
variable  response  can  result  in  earthquake-induced  problems,  such  as  differential  settling, 
etc. 

Figure  10  shows  the  faults  in  the  San  Diego  area  which  are  considered  to  be  either  "ac- 
tive" or  "potentially  active."  The  La  Nacion  fault  has  a  verified  length  of  approximately 
15  miles.  As  this  fault  offsets  Holocene  materials  at  least  locally  and  demonstrates  repeated 
movement  of  late  Pleistocene  materials,  it  must  be  considered  an  active  fault.  The  maxi- 
mum credible  event  that  could  be  expected  would  be  of  magnitude  6.8  with  expected 
acceleration  approaching  0.4  g. 

Determining  the  maximum  credible  surface  breakage  for  basement  faults  deduced 
from  gravity  data  is  not  a  reliable  method  of  estimating  maximum  credible  earthquake 
magnitudes.  The  approximate  locations  and  sense  of  movement  of  these  faults  (Fig.  1 1)  do, 
however,  provide  clues  to  the  thickness  of  the  less  competent  overlying  sediment,  to  the  lo- 
cation of  zones  along  which  aftershocks  remote  to  large  events  are  apt  to  occur,  and  to  the 
location  of  zones  along  which  local  earthquakes  are  more  apt  to  be  centered. 

Response  of  surficial  materials. —Seed  (1969)  stated  that  "analysis  of  the  effects  of  soil 
conditions  on  damage  due  primarily  to  the  effects  of  ground  shaking  requires  an  under- 
standing of  the  complex  interrelationships  between  the  effects  of  soil  types,  soil  depth,  the 


a 
o 


SURFACE      OUTPUT 


0 

Hydraulic     Fill 
Avg.    BC=  21  (0=75%) 

9 

^  Saturated     Hydraulic    Fill 
Avg.    BC=  21  (D,=  6S%) 
"2=35    /j=95    /,  =  120 

13 
20 

Boy     D*po>ilt 
Avg,      BC  =  S    (0,=  40%) 
Kj  =  30    ^-^=82    J',  =  II3 

3t 

T«r  r  ac«     O«potit  t 
Avg.        BC=24   (  D,=  80%) 
K2=65     J'd  =  102     /,  =  I26 

San    Di*go     Formation 
Avg.      BC  =  59(  D,=  90IOO%J 
'<2="     (|'d='05    ^".=  128 

<fd~  "'V    D«n«'«V 

^,=  Wot    D«n»ily 

D,  =   Rolaliv* 
D«n«ity 

1 
^ 

'. 

BC  =   Blow    Count 
G=    Kj(0)V2 

300 

fT  -    Ov«rburd«n 
Strsst 

BEDROCK    IMPUT 


Figure  20.  Input  acceleration,  surficial  geology,  and  resulting  surface  acceleration  output  (Equivalent  damping 

7.8%). 


52 


MISSION    BAT 


1857    Boundoiy 
Preieni     Boundo 


SAN     DIEGO 


CHUIA     VISTA 


Figure  21.  Areas  which  have  been  filled  since  1857.  (Sources:  U.S.G.S.  Water  Supply  Paper  446;  San  Diego  Bay, 
1859  Coast  Survey  Office;  San  Diego  Union,  June  29,  1971). 

amplitude  of  ground  motions,  the  frequency  characteristics  of  ground  motions,  and  the 
structural  characteristics  of  buildings  in  order  to  analyze  damage  resulting  from  past  earth- 
quakes or  prevent  damage  in  future  earthquakes." 

From  the  standpoint  of  local  zoning,  a  mappable  parameter  which  reflects  the  max- 
imum expected  shaking  is  desired.  One  approach,  illustrated  in  Figure  19,  is  to  calculate  the 
maximum  probable  shaking  at  the  bedrock-soil  interface  and  compute  the  theoretical  re- 
sponse expected  at  the  surface  of  the  ground  (see  Seed,  1969).  From  these  data  one  can 
compile  a  series  of  maps  which  allow  rough  reconstruction  of  the  maximum  expected  shak- 
ing for  any  particular  period  of  shaking.  Housner  (1952)  suggested  that  the  area  under  the 
particle  velocity  spectrum  over  a  range  of  shaking  rates  be  used  to  define  the  local  intensity 
of  shaking.  This  measure  of  intensity  would  be  obtainable  directly  from  such  maps. 

To  calculate  the  theoretical  response  at  the  ground  surface,  values  of  the  shear  modu- 
lus, unit  weight,  and  damping  factor  are  needed  as  a  function  of  depth  from  the  surface  to 
bedrock.  Care  must  be  taken  in  selecting  these  values  due  to  the  nonlinearity  of  some  of 
these  "constants"  with  increasing  strain.  The  seismic  input  should  be  either  an  average 
smoothed  response  compiled  from  earthquakes  having  magnitude  and  epicentral  distance 
similar  to  the  maximum  probable  event  expected  to  affect  the  San  Diego  area,  or  a  "white" 
input.  An  advantage  of  using  a  "white"  input  is  that  it  allows  separation  of  effects  due  to 
surficial  materials  from  those  due  to  the  transmission  path;  it  also  provides  a  mappable  out- 
put from  which  the  response  for  any  input  is  readily  calculable. 

We  have  carried  out  recently  a  similar  analysis  to  determine  the  response  of  the  sur- 
ficial materials  to  shaking  and  response  of  simple  structures  to  the  resultant  velocity  of  sur- 
face shaking.  Figure  20  shows  the  acceleration  time  history  input  at  the  top  of  the  Cre- 
taceous sediments,  the  assumed  geologic  column  overlying  these  sediments,  and  the  theo- 
retically predicted  acceleration  time  history  which  would  result  at  the  surface.  For  the 
geologic  parameters  and  acceleration  levels  assumed,  the  maximum  acceleration  is  in- 
creased by  more  than  a  factor  of  two  (i.e.  from  0.06  g  to  0.13  g)  in  traversing  the  300  foot 
column.  Most  of  this  amplification  occurs  in  the  upper  28  feet,  which  is  assumed  to  be  com- 
posed of  loosely  consolidated  sediments.  The  stratagraphic  column  assumed  is  typical  of 
much  of  the  area  surrounding  the  bays  to  which  fill  has  been  added  (Fig.  21).  These  filled 
areas  represent  zones  of  maximum  seismic  risk. 

The  spectrum  shown  in  Figure  22  approximates  the  particle  velocity  spectrum  at  the 
earth's  surface  for  the  conditions  and  assumptions  described  on  Figure  20.  If  data  of  this 


53 


1.0 
PERIOD-SECONDS 


Figure  22.  Spectral  velocity  derived  from  surface  acceleration. 

sort  were  available  throughout  areas  of  maximum  expected  seismic  risk,  considerable  prog- 
ress could  be  made  toward  rational  seismic-risk  zoning. 


CONCLUSIONS  AND  RECOMMENDATIONS 

In  this  paper  we  have  consolidated  the  background  information  developed  by  various 
experts  in  the  field  of  geology,  seismology,  and  earthquake  engineering  with  specific  appli- 
cation to  the  San  Diego  area,  and  have  formed  the  following  conclusions  and  recommenda- 
tions. 

1.  San  Diego  is  in  an  active  seismic  area. 

2.  A  Richter  magnitude  7.3  earthquake  on  the  Elsinore  fault  having  a  repeat  interval 
of  60  years  appears  to  be  the  "maximum  probable"  earthquake  for  San  Diego.  For  most 
construction  the  "maximum  probable"  earthquake  is  recommended  for  design.  In  the  case 
of  structures  such  as  hospitals  which  must  remain  operative  during  times  of  disaster  and 
special  installations  such  as  nuclear  reactors  design  should  be  based  on  the  "maximum 
credible"  earthquake. 

3.  Design  studies  should  further  consider  the  possible  effects  of  a  magnitude  7.8  "max- 
imum probable"  earthquake  on  the  San  Jacinto  fault  because  of  its  longer  expected  dura- 
tion of  shaking. 

4.  Structures  built  on  filled  areas  underlain  by  loose  embayment  type  soils  such  as 
those  found  in  San  Diego  and  Mission  bays  are  particularly  susceptible  to  earthquake  dam- 
age. Because  of  the  numerous  structures  planned  for  such  areas,  a  comprehensive  study  of 
the  effects  of  earthquake-induced  forces  in  embayment  deposit  areas  is  strongly  recom- 
mended. 

5.  A  seismic  sea  wave  (tsunami)  initiated  within  the  offshore  California  Continental 
Borderland  is  possible.  Such  a  wave  could  have  a  damaging  eff'ect  on  low-lying  shoreline 
areas  along  the  Pacific  Ocean  and  in  mouths  of  bays. 

ACKNOWLEDGMENTS 

Publication  of  this  paper  has  been  made  possible  in  part  by  financial  support  provided  by  the  San  Diego  Asso- 
ciation of  Geologists.  This  study  was  partially  supported  by  a  research  grant  from  the  Professional  and  Geotechni- 
cal  Development  Program  of  Woodward-Gizienski  &  Associates.  This  support  is  gratefully  acknowledged. 

We  would  like  to  thank  L.  J.  Lee,  S.  P.  Gizienski.  and  R.  G.  Gastil  for  reviewing  this  manuscript.  R.  P.  Phillips 
for  permission  to  publish  his  compilation  of  epicenters,  and  T.  M.  Gavin  for  his  help  with  the  computer  analysis. 


54 


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1969.  Seismic  risk  studies  in  the  United  States,  p.  14-27.  In,  Proceedingsof  fourth  World  conference  on  earth- 
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Allen,  C.  R. 

1964.  Engineering  implications  of  seismic  geology.  In.  Proceedings  of  the  earthquake  and  geologic  hazards 
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Allen,  C.  R.,  P.  St.  Amard.  C.  R.  Richter,  and  J.  M.  Nordquist 

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Allen.  C.  R..  L.  T.  Silver,  and  P.  G.  Stehh 

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Artim.  E.  R.,  and  C.  J.  Pinckney 

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1967.  Historic  surface  faulting  in  continental  United  States  and  adjacent  parts  of  Mexico.  U.S.  Geol.  Sur., 
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Buffington,  E.  C. 

1964.  Structural  control  and  precision  bathymetry  of  La  Jolla  Submarine  Canyon,  California.  Marine  Geol. 
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Clements,  T.,  and  K.  O.  Emery 

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Coffman,  J.  L.,  and  W.  K.  Cloud 

1970.  United  States  earthquakes  1968.  U.S.  National  Earthquake  Information  Center.  Washington.  D.C. 

Cluff.  L.  S. 

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Elders,  W.  A.,  R.  W.  Rex,  T.  Meidav,  and  P.  T.  Robinson 

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Elliott,  W.  J. 

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Elliott,  W.  J. 

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Esteva,  L. 

1970.  Seismic  risk  and  seismic  design  decisions,  p.  142-152.  In.  Seismic  design  nuclear  power  plants.  M.I.T. 
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Esteva.  L..  and  E.  Rosenblueth 

1964.  Espectros  de  temblores  a  distancias  moderadas  y  granelas.  Soc.  Mex.  Ing.  Sis.  Bol.  2(1). 

Evemden.  J.  R. 

1970.  Study  of  regional  seismicity  and  associated  problems.  Seism.  Soc.  Amer.  Bull.  60:  393-446. 

Gastil.  R.  G..  E.  C.  Allison,  and  R.  P.  Phillips 

Reconnaissance  geology  of  the  State  of  Baja  California.  Geol.  Soc.  Amer.  Mem.  (in  press). 

Gutenberg,  B. 

1 94 1 .  Mechanism  of  faulting  in  southern  California  indicated  by  seismograms.  Seism.  Soc.  Amer.  Bull.  3 1 :  263- 
302. 
Gutenberg,  B.,  and  C.  F.  Richter 

1954.  Seismicity  of  the  earth.  Princeton  L'niversity  Press,  Princeton,  2nd  ed.:  310  p. 

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1952.  Intensity  of  ground  motion  during  strong  earthquake.  California  Inst.  Tech.,  Earthquake  Res.  Lab., 
August  1952. 

Housner,  G.  W. 

1969.  Engineering  estimates  of  ground  shaking  and  maximum  earthquake  magnitude,  p.  1-13.  In.  Proceed- 
ings of  fourth  World  conference  on  earthquake  engineering. 


55 


lida,  K. 

1970.  The  generation  of  tsunamis  and  the  local  mechanism  of  earthquakes,  p.  3-18.  In,  Tsunamis  in  the  Paci- 
fic Ocean,  Proceedings  of  the  Int.  Symp.  on  Tsunamis  and  Tsunami  Research.  East-West  Center  Press, 
Honolulu. 

Joy,  J.  W. 

1968.  Tsunamis  and  their  occurrence  along  the  San  Diego  County  coast.  Westinghouse  Ocean  Research  Lab 
report  for  the  Unified  San  Diego  County  Civil  Defense  and  Disaster  Organization.  35  p. 

Kennedy.  M.  P. 

1969.  Preliminary  geologic  map  of  a  portion  of  northwestern  San  Diego  City,  California.  California  Div. 
Mines  Geol.  Open-File  Rept..  Scale  1:9,600. 

Linehan, D. 

1970.  Geological  and  seismological  factors  influencing  the  assessment  of  a  seismic  threat  to  nuclear  reactors, 
p.  69-90.  In.  Seismic  design  for  nuclear  power  plant.  M.I.T.  Press,  Cambridge. 

Moore,  D.  G. 

1969.  Reflection  profiling  studies  of  the  California  Continental  Borderland:  structure  and  Quaternary  turbi- 
dite  basins.  Geol.  Soc.  Amer.  Spec.  Paper  107. 

Peterson,  G.  L. 

1970.  Quaternary  deformation  patterns  of  the  San  Diego  area,  southwestern  California,  p.  120-126.  In.  Amer. 
Assoc.  Geol.  Guidebook  1970  Fall  Field  Trip  of  Pac.  Sec.  of  AAPG.  SEPM.  and  SEG. 

Richter.  C.  F. 

1958.  Elementary  seismology.  W.  H.  Freeman  and  Company,  San  Francisco. 

Richter,  C.  F. 

1959.  Seismic  regionalization.  Seism.  Soc.  Amer.  Bull.  49:  123-162. 

Savage,  J.  C,  and  R.  D.  Burford 

1970.  Accumulation  of  tectonic  strain  in  California.  Seism.  Soc.  Amer.  Bull.  60:  1877-1896. 

Scholz,  D.  H.,  and  T.  J.  Fitch 

1969.  Strain  accumulation  along  the  San  Andreas  fault.  J.  Geophys.  Res.  74:  6649-6666. 

Schwiderski,  E.  W. 

1968.  Mantle  convection  and  crustal  tectonics  inferred  from  a  satellite's  orbit:  a  diflferent  view  of  sea-floor 
spreading.  J.  Geophys.  Res.  73:  2828-2833. 

Seed,  B.  H. 

1969.  The  influence  of  local  soil  condition  on  earthquake  damage.  In.  Soil  dynamics.  Proc.  of  specialty.  Ses- 
sion 2.  Seventh  International  Conference  on  Soil  Mechanics  and  Foundation  Engineering. 

Seed,  B.  H.,  I.  M.  Idreiss,  and  F.  W.  Kiefer 

1969.  Characteristics  of  rock  motions  during  earthquakes.  Soil  Mechanics  and  Foundations  Division.  Amer. 
Soc.  Civil  Eng.  95:  1199-1218. 

Steinbrugge,  C. 

1966.  Engineering  seismicity  aspects  of  Prince  William  Sound,  Alaska  earthquakes,  preliminary  report, 
March-April,  1964.  U.S.  Coast  and  Geodetic  Survey:  58-75. 

Von  Hake.  C.  A.,  and  W.  K.  Cloud 

1966.  United  States  earthquakes  1964.  U.S.  Coast  and  Geodetic  Survey.  Washington,  D.C. 

Whalen,  R.  W..  D.  R.  Bucci,  and  J.  N.  Strange 

1970.  A  model  studv  of  wave  run-up  at  San  Diego.  California,  p.  427-452.  In.  Tsunamis  in  the  Pacific  Ocean, 
Proceedings  of  the  Int.  Symp.  on  Tsunamis  and  Tsunami  Research.  East-West  Center  Press.  Honolulu. 

Wiegand.  J.  W. 

1970.  Evidence  of  a  San  Diego  Bay-Tijuana  fault.  Assoc.  Eng.  Geol.  Bull.  7:  107-121. 


Department  of  Geology,  San  Diego  State  College,  San  Diego,  California,  and  Wood- 
ward-Gizienski  &  Associates,  San  Diego,  California. 


56 


APPENDIX 
Important   Earthquakes,   Modified  and  Up-dated  from  Joy  (1967). 


Type,  Date,  and  Time 
z  indicates  GMT 
*  indicates  PST 


Generating  region  and 
epicenter,  if  known. 
Magnitude  (Arabic) 
Modified  Mercaili 
Intensity  (Roman) 


Remarks  and  References 


Local  earthquake 
April  II,  1769 

Coastal  earthquake 
November  22,  1800, 
1330* 


Local  earthquake 
May  25,  1803 
(no  tsunami  noted) 
Regional  earthquakes 
May,  1812 


Local  earthquakes 
October  12,  1812 
Coastal  earthquake 
December  8,  1812 
about  0700* 


Local  earthquake 
June  23, 1843, 1530* 
Local  earthquake 
September  16,  1849 
Local  earthquake 
September  22,  1849 
Local  earthquake 
April  12,  1852 
Local  earthquakes 
October  26,  1852 
November  27-30,  1852 
(no  tsunami  noted) 
Possible  remote  tsunami 
November  1853 


Remote  tsunami 

December  23,  1854, 

0015Z 

December  24,  1854, 

0800z 

Local  earthquake 

September  20,  1856 

Several  local  earthquakes 
May  27,  June  13-14,  1862 
October  21,  1862 
January  25,  1863,  0200* 
July  7,  1963,  1311* 
Local  earthquake 
April  19,  1865 
Remote  tsunami 
April  2,  1868 


San  Diego  (severe) 


Southern  Cahfornia  (VII) 


San  Diego,  near  32.5°N 
117°W 

Southern  California 


San  Diego 

Coastal  Southern  Cali- 
fornia (VIII-IX) 


San  Diego  (very  severe) 
Baja  California 
Santa  Ysabel 
San  Diego  County 
Carrizo  Creek 
San  Diego  County 
San  Diego 

San  Diego 
(IX-November  29,  1852) 


Kuril  Islands 


Ansei,  Tokkaido,  Japan 
34.1°N;  137.8°E(8.4) 

33.2°N;  135.6°E(8.4) 

San  Diego  County 
(VII) 

All  at  San  Diego 


San  Diego 
S.  E.  Hawaii 


Diary  of  Miguel  Costanso,  Portola  Expedition 
1769-1770.  Bancroft  (Works,  VoL  18,  p.  127); 
Wood  (1916);  Townley  &  Allen  (1939). 
The  adobe  walls  of  San  Diego  Presidio  barracks 
were  cracked.  California  Archives  Provincial  State 
Papers  XXI,  p.  135;  HitteU  (1898);  Wood  &  Heck 
(1966);  Trask  (1864);  Bancroft  (1883)  p.  86, 
654,658. 

Slightly  Damaged  San  Diego  Mission  Church, 
Wood  &  Heck  (1961);  Bancroft  (1883)  p.  106, 
114;  Wood  (1916). 

Southern  California  was  subjected  to  nearly 
continuous  shocks  for  4  Vi   months.  The  inhabi- 
tants abandoned  their  houses  and  lived  out  of 
doors.  Townley  &  Allen  (1939). 
Shocks  for  40  days.  Townley  &  Allen  (1939). 

Mission  San  Juan  Capistrano  destroyed.  Strongly 

felt  at  San  Diego,  but  no  damage  to  San  Diego 

Mission.  Probably  on  Inglewood-Newport  fault. 

Bancroft  (1883)  p.  347-348;  Lounderback  (1948); 

Heizer  (1941)  p.  221;  5.  F.  Bulletin,  March  5, 

1864,  March  19,  1864,  p.  3,  c.  4. 

Southern  California  to  Mexico.  Townley  & 

AUen  (1939);  Wood  &  Heck  (1966). 

Probably  on  the  Elsinore  fault.  Townley  &Allen 

(1939). 

Probably  on  Elsinore  fault.  Townley  &  Allen 

(1939). 

Townley  &  Allen  (1939). 

November  29  two  minute  shock  in  San  Diego 
followed  by  light  quakes  for  several  days.  Trask 
(1856);  Wood,  (1916);  Townley  &  Allen  (1939). 

Reported  to  be  a  large  tsunami.  Small  waves 
possibly  recorded  on  newly  installed  San  Diego 
gage.  Solov'ev  and  Ferchev  (1961);  lida,  et  al., 
(1967);  U.S.  Coast  Survey  Report  for  1855 
(1856)  p.  99. 

A  very  large  tsunami.   Recorded  at  San  Diego 
+  12.6  hrs.  later,  0.5  feet,  31  min.  avg.  period. 
Bache  (1856);  lida,  et  al,  (1967);  Shuck  (1869). 
Two  tsunamis,  whose  effects  along  this  coast 
were  more  or  less  merged  together. 
Walls  cracked,  ceilings  fell,  and  local  Indians  were 
terrified.  Cattle  stampeded  at  Santa  Ysabel. 
Wood  &  Heck  (Rev.  1966). 
May  27,  severe  shock  at  San  Diego,  Temecula, 
Probably  on  Elsinore  fault.  Track  (1864); 
Bancroft,  MMs  cited  by  Townley  &  Allen  (1939). 


Severe  shock.  Townley  &  Allen  (1939). 

Recorded  at  San  Diego,  0.33  ft;  30  min.  avg. 
period.  S.  F.  Bulletin,  June  13,  1868;  lida,  etai, 
(1967);  Heck  (1947);  Townley  &  Allen  (1939). 


57 


Type,  Date,  and  Time 
z  indicates  GMT 
*  indicates  PST 


Generating  region  and 
epicenter,  if  known. 
Magnitude  (Arabic) 
Modified  Mercalii 
Intensity  (Roman) 


Remarks  and  References 


Remote  Tsunami 
August  13,  1868, 
1645Z 


N.  Chile;  So.  Peru 
18.5°S;71°W 


Remote  Tsunami 
August  23,  1872 


Remote  tsunami 
September  16-17,  1872 


Hawaii  (?) 


(?) 


Remote  tsunami 

November  22,  1878 

Probably  either  that  of 

May  10,  1877,  or  August 

13,  1868,  but  misdated. 

(no  local  earthquakes  reported) 

Offshore  or  coastal  earthquake  Los  Angeles  region 

earthquakes  and  possible  (VI-V) 

local  tsunami  August  10, 

1879,  1315* 


Local  earthquakes 

December  21,  1880 

2300* 

Local  earthquakes 

March  11,  1882,  1600* 

March  30,  1882,  2300* 

Octobers,  1882,0200* 

(no  tsunamis) 

Local  inland  earthquake 

February  9,  1890,0406* 

(no  tsunami) 

Offshore  earthquakes 

February  23,  1892,  2320* 

Several  aftershocks 

(no  tsunamis) 

Local  earthquake 

October  23,  1894 

Local  earthquake 

July  3,  1896,2127* 

(no  tsunami) 

Local  inland  earthquake 

December  23,  1899,0425* 

Remote  tsunami 

January  31,  1906,  1536z 


San  Diego 
(V-strongest) 

San  Diego 
(Ill-V) 


San  Jacinto  fault  (VI) 


Off  coast  N.W.  of 
Ensenada,  B.C.,  Mexico 
31.5°N;  116.5°W(VII-1X) 


San  Diego-Poway  region 
33°N;  117°W(V1I) 
San  Diego  (Small) 


San  Jacinto  fault 

(IX) 

Columbia  -  Ecuador 

1°N;  81.5%  (8.6) 


Great  Africa,  Peru,  Tsunami.  Recorded  at  San 
Diego  1 1.9  hrs.  later,  1.0  ft,  16  min.  avg.  period. 
Noted  at  San  Pedro  and  Wilmington,  6.0  ft,  20 
min.  avg.  period.  Proctor's  article  gives  heights 
at  San  Pedro  which  are  10  times  too  great  ac- 
cording to  local  newspaper  accounts.  Alta  Cali- 
fornia, September  12,  1868;Z,./4.  Star,  August 
14-19,  1868;  lida,  era/.,  (1967);  von  Hockstetter 
(1868);  (1869);  Berninghausen  (1962);  Proctor 
(1869). 

lida,  et  at.,  (1967),  lists  a  tsunami  noted  in 
Hawaii  on  this  date.  Davidson  (1872)  gives 
August  24th  as  the  date  that  a  tsunami  was  ob- 
served at  Astoria,  San  Diego,  and  San  Francisco. 
No  details. 

Davidson  (1872)  states  that  tsunami  activity  was 
noted  on  these  two  dates  on  the  San  Diego,  San 
Francisco,  and  Astoria  Ore.  tide  gages.   No  de- 
tails or  confirmation  from  other  sources. 
Angel  (1883)  reports  6  ft.  waves  at  Wilmington. 
Waves  and  damage  done  at  San  Luis  Obispo, 
Point  Sal,  Avila,  Port  Harford,  Surf,  Pismo, 
Morro  Bay,  and  Cayucos.  Not  noticed  in  San 
Diego.   No  reports  from  any  other  source  for 
tsunamis  of  this  date. 

Sea  wave  noted  in  Santa  Monica  Bay.  Not  report- 
ed in  San  Diego.   The  sea  wave  was  originally 
reported  by  Rockwood  who  was  cited  by 
Townley  &  Allen;  no  detail,  small  earthquake. 
No  tsunami  noticed  elsewhere.  lida,  et  ai, 
(1967)  cite  Wood  (1916)  as  authority  for  this 
event  and  accept  it  as  a  valid  tsunami.  Rockwood 
(1879);  Wood  (1916);  Townley  &  AUen  (1939); 
lida,  era/.,  (1967). 
Townley  &  AUen  (1939);  Rockwood  (1881). 


Townley  &  AUen  (1939). 


Felt  in  San  Diego.  Wood  &  Heck  (Rev.  1966). 


Considerable  datnage  in  San  Diego.   Possibly 
Aqua  Blanca  fault.  Wood  &  Heck  (1966); 
Richter  (1965);  Holden  (1898);  Allen,  Silver,  & 
StehU  (1960);  Townley  &  AUen  (1934). 
Walls  cracked,  but  no  serious  damage  in  San 
Diego.  Wood  &  Heck  (1966). 
Townley  &  AUen  (1939). 


Felt  in  San  Diego.  Wood  &  Heck  (Rev.  1966). 

A  major  tsunami;  said  to  have  been  recorded  in 
San  Diego  although  heights  not  given.  Heck 
(1947);  lida,  era/.,  (1967). 


58 


Type,  Date,  and  Time 
z  indicates  GMT 
*  indicates  PST 


Generating  region  and 
epicenter,  if  known. 
Magnitude  (Arabic) 
Modified  Mercalli 
Intensity  (Roman) 


Remarks  and  References 


Local  inland  earthquake 
May  15,  1910,0747* 
(no  tsunami) 
Remote  tsunami 
November  10,  1922 
0433z 

Remote  tsunami 
February  4,  1923 
1602z 

Remote  tsunami 
April  14,  1923, 
1531Z 

Offshore  earthquakes 
and  local  tsunami 
November  4,  1927,  1351z 
0551* 


Remote  Tsunami 
March  3,  1933 
March  2,  1933,  17:31z 
October  2,  1933, 
Aftershock 


Coastal  earthquake 
March  10,  1933  1754* 


Local  inland  earthquake 
December  30-31,  1934 

Large  local  earthquake 
May  18,  1940,  2036* 
(  no  tsunami) 

Possible  remote  tsunami 
February  9,  1941,  0144* 


Lake  Elsinore  (6.0) 


Atacama,  No.  Chile 
25.5°S;  70°W  (8.3) 

Kamchatka 
54°N;  161°E(8.3) 

Kamchatka 
56.5°N;162.5°E(7.2) 

Off  Cape  Arguello 
34.5°N;121.5°W 
(7.3,  IX-X) 


Sanriku,  Japan 
Tuscarora  Deep 
39.l'^N;144.7^E 


Long  Beach,  California 
33.6^N;  118°W(6.25,IX) 


Imperial  Valley  in  Mexico 
32°N;  114.75^W(7.1,IX,  X) 

S.E.  of  El  Centro 
32.7°N;  115.5°W(7.1,X) 


Cape  Mendocino,  California 
40.9°N;125.4°W(6.6) 


Felt  in  San  Diego.  Probably  Elsinore  fault. 
Wood  &  Heck  (1966). 

At  San  Diego  13  hrs.  later,  1.3  ft.  max.,  15  min. 
period.  Willis,  (1929);  Berninghausen  (1963); 
lidai,  etal.,  (1967). 

At  San  Diego  10  hrs.  later,  1.3  ft.  max.,  10  min. 
period,  lida,  etal.,  (1967);  Miller  (1964)  RED 
in  Honolulu. 

At  San  Diego  14.3  hrs.  later,  0.75  ft.  max.,  43 
min.  period,  lida,  etal.,  (1967);  MiUer  (1964) 
RED  in  Honolulu. 

This  is  the  only  well  documented  locally  gener- 
ated tsunami  in  California  history.    Byerly's 
paper  reproduces  marigrams.  This  was  a  very 
small  tsunami;  very  small  at  La  Jolla  and  San 
Diego,  but  was  detected  at  Hilo,  Hawaii  5.1  hrs. 
later  where  it  produced  water  level  excursions 
of  about  8  in.  Along  this  coast  the  max.  heights 
nowhere  exceed  those  attained  by  great  tsunamis 
of  distant  origin.  According  to  Wilson  the  earth- 
quake was  felt  at  sea  by  the  S.S.  Socony  which 
at  the  time  was  at  34°54'34"N;  121°01'00"W. 
La  JoUa  +0.9  hrs.,  0.02  ft.      (6mm) 

Nr.  Port  San  Luis  5  ft. 

Surf-Pismo  6  ft. 

San  Diego  0.02  ft. 

San  Francisco  +1.2  hrs.,  0.02  ft.      (4mm) 

(15  min.  period  at  La  Jolla,  12  min.  period  at 
San  Francisco)  Byerly  (1930);  Wilson  (1928); 
Miller  (1964)  RED;  Wood  &  Heck  (1961);  lida, 
etal.,  (1967);  Cox  (1964). 
Great  Sanriku  Tsunami. 

Santa  Monica  0.35  ft. 

San  Pedro  +11.5  hrs.  0.75  ft. 

La  JoUa  0.25  ft. 

San  Diego  -  probably  recorded  but  very  small. 
(14  min.  period  at  San  Pedro,  1 1  min.  period 
at  La  JoUa).  Neumann  (1935)  Heck  (1947); 
lida,  etal.,  (1967);  Miller  (1964)  BLUE  in 
Hawaiian  Island. 

Destructive  Long  Beach  earthquake.   Newport- 
Inglewood  fault.  Tide  gage  records  ground 
motion  at  Long  Beach.  Nothing  noticed  on 
San  Diego  tide  gage  record.  Slight  activity  no- 
ticed at  La  Jolla;  possible  shelf  seiche.  The 
earthquake  was  felt  aboard  ships.   Aftershock 
felt  in  San  Diego.  Emery  (1960)  p.  125;  Wood 
(1933);  Bittinger  (1933);  Wood  &  Heck  (1961); 
Eaton  (1933);  Clements  &  Emery  (1947); 
Buwalda(1933). 

Strongly  felt  in  Tiajuana  and  San  Diego.   Also 
felt  in  Arizona  and  Nevada.  Wood  &  Heck 
(1966). 

The  surface  rupture  produced  during  this  earth- 
quake was  over  40  miles  long.  Shocks  strongly 
felt  throughout  San  Diego  County  with  some 
damage.  Wood  &  Heck  (1966). 
From  inspection  of  local  tide  records;  not  re- 
ported elsewhere.  San  Diego:   +14  hrs.,  general 
increase  in  harbor  seiching.   La  Jolla:   no  unusual 
activity  noticed.  Port  Hueneme:   +36  hrs., 
general  increase  in  harbor  seiching.  San  Fran- 
cisco:  +  14  hrs..  some  harbor  seichine. 


59 


Type,  Date,  and  Time 
z  indicates  GMT 
*  indicates  PST 


Generating  region  and 
epicenter,  if  known. 
Magnitude  (Arabic) 
Modified  Mercalli 
Intensity  (Roman) 


Remarks  and  References 


Remote  tsunami 
December  7,  1944,  0435z 


Local  earthquake 
January  1,  1946, 

Remote  Tsunami 
April  1,  1946,  1229* 


Kii,  Japan 

33.75^N;  136°E  (8.0) 


32°43'N;117°25'W(3.3) 


Aleutian  Islands 
53.5°N;  163^(7.4) 


Local  inland  earthquake  Anza  Desert 

September  5,  1950,  1120*  33.7'^N;  116.8°W  (4.8) 
(no  tsunami) 

Offshore  earthquake  San  Clemente  Island 

December  25,  1951,  1647*  32.8°N;  118.3°W  (5.9,  VI) 


Remote  tsunami 
March  4,  1952 


Hokkaido,  Japan 
42.2°N;  143.8°E  (8.1) 


Remote  tsunami  Kamchatka 

November  5,  1952,  1658z         57.75°N;  159.5°E  (8.25) 


Local  inland  earthquake 
June  13,  1953,2017* 
Local  earthquake 
March  19,  1954,  0154* 


Local  inland  earthquake 
March  22,  1954,  2015* 
Local  inland  earthquake 
October  17,  1954,  1457* 

Local  inland  earthquake 
October  24,1954,0144* 

Local  inland  earthquake 
November  12,  1954,  0427* 
Two  aftershocks 
Local  earthquakes 
January  3,  1956 
February  9,  1956,  0633* 

Local  earthquakes 
February  14,  1956,  1033* 
1720* 


Imperial  Valley 
32.8°N;  115.7°W(VII) 


Borrego  Springs 
33°17'N;  116^11' 


W(VI) 


Santa  Rosa  Mountains 
33.3°N;  116.2°W(5.1,VI) 
Baja  California 
31.5°N;116.5°W(5.7) 

Baja  California 
31.5°N;116°W(6.0) 

Baja  California 
31.5°N;116°W(6.3) 

Baja  California 

bothat31.8°N;  115.9°W 

1/3/56  (4.7) 

2/9/56  (6.8,  VI-VIII) 

Baja  California 

31.8°N;  115.9°W(V-VIII) 

1033*  =  6.3 

1  T^n*  -   C    A 


San  Diego 
La  Jolla 
Los  Angeles 
Port  Hueneme 
San  Luis  Obispo 
Avila,  Calif. 


San  Diego,  +13.9  hrs.,  0.33  ft.,  14  min.  period. 
Terminal  Island,  Long  Beach,  0.33  ft.,  16  min. 
period.  lida,  etal,  (1967);  Heck  (1947); 
Bodle(1946). 

Locahzed  shock  in  the  coastal  area  running  from 
La  Jolla  through  San  Diego  and  National  City 
"Buildings  swayed." 
A  great  tsunami 

1.3  ft. 
+6.2  hrs.  1.4  ft. 

2.6  ft. 

5.5  ft. 
+5.6  hrs.,  8.0  ft. 

3.8  ft. 

Munk  (1953);  Green  (1946);  Bodle  &  Murphy 
(1948);  lida,  e/fl/.,  (1967);  MiUer  (1964) 
BLUE. 
Felt  in  San  Diego.  Wood  &  Heck  (1966). 


Slight  damage  in  San  Diego  and  northern  parts 
of  the  county.  No  tsunami.  Richter  (1965); 
Wood  &  Heck  (1966). 
"There  is  some  coherence  between  the  two 
records,  with  similar  phases  occurring  at  Ocean- 
side  2  to  3  min.  after  La  Jolla."  (Munk,  1953). 
Los  Angeles,  2.6  ft:   Oceanside,  1.5  in.;  La  Jolla, 
1.0  in.  (10  min.  avg.  period  at  Oceanside  and 
La  JoUa).  Munk  (1953);  Miller  (1964)  RED; 
Murphy  &  Cloud  (1954);  lida,  etal,  (1967). 
A  great  tsunami. 
San  Diego 
La  Jolla 

L.A.  (Berth  174) 
San  Pedro  Bkwtr 
Santa  Monica 
Port  Hueneme 
Avila,  Calif. 

lida,  et  al.,  (1967);  Zerbe  (1953);  MUler  (1964) 
RED. 

Felt  from  San  Diego  to  Phoenix,  Arizona.  Wood 
&  Heck  (1966). 

Intensity  VI  at  La  Jolla.  Felt  by  and  awakened 
many  in  La  Jolla;  frightened  few.  Cracked  plastt 
and  a  few  walls  in  La  Jolla.  Murphy  &  Cloud 
(1956). 

Slight  damage  from  Palm  Springs  to  San  Diego, 
Wood  &  Heck  (1966). 
Felt  in  San  Diego  County,  Probably  Agua 
Blanca  fault.  Murphy  and  Cloud  (1956)  Richter 
(1958). 

Felt  in  San  Diego  County.  Probably  Agua 
Blanca  fault.  Murphy  and  Cloud  (1956);  Richter 
(1958). 

Extensive  damage  at  El  Alamo,  Mexico.  Felt 
in  San  Diego  and  much  of  southern  California. 
Wood  &  Heck  (1966). 

Tecate,  B.C.,  Mexico  to  San  Diego.  Probably 
San  Miguel  fault.   Richter  (1965);  Wood  & 
Heck  (1961). 


San  Diego 

+9.6  hrs. 

2.3  ft. 

La  Jolla 

+9.6  hrs. 

0.8  ft. 

L.A.  (Berth  174) 

+9.6  hrs. 

2.3  ft. 

San  Pedro  Bkwtr. 

+9.5  hrs. 

1.7  ft. 

Santa  Monica 

+9.6  hrs. 

4.7  ft. 

Port  Hueneme 

+9.0  hrs. 

2.3  ft. 

Avila,  Calif. 

+9.0  hrs. 

3.3  ft. 

Probably  on  San  Miguel  fault. 
Wood  &  Heck  (1961). 


Richter  (1965); 


60 


Type,  Date,  and  Time 
z  indicates  GMT 
*  indicates  PST 

Generating  region  and 
epicenter,  if  known. 
Magnitude  (Arabic) 
Modified  Mercalli 
Intensity  (Roman) 

Remarks  and  References 

Remote  tsunami 
March  9,  1957,  1422* 


L.A.  Harbor  Berth 


Remote  tsunami 
May  22,  1960,  1911z 


Remote  tsunami 
March  27,  1964,  0336z 


Offshore  earthquake 
December  22,  1964 
(No  tsunami  noted) 


Aleutian  Islands 

A  great  tsunami. 

51.3  N;175.8°W  (8.0-8.5) 

Ensenada,  B.C. 

Mexico 

+6.8  hrs.  3.4  ft. 

San  Diego 

+6.9  hrs.  1.5  ft. 

La  JoUa 

+6.6  hrs.  2.0  ft. 

Newport  Bay 

+6.6  hrs.  0.9  ft. 

Anaheim  Landing 

+6.7  hrs.  2.6  ft. 

Long  Beach 

+6.6  hrs.  1.7  ft. 

San  Pedro  Bkwtr 

+6.6  hrs.  1.2  ft. 

L.A.  Harbor  Term.I  +6.6  hrs.  0.6  ft. 

L.A.  Harbor 

Berth  60 

+7.0  hrs.  2.1  ft. 

L.A.  Harbor  Berth 

174 

+6.9  hrs.  3.1  ft. 

Santa  Monica 

+6.6  hrs.  3.0  ft. 

Port  Hueneme 

+6.5  hrs.  3.5  ft. 

Avila,  Calif. 

+5.8  hrs.  3.5  ft. 

lida,  era/., (1967); 

Salsman(1959);MiUer 

(1964)  BLUE. 

Southern  Chile 

A  great  tsunami. 

41.0°S;73.5"W 

Ensenada,  B.C. 

(8.25-8.5) 

Mexico 

+  13.6  hrs.  8.1  ft. 

San  Diego 

+  14     hrs.  4.6  ft. 

La  JoUa 

+  14     hrs.  3.3  ft. 

Wilson  Cove, 

San  Clemente 

Island 

+  14     hrs.  4.1ft. 

Alamitos  Bay, 

Long  Beach 

+  14.5  hrs.  4.0  ft. 

L.B.  Naval  Ship 

Yard 

+  14.4  hrs.  5.7  ft. 

San  Pedro  Bkwtr. 

+  14.5  hrs.  3.0  ft. 

L.A.  Harbor  Berth 

60 

+  14.5  hrs.  5.0  ft. 

Santa  Monica 

+  14.4  hrs.  9.1  ft. 

Port  Hueneme 

+  14.3  hrs.  8.8  ft. 

Berkman  &  Symons  (1964);  lida,  etai,  (1967); 

Miller  (1964)  RED, 

Prince  William  Sound 

Great  Alaska  Earthquake  and  Tsunami. 

&GuIf  of  Alaska  (8.4) 

Ensenada,  B.C. 

Mexico 

+6.1  hrs.  7.8  ft.+ 

San  Diego 

+6.2  hrs.  3.7  ft. 

La  JoUa 

+5.8  hrs.  2.2  ft. 

Newport  Bay 

+5.8  hrs.  1.3  ft. 

Alamitos  Bay, 

Long  Beach 

+5.9  hrs.  2.8  ft. 

L.A.  Harbor  Berth 

60 

+5.8  hrs.  0.4  ft. 

Santa  Monica 

+5.7  hrs.  2.5  ft. 

Avila,  Calif. 

+5.4  hrs.  5.0  ft. 

N.W.  of  Ensenada,  B.C. 
Mexico;  31.9°N; 
117.1°W(5.5) 


Spaeth  &  Berkman  (1965);  lida,  era/.,  (1967). 
Possibly  on  submerged  portion  of  Agua 
Blanca  fault.   Richter  (1965). 


Remote  tsunami 
October  17,  1966 
Local  inland  earthquake 
April  8,  1968,  1828* 


Local  inland  earthquake 
April  28,  1969,  1521* 


Near  coast  of  Peru 
10.7^S;78.7°W(7.5) 
Ocotillo  Wells,  Calif. 
33°10.5'N;  1 16^^07. 3'W 
(6.5,  VII) 


Borrego  Springs,  C 
33.°21'N;  116^21' 
(5.9,  VII) 


s,  Calif. 
W 


San  Diego,  +10.1  hrs.,  0.25  ft.  Berkman  & 
Carrier  (1967);  lida,  e/ a/.,  (1967). 
Felt  by  and  frightened  all  in  San  Diego.  In- 
tensity VI  in  San  Diego.  Cracks  opened  on  the 
west  side  of  Sunset  CUffs  Boulevard.  Plaster 
cracked  and  fell  in  several  San  Diego  buildings. 
A  9  ft.  concrete  retaining  wall  had  1/8-in.  crack 
from  top  to  bottom,   von  Hake  &  Cloud  (1970). 
Intensity  V  in  San  Diego. 


61 


APPENDIX  REFERENCES 

Allen.  C.  R..  L.  T.  Silver,  and  F.  G.  Stehli 

1960.  Agua  Blanca  Fault-a  major  transverse  structure  of  northern  Baja  California.  Mexico  Geo!  Soc 
Amer.  Bull.  71:457-482. 

Angel.  M. 

1882.  The  history  of  San  Luis  Obispo  County.  Publisher  unknown,  p.  329-330.  (available  Los  Angeles  Main 
Public  Library). 

Bache.  A.  D. 

1856.  Report  of  Superintendent  of  the  United  States  Coast  Survey  for  1855.  Notice  of  earthquake  wave  on 
the  western  coast  of  the  U.S.  on  the  23rd  and  25th  of  December,  1854.  Washington.  D.C..  p.  99;  Appen- 
dix 50,  342-346.  (also  published  in  Amer.  J.  Sci..  Series  H.  21(61):  37-45). 

Bancroft.  H.  H. 

1883.  History  of  California.  A.  L.  Bancroft.  San  Francisco  2:  201,  263.  268.  346-358.  363-368. 

Berkman.  S.  C.  and  D.  C.  Carrier 

1967.  The  tsunami  of  October  17.  1966  as  recorded  by  tide  gauges.  Coast  and  Geodetic  Survey  ms.,  5  p.  (on 
file  in  International  Tsunami  Information  Center.  U.  Hawaii,  Honolulu). 

Berman.  S.  C.  and  J.  M.  Svmons 

1964.  The  tsunami  of  May  22.  1960  as  recorded  at  tide  stations,  U.S.  Dept.  Comm.,  Coast  and  Geodetic  Sur- 
vey. Washington.  D.C..  79  p. 

Berninghausen.  W.  J. 

1962.  Tsunamis  reported  from  the  west  coast  of  South  America.  1562-1960.  Seism.  Soc.  Amer.  Bull.  52(4): 
915-921. 

Bittinger.  C. 

1933.  Experiences  over  a  submarine  epicenter.  Amer.  Geophys.  Union.  Trans  14th  Ann.  Mtg.,  p.  260.  (also  in 
Earthquake  notes.  Seism.  Soc.  Amer.  (Eastern  Section)  June  5(1&2),  p.  260). 

Bodle,  R.  R..  and  L.  M.  Murphy 

1948.  U.S.  earthquakes,  1946.  U.S.  Dept.  Comm..  Coast  and  Geodetic  Survey,  Washington,  D.C.,  Ser.  715. 
45  p. 

Buwalda.  J.  P. 

1933.  The  Long  Beach  earthquake— what  happened  geologically.  Sci.  70(2016):  148-149. 

Byerly.  P. 

l"930.  The  California  earthquake  of  November  4.  1972.  Seism.  Soc.  Amer.  Bull.  20(2):  53-66. 

Clements.  T..  and  K.  O.  Emery 

1947.  Seismic  activity  and  topography  of  the  sea  floor  off  southern  California.  Seism.  Soc.  Amer.  Bull.  37(4): 
309-313. 

Cox.  D.  C. 

1964.  Unpublished  list  of  tsunamis,  p.  96.  In  Weigel,  Oceanographic  engineering.  Prentice-Hall. 

Davidson,  G. 

1872.  Remarks  on  recent  earthquake  waves.  California  Acad.  Sci.,  Proc.  4(5):  268. 

Eaton.  J.  E. 

1933.  Long  Beach.  California  earthquake  of  March  10.  1933.  Amer.  Assoc.  Pet.  Geol.  Bull.  17:  732-738. 

Emery.  K.  O. 

1960.  The  sea  off  southern  California.  John  Wiley  and  Sons,  New  York. 

Green.  C.  K. 

1946.  Seismicsea  wave  of  April  1.  1946  as  recorded  on  tide  gauges.  Amer.  Geophvs.  Union.  Trans.  27(4):  490- 
500. 

Heck.  N.  H. 

1947.  List  of  seismic  sea  waves.  Seism.  Soc.  Amer.  Bull.  37(4):  269-284. 

Heizer,  R.  F. 

1941.  California  earthquakes  of  the  mission  period.  1769-1838,  p.  219-223.  California.  J.  Mines  Geol.,  Rep. 
No.  34  of  the  state  mineralogist,  April  1941. 

Hittell.  T.  H. 

1898.  History  of  California.  N.  J.  Stone  and  Co..  San  Francisco.  2:  547  p. 

Holden.  E.  S. 

1898.  A  catalog  of  earthquakes  of  the  Pacific  coast  from  1769-1897.  Smithsonian  Misc.  Coll..  Washington. 
D.C..  No.  1087. 

lida.  K..  D.  C.  Cox.  and  F.  Parares-Caravannis 

1967.  Preliminary  catalog  of  tsunamis  occurring  in  the  Pacific  Ocean.  Hawaii  Inst.  Geophys.,  Rep.  HIG-67- 
25,  U.  Hawaii. 

Louderback.  G.  D. 

1948.  California  earthquakes  in  1812.  Paper  presented  April  9.  1948,  Seism.  Soc.  Amer.  Ann.  Mtg.  (Abstract: 
Geol.  Soc.  Amer.  Bull..  59(12).  part  2.  December  1948). 


62 


Miller,  G.  R. 

1964.  Tsunamis  and  tides.  PhD.  Thesis.  Univ.  Calif.,  Scripps  Inst.  Oceanogr.,  120  p. 

Munk.  W.  H. 

1953.  Small  tsunami  reaching  California  from  the  Japanese  earthquake  of  March  4,  1952.  Seism.  Soc.  Amer. 
Bull.  43(3):  219-222. 

Murphy,  L.,  and  W.  Cloud 

1954.  U.S.  earthquakes,  1952.  U.S.  Dept.  Comm.,  Coast  and  Geodetic  Survey,  Washington,  D.C.,  ser.  773. 

Murphy,  L.,  and  W.  Cloud 

1956.  U.S.  earthquakes,  1954.  U.S.  Dept.  Comm.,  Coast  and  Geodetic  Survey,  Washington,  D.C..  ser.  793. 

Neumann,  F. 

1935.  U.S.  earthquakes,  1933.  U.S.  Dept.  Comm.,  Coast  and  Geodetic  Survey,  Washington,  D.C.,  ser.  579. 

Proctor,  R.  A. 

1869.  Earthquake  waves  in  the  Pacific.  Nature  1:  54-56. 

Rockwood,  C.  G. 

1879.  Notes  on  American  earthquakes.  Amer.  J.  Sci..  3rd  series,  vols,  for  1872-87. 

Salsman,  G.  G. 

1959.  The  tsunami  of  March  9,  1957  as  observed  at  tide  stations.  U.S.  Dept.  Comm.,  Coast  and  Geodetic 
Survey,  Washington,  D.C.,  Tech.  Bull.  6,  18  p. 

Shuck,  O.  T. 

1869.  Cahfornia  scrap-book;  a  repository  of  useful  information  and  select  readings,  p.  246,  274, 298,  386,  476. 
H.  H.  Bancroft  and  Co.,  San  Francisco  and  New  York. 

Spaeth.  M.  G.,  and  S.  C.  Berkman 

1967.  The  tsunami  of  March  28,  1964  as  recorded  at  tide  stations.  U.S.  Dept.  Comm.,  Coast  and  Geodetic 
Survey,  Washington,  D.C.,  No.  33,  86  p. 

Townley,  S.  D.,  and  M.  W.  Allen 

1939.  Descriptive  catalog  of  earthquakes  of  the  Pacific  coast  of  the  United  States  1769-1928.  Seism.  Soc. 
Amer.  Bull.  29(1):  1-297. 

Trask,  J.  B. 

1856.  Untitled  paper  of  earthquakes  in  California  from  1812  to  1855,  p.  84-86.  California  Acad.  Nat.  Sci., 
Proc.  1  (January  14,  1856). 

Trask,  J.  B. 

1864.  Earthquakes  in  California  from  1800  to  1864,  p.  130-153.  California  Acad.  Nat.  Sci.,  Proc.  3  (April  18, 
1864). 

von  Hockstetter,  F. 

1868.  On  the  earthquake  in  Peru  on  13  August  1868  and  the  flood  wave  caused  by  it  in  the  Pacific  Ocean, 
particularly  on  the  coasts  of  Chile  and  New  Zealand  (in  German).  Sitzungsher,  d.k.  Akad.  d.  Wiss- 
ensch.,  Vienna,  59,  p.  109. 


SAl\J 

(0(0%  LIBRARY      ■ 

Dlu    7  1^7^ 

i' 

J  HARVARO 

t\\  UNIVERSITY 


THE  FEEDING  TECHNIQUES 

OF  STILT  SANDPIPERS  AND  DOWITCHERS 


P.J.  K.  BURTON 


TRANSACTIONS 

OF  THE   SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 


VOL.  17,  NO.  5         16  AUGUST  1972 


THE  FEEDING  TECHNIQUES 

OF  STILT  SANDPIPERS  AND  DOWITCHERS 

P.  J.  K.  BURTON 


ABSTRACT.— This  paper  presents  descriptive  and  quantitative  observations  of  feeding  behavior  of  Stih 
Sandpipers  (Micropalama  himantopus)  and  dowitchers  (Limnodromus  spp.).  Stilt  Sandpipers  make  frequent 
use  of  "stitching" -a  series  of  extremely  rapid  jabs  into  the  mud  surface,  performed  while  pivoting  the  body  or 
walking.  This  is  probably  a  means  of  tactile  foraging;  visual  searching  behavior  is  also  used.  Dowitchers  do 
not  "stitch"  but  employ  isolated  (though  frequent)  deep  jabs  and  probes,  which  are  often  prolonged  and  vig- 
orous. They  show  little  evidence  of  hunting  by  sight.  The  differences  provide  further  evidence  for  assigning 
the  Stilt  Sandpiper  and  dowitchers  to  the  Calidridinae  and  Scolopacinae,  respectively.  Significant  differences 
in  anatomy  of  the  feeding  apparatus  are  summarized. 

The  great  diversity  in  bill  shape  and  size  among  shorebirds  leads  one  to  expect  a  corre- 
sponding diversity  in  feeding  techniques  and  methods;  yet  there  have  been  few  really  de- 
tailed and  quantitative  studies  of  these  techniques.  One  of  the  most  thorough,  concerning 
European  shorebirds,  is  a  little  known  study  by  Streefkerk  (1960).  Detailed  descriptions  of 
feeding  methods  in  five  species  of  shorebirds,  with  a  summary  of  information  available  for 
others,  are  given  by  Burton  (1969,  and  in  press).  The  present  paper  concerns  dowitchers 
{Limnodromus  spp.)  and  the  Stilt  Sandpiper  {Micropalama  himantopus).  Various  descrip- 
tions in  the  literature  suggest  a  close  similarity  between  the  feeding  methods  of  these  birds, 
and  both  are  often  stated  to  employ  a  "sewing-machine  action"  (e.g.,  by  Peterson,  1947). 
The  best  accounts  are  those  of  Bent  (1927)  and  Palmer  (1967),  but  these  include  observa- 
tions from  a  variety  of  sources  and  are  insufficiently  detailed  for  comparative  purposes. 

Dowitchers  and  Stilt  Sandpipers  have  often  been  considered  closely  related,  as  in- 
dicated by  their  juxtaposition  in  many  North  American  lists  until  recently.  According  to  an 
alternative  view  (Lowe,  1931;  Peters,  1934)  the  Stilt  Sandpiper  is  closely  related  to  Calidris 
(Scolopacidae,  subfamily  Cahdridinae),  while  dowitchers  are  allied  to  the  snipes,  Gallinago 
spp.  (subfamily  Scolopacinae).  This  view  is  reinforced  by  recent  work  (Jehl,  1968;  Burton, 
in  press).  As  the  Calidridinae  and  Scolopacinae  differ  extensively  in  feeding  methods,  the 
alleged  similarity  in  this  respect  between  dowitchers  and  the  Stilt  Sandpiper  is  surprising. 
Therefore,  during  the  course  of  a  visit  to  Texas  during  the  latter  half  of  April,  1969,  I  took 
the  opportunity  to  make  detailed  observations  on  their  feeding  behavior. 

METHODS 

Stilt  Sandpipers  were  watched  principally  at  a  brackish  pool  lying  between  agricultural 
land  and  mesquite  brush  near  Alamo,  lower  Rio  Grande  valley,  Texas.  Dowitchers  (mostly 
L.  griseus)weTe  watched  at  tidal  pools  on  the  mudflats  of  the  west  (Laguna  Atascosa)  shore 
of  Padre  Island,  near  Port  Isabel,  Texas.  Observation  was  by  telescope  (30  X  to  60  x). 
Quantitative  aspects  were  studied  by  dictating  running  commentaries  into  a  portable  tape 
recorder.  This  provided  data  amenable  to  statistical  treatment,  and  to  timing  of  various 
items;  some  timed  data  were  also  obtained  in  the  field,  using  a  stop  watch.  Prolonged  exam- 
ination of  qualitative  aspects  of  behavior  were  also  made,  in  addition  to  the  taped  recorded 
commentaries.  The  total  observation  time  (about  20  hours  for  each  species)  is  relatively 
brief  but  served  to  clarify  considerably  the  similarities  and  diff'erences  between  the  two  spe- 
cies in  feeding  methods. 

STILT  SANDPIPER 

Three  principal  types  of  feeding  action  were  distinguished: 
a.  Pecks.— These  are  extremely  brief  movements  made  into  water,  or  at  its  surface,  or 

SAN  DIEGO  SOC.  NAT.  HIST.,  TRANS.  17  (5):  63-68,  16  AUGUST  1972 


64 


into  the  surface  of  exposed  mud. 

b.  Probes.— LongeT  movements  in  which  the  bill  is  thrust  into  the  mud  for  some  depth.  As 
in  the  case  of  Dunlin  {Calidris  alpina)  studied  by  Burton  (in  press),  these  are  normally 
made  with  a  very  rapid  up-and-down  quivering  action.  This,  and  their  distinctly 
longer  duration,  are  the  only  means  of  distinguishing  them  from  pecks  when  (as  fre- 
quently happens)  the  insertion  of  the  bill  into  the  mud  cannot  be  seen. 

c.  Stitching.— This  term  is  used  by  Burton  (in  press)  to  refer  to  a  feeding  action  which 
appears  to  be  characteristic  of  the  Calidridinae.  It  has  previously  been  described  from 
several  members  of  the  subfamily  by  Streefkerk  (1960),  Holmes  (1966), and  others. 
Basically,  stitching  consists  of  very  rapid  series  of  shallow  jabs  into  the  mud  surface, 
made  while  on  the  move;  Holmes  (1966)  refers  to  it  simply  as  a  "rapid  series  of  jabs." 
Stitching  and  probes  may  appear  closely  similar,  and  in  fact  intergrade;  probes  are 
generally  made  in  one  spot,  and  usually  deeper  and  more  vigorously  than  stitching. 

The  Stilt  Sandpipers  invariably  fed  on  mud  covered  by  a  layer  of  water,  sometimes 
barely  covering  the  feet,  but  usually  to  about  tarsus  length,  and  commonly  to  belly  depth.  A 
notable  characteristic  mentioned  by  several  authors  is  their  lack  of  mobility.  This  is  espe- 
cially striking  by  comparison  with  other  species  feeding  near  them  in  similar  situations,  in 
this  case  Lesser  Yellowlegs  {Tringa flavipes)  and  Wilson's  Phalarope  {Phalaropus  tricolor). 
This  lack  of  mobility  is  also  a  contrast  to  most  other  members  of  the  Cahdridinae,  which, 
however,  generally  feed  in  shallower  water  or  on  exposed  substrates. 

Several  accounts  of  the  habits  of  the  Stilt  Sandpiper  mentioned  a  characteristic  atti- 
tude, with  neck  outstretched  and  bill  pointed  vertically  down.  This  attitude  is  indeed  well 
marked  in  this  species,  though  a  similar  attitude  is  quite  often  assumed  by  other  shorebirds 
(e.g.  Redshank,  Tringa  totanus)vj2iding  in  fairly  deep  water.  It  is  restricted  to  spells  of  feed- 
ing by  means  of  pecks;  stitching  series  are  carried  out  with  the  bill  inclined  at  about  80°  to 
the  horizontal,  as  in  other  Calidridinae.  The  more  perpendicular  bill  carriage  where  pecks 
predominate  is  probably  related  to  the  fact  that  these  are  used  in  hunting  by  sight;  the  per- 
pendicular attitude  may  serve  to  minimize  errors  due  to  refraction.  Stitching,  on  the  other 
hand,  appears  to  be  a  form  of  trial  probing  with  the  object  of  detecting  prey  by  tactile 
means.  The  behavior  of  the  birds  while  making  pecks  gave  a  strong  impression  that  they 
were  engaged  in  visual  search.  They  would  walk  slowly  about,  on  a  zig-zag  path  or  back- 
wards and  forwards,  not  covering  a  great  amount  of  ground,  but  maintaining  the  out- 
stretched neck  and  perpendicular  attitude  throughout. 

Stitching  usually  involves  even  less  mobility,  and  one  bird  may  spend  an  hour  or  more 
in  an  area  only  a  yard  or  so  across;  similar  lethargy  mentioned  by  Bent  (1927)  and  Palmer 
( 1967)  probably  refers  to  birds  feeding  in  this  way.  By  contrast  with  most  Calidris  species, 
which  usually  stitch  while  walking,  Stilt  Sandpipers  generally  carry  this  out  while  standing 
still,  the  only  movement  usually  being  a  side  to  side  pivoting  at  the  pelvis  combined  with 
neck  action,  so  that  the  stitching  jabs  are  made  around  it  in  a  semicircle.  The  tracks  left  by 
this  process  should  have  been  highly  characteristic  if  there  were  any,  but  the  mud  surface 
was  much  too  soft  to  retain  indentations.  The  bird  seen  swinging  its  immersed  bill  from  side 
to  side  mentioned  by  Bent  and  Palmer  was  very  probably  stitching.  Palmer's  comparison 
with  a  side  to  side  action  seen  in  the  Greater  Yellowlegs  (Tringa  melanoleuca)  is  probably 
misleading.  A  side  to  side  action  is  shown  by  a  variety  of  Tringinae,  especially  Redshank  (T. 
totanus)  and  Willet  (Catopfrophorus  semipalmatus),  but  close  examination  shows  it  to  be 
accompanied  by  very  rapid  opening  and  shutting  jaw  movements,  a  feature  never  seen  in 
the  stitching  of  Calidridinae  which  it  superficially  resembles. 

Stitching  was  usually  performed  in  more  shallow  water  than  feeding  by  pecks,  though 
some  prolonged  spells  of  stitching  took  place  with  the  head  completely  immersed,  only  the 
quivering  and  slow  pivoting  of  the  body  indicating  what  was  happening.  On  the  rare  occa- 
sions when  stitching  was  carried  out  on  virtually  exposed  mud,  it  could  be  seen  that  the  bill 
was  very  slightly  open  at  the  tip,  as  in  Dunlin.  The  duration  of  321  stitching  sequences  timed 
gave  a  mean  of  2.3  sees  with  a  maximum  of  13.7  sees.  This  is  somewhat  shorter  than  that 
recorded  for  Dunlin  (mean  3.9,  maximum  25.7)  on  a  tidal  mudflat  by  Burton  (in  press) 
though  it  is  pointless  to  pursue  the  comparison  too  closely. 

Probes  were  relatively  brief,  the  great  majority  lasting  under  one  second.  Since  most 


65 


were  made  under  water  (usually  of  belly  depth),  the  depth  to  which  the  bill  was  inserted 
into  the  mud  could  not  normally  be  seen;  when  visible,  the  amount  of  insertion  appeared 
not  less  than  half  the  bill  length,  and  frequently  its  full  length.  No  changes  of  bill  orien- 
tation were  seen  during  the  course  of  a  probe.  The  great  majority  of  probes  were  isolated, 
but  up  to  six  have  been  observed  in  one  spot,  presumably  in  efforts  to  capture  a  particularly 
difficult  prey  animal.  Most  obvious  captures  (indicated  by  head  jerking  and  swallowing 
movements)  followed  probes  rather  than  pecks;  probably  the  items  acquired  by  pecking 
were  mostly  so  small  that  their  capture  went  unobserved. 

The  proportions  of  pecks,  probes  and  stitching  sequences  in  52  minutes  of  timed  obser- 
vations were  recorded  (Table  1).  Each  stitching  sequence  was  counted  as  a  single  move- 
ment. The  mean  number  of  feeding  movements  per  minute  was  40.2  (min.  10,  max.  82). 
High  rates  were  associated  with  a  large  proportion  of  pecks— not  surprisingly,  since  pecks 
are  the  most  rapid  movements.  Conversely,  low  rates  are  associated  with  a  high  proportion 
of  spells  of  stitching,  which  are  of  longer  duration  than  pecks  or  probes.  Interestingly,  high- 
est probing  rates  occur  around  the  middle  of  the  range  of  total  frequencies,  and  relatively 
few  probes  followed  stitching  sequences,  contrasting  with  the  Dunlin  studied  previously 
(Burton,  in  press).  Evidently  visual  signs  provided  the  clues  leading  to  a  probe  in  the  major- 
ity of  cases.  Stitching  thus  appeared  a  relatively  inefficient  method  of  locating  prey  in  this 
area,  though  it  may  have  been  more  important  near  the  edges  of  the  pool,  where  prey  were 
possibly  deeper  lying.  In  other  situations,  and  especially  at  night,  it  may  well  be  of  much 
greater  value. 

Table  1.     Summary  of  timed  observations  on  feeding  movements  of  individual  Stilt  Sandpipers. 


Rate  (Total  move- 

Number of 

Combined 

Stitching 

Peci<s 

Probes 

ments  per  minu 

te) 

minutes 

totals 

10  to  19 

5 

79 

77 

2 

0 

20  to  29 

8 

192 

123 

32 

37 

30  to  39 

14 

483 

8 

197 

278 

40  to  49 

13 

598 

19 

273 

306 

50  to  59 

6 

317 

34 

269 

14 

60  to  69 

3 

190 

3 

151 

36 

70  to  79 

2 

145 

6 

126 

13 

80  to  89 

1 

82 

0 

44 

38 

Overall  total 

52 

2086 

270 
(12.9%) 

1094 

(52.4%) 

722 
(34.5%) 

DOWITCHER 

Only  two  types  of  feeding  action  could  be  distinguished: 

a.  ya^5.— These  are  simple,  brief  movements  in  which  the  bill  is  thrust  into  the  mud  and 
immediately  withdrawn. 

b.  Probes.— More  prolonged  movements  in  which  the  bill  is  thrust  into  the  mud  and  held 
there  for  a  short  time,  usually  accompanied  by  a  rapid  up-and-down  quivering  action. 

Probes  are  generally  deeper  than  jabs,  mostly  between  one  third  and  the  full  length  of 
the  bill  (where  depth  of  insertion  could  be  clearly  seen),  whereas  most  jabs  were  to  less  than 
half  the  bill  length.  However,  the  main  distinction  was  the  brevity  of  jabs,  which  were  gen- 
erally too  rapid  for  accurate  timing,  though  apparently  under  0.5  seconds  in  duration.  103 
probes  timed  averaged  1.7  seconds,  with  maxima  of  4.1  and  7.3  seconds. 

This  classification  is  to  some  extent  arbitrary,  but  probably  most  jabs  are  trials  made  in 
searching  for  prey  by  tactile  means,  while  probes  include  most  of  the  actions  in  which  prey 
are  actually  captured.  Probes  were  often  grouped  in  one  place.  When  this  was  the  case,  they 
were  often  made  with  obvious  vigor,  and  frequently  with  the  capture  of  a  prey  animal,  as 
indicated  by  swallowing  movements.  Presumably,  in  such  cases,  prey  had  been  located,  but 


66 


several  attempts  were  needed  to  complete  its  extraction. 

The  dowitchers  were  nearly  all  feeding  in  tidal  pools,  frequently  up  to  belly  depth;  a 
few  were  watched  feeding  on  exposed  mud.  Though  none  ever  showed  the  remarkable  at- 
tachment to  one  spot  displayed  by  some  Stilt  Sandpipers,  their  mobility  was  not  great.  Most 
commonly,  a  bird  would  concentrate  on  a  small  area  for  about  30  seconds,  probing  around 
itself  with  pivoting  movements  of  the  body  and  a  leisurely  step  or  two;  then  walk  on  more 
briskly  for  a  few  seconds,  and  pause  to  repeat  the  process.  Long  series  of  jabs  were  some- 
times made  while  walking  steadily  forwards;  however,  these  could  not  be  confused  with  the 
stitching  of  Stilt  Sandpipers  and  Calidris  spp.,  as  the  frequency  of  jabs  was  far  less  rapid, 
and  the  bill  was  raised  well  clear  of  the  mud  between  each. 

Rates  of  feeding  movements  were  generally  high.  The  mean  rate  during  61  minutes  of 
timed  observations  (Table  2)  was  60.6  (min.  36,  max.  110).  Not  surprisingly,  high  rates  coin- 
cided with  high  proportions  of  jabs.  Overall,  there  were  slightly  less  (48%)  jabs  than  probes. 
Highest  rates  were  recorded  from  birds  feeding  on  exposed  mud,  which  employed  a  high 
proportion  of  jabs,  and  apparently  met  with  little  success.  Birds  feeding  in  this  situation 
were  occasionally  seen  to  make  short  runs  and  sudden  turns,  suggesting  pursuit  of  prey  lo- 
cated by  sight. 

Between  feeding  actions,  dowitchers  held  the  bill  inclined  at  about  70°  or  80°  to  the 
horizontal.  An  attitude  with  neck  outstretched  and  bill  pointed  vertically  down,  as  in  Stilt 
Sandpiper  was  never  seen.  The  orientation  of  the  bill  was  rarely  altered  to  any  significant 
extent  during  the  course  of  a  probe,  though  on  one  occasion  the  bird  turned  a  full  circle 
around  its  bill  during  a  single  probe.  The  probes  themselves  were  sometimes  made  with 
considerable  force  and  vigor,  quite  unlike  anything  seen  in  the  Stilt  Sandpiper. 

No  prey  item  was  at  any  time  seen.  Several  samples  of  mud  in  areas  favored  by  dowit- 
chers were  dug  up  and  carefully  sifted,  but  the  only  animal  species  found  was  the  small  (5  to 
9  mm.)  bivalve  Lyonsia  hyalina  Conrad.  This  moUusk  is  evidently  abundant  in  the  area,  and 
may  well  have  been  the  main  prey  of  the  dowitchers  observed. 

Table  2.     Summary  of  timed  observations  on  feeding  movements  of  individual  dowitchers. 


Rate  (Total  movements 

Number  of 

Combined 

Jabs 

Probes 

per  minute) 

minutes 

totals 

30  to  39 

2 

74 

10 

64 

40  to  49 

8 

359 

96 

263 

50  to  59 

20 

1099 

455 

644 

60  to  69 

18 

1164 

591 

573 

70  to  79 

10 

726 

402 

324 

80  to  89 

2 

162 

117 

45 

110 

1 

110 

93 

17 

Overall  total 

61 

3694 

1764 

(47.8%) 

1930 

(52.2%) 

COMPARISON 

Dowitchers  and  Stilt  Sandpipers  certainly  show  some  similarities  while  feeding.  Both 
forage  largely  in  pools  and  show  a  generally  high  rate  of  feeding  movements  combined  with 
low  mobility.  Their  feeding  actions  are  mostly  simple,  fairly  regular  movements,  made 
more  or  less  straight  downwards.  The  impression  of  similarity  is  heightened  by  contrast 
with  other  waders  feeding  in  the  same  situations,  notably  Greater  and  Lesser  Yellowlegs, 
whose  brisk  actions  include  dashes  and  sudden  turns. 

Nevertheless,  there  are  well  marked  and  important  differences.  The  most  obvious  of 
these  is  the  absence  of  "stitching"  in  Dowitchers— notwithstanding  the  fact  that  the  actions 
of  both  birds  have  been  likened  with  some  justice  to  a  sewing  machine.  Despite  the  sim- 
ilarity of  imagery,  it  must  be  remembered  that  the  term  "stitching"  as  used  here  and  else- 


67 


where  (Burton  1971,  and  in  press)  applies  specificially  to  an  extremely  rapid  series  of  shal- 
low jabs,  made  with  minimum  head  movement.  This  action,  seen  in  many  calidridine 
sandpipers,  including  the  Stilt  Sandpiper,  was  never  observed  from  dowitchers  during  the 
course  of  these  observations.  Conversely,  Stilt  Sandpipers  rarely  used  deep  test  probes, 
whereas  the  jabs  of  dowitchers  regularly  penetrate  to  a  third  or  more  of  their  considerable 
bill  length.  Probes  in  both  species  are  made  with  a  similar  quivering  action,  but  Stilt  Sand- 
pipers never  exhibit  the  vigor  and  forcefulness  which  is  often  shown  by  probing  Dowitchers. 

The  attitude  with  neck  outstretched  and  bill  pointed  perpendicularly  down  is  charac- 
teristic of  Stilt  Sandpipers  but  is  rarely  shown  by  Dowitchers.  As  explained  earlier,  this  atti- 
tude is  probably  connected  with  hunting  by  sight,  and  indicates  the  much  greater  impor- 
tance of  vision  for  feeding  in  the  Stilt  Sandpiper— a  factor  which  underlies  other  differences 
between  their  feeding  techniques.  Stitching  as  a  means  of  tactile  foraging  increases  the 
chances  of  contact  with  prey  lying  near  the  surface  in  a  given  time,  but  must  be  relatively 
inefficient  for  detecting  deeper  lying  prey.  The  individual  jabs  in  a  stitching  series  are  shal- 
low, and  probably  only  penetrate  the  soft  surface  layer  of  water  covered  mud;  they  require 
relatively  little  anatomical  specialization,  and  form  part  of  a  generally  more  versatile  range 
of  feeding  techniques.  The  generally  deeper  jabs  of  dowitchers  stand  a  greater  chance  of 
detecting  deep  lying  prey,  but  there  are  many  fewer  in  a  given  time.  Also,  since  the  head  is 
fully  raised  and  lowered  between  each  one,  and  the  deeper  penetration  involves  entering  a 
harder  substrate,  the  amount  of  energy  expended  in  proportion  to  the  number  of  contacts 
with  prey  may  well  be  greater  in  dowitchers.  This  is  probably  offset  to  some  extent  by 
greater  tactile  sensitivity  in  dowitchers.  Moreover,  dowitchers  are  capable  of  handling  con- 
siderably larger  prey  than  Stilt  Sandpipers,  and  since  these  tend  to  be  deeper  lying  dowit- 
chers may  be  expected  to  encounter  more  of  them.  The  feeding  technique  and  anatomy  of 
dowitchers  thus  probably  depends  on  relatively  infrequent  contacts  with  larger  prey. 

Detailed  information  on  anatomy  of  the  feeding  apparatus  in  shorebirds  is  given  by 
Kozlova  (1961-62)  and  Burton  (in  press).  The  points  of  difference  between  dowitchers  and 
Stilt  Sandpipers  summarized  below  appear  particularly  relevant  to  a  comparison  of  feeding 
methods. 

a.  The  bill  axis  is  considerably  more  downwardly  directed  relative  to  the  cranium  in 
dowitchers. 

b.  The  dorsal  bar  of  the  upper  jaw  is  greatly  reinforced  in  dowitchers,  and  is  almost  in 
contact  with  the  ventral  bar.  In  the  Stilt  Sandpiper,  both  ventral  and  dorsal  bars  are 
thin  and  widely  separated. 

c.  Hexagonal  pits,  indicating  clusters  of  tactile  receptors  (Herbst's  corpuscles)  are  much 
more  numerous  at  the  tips  of  the  jaws  in  dowitchers. 

d.  M.  protractor  quadrati,  which  raises  the  tip  of  the  upper  jaw,  is  enormous  in  dowit- 
chers by  comparison  with  the  Stilt  Sandpiper. 

e.  M.  adductor  externus  (of  major  importance  for  jaw  closure  and  gripping  prey)  is  rela- 
tively larger  in  dowitchers,  and  of  more  complex  structure,  with  more  pinnate  fiber 
arrangements— a  modification  to  increase  the  force  of  contraction  over  short  distances. 

f.  M.  rectus  capitis  superior,  a  flexor  of  the  head  and  anterior  part  of  the  neck,  lacks  at- 
tachment to  vertebra  4  in  the  Stilt  Sandpiper.  This  curious  feature,  unique  among 
shorebirds,  is  probably  connected  with  its  characteristic  head  attitude  with  bill 
pointed  straight  down,  while  feeding  in  water. 

In  most  of  these,  and  other  anatomical  features  of  head  and  neck,  the  Stih  Sandpiper  is 
typical  of  the  Calidridinae,  whereas  dowitchers  closely  approach  the  Scolopacinae,  though 
showing  some  similarity  to  members  of  the  Tringinae.  Dowitchers  have  by  some  authors 
(e.g.,  Kozlova.  1961-2)  been  considered  more  closely  allied  to  godwits,  but  Jehl  (1968)  has 
produced  strong  evidence  for  their  close  relationship  to  the  Scolopacinae,  first  proposed  by 
Lowe  ( 1931).  The  results  of  this  study  bear  out  Jehl's  view.  The  feeding  technique  of  dowit- 
chers closely  resembles  that  of  Snipe  (Gallinago  gallinago ),  described  in  detail  by  Burton  (in 
press),  in  the  great  reliance  of  both  on  simple  probing,  and  in  the  manner,  timing,  and  dis- 
position of  probes.  They  certainly  show  little  resemblance  to  the  versatile  techniques  of  the 
much  more  mobile  godwits.  Similarly,  the  feeding  behavior  of  Stilt  Sandpipers,  with  its 
frequent  use  of  "stitching"  is  very  similar  to  that  of  other  Calidridinae,  though  with  modifi- 


68 


cations  for  feeding  in  deeper  water  than  most  of  the  subfamily. 

In  any  further  study  of  feeding  in  dowitchers  and  Stilt  Sandpipers,  it  would  be  desir- 
able to  observe  them  in  an  area  where  both  forage  together.  I  saw  them  in  close  proximity 
on  various  stretches  of  shore  in  the  Laguna  Atascosa  Refuge,  but  was  not  able  to  prolong 
my  observations  there.  Such  a  comparison  might  throw  further  light  on  the  results  obtained 
by  Recher  (1966)  in  a  comparison  of  waders  sharing  a  stretch  of  shore.  It  would  be  particu- 
larly interesting  to  know  whether  dowitchers  (the  larger  species)  take  a  narrower  spectrum 
of  prey,  including  more  large  items,  than  the  Stilt  Sandpiper.  Such  a  difference  might  be 
expected  from  Recher's  analysis  of  diets  in  relation  to  body  size,  though  in  the  experience  of 
Jehl  (pers.comm.)  the  reverse  seems  to  be  the  case  at  Churchill,  Manitoba,  where  the  two 
species  often  feed  in  close  proximity. 

ACKNOWLEDGMENTS 

I  am  deeply  indebted  to  Mr.  and  Mrs.  John  J.  Morony  and  John  J.  Morony,  Jr.  and  to  Mr.  and  Mrs.  John 
Arvin;  their  kindness  and  hospitahty  during  my  visit  to  Texas  made  this  study  possible.  I  am  grateful  to  Mr. 
J.  Peake  of  the  British  Museum  (Natural  History)  for  identifying  the  mollusk  collected  at  Padre  Island. 

LITERATURE  CITED 

Bent,  A.C. 

1927.  Life  histories  of  North  American  Shore  Birds.  Part  I.  U.S.  Natl.  Mus.  Bull.  146. 

Burton,  P.  J.  K. 

1969.  Anatomy  and  adaptive  modifications  of  the  feeding  apparatus  in  waders  (Aves:  Charadrii).  Ph.D. 
thesis.  University  of  London. 

1971.  Comparative  anatomy  of  head  and  neck  in  the  Spoon-billed  sandpiper,  Eurynorhynchus  pygmeus  and 
its  allies.  J.  Zool.,  Lond.,  163:  145-163. 

In  press.  Feeding  and  the  feeding  apparatus  in  waders.  British  Museum  (Natural  History). 

Holmes,  R.  T. 

1966.  Feeding  ecology  of  the  Red-Backed  Sandpiper  {Calidris  alpina)  in  Arctic  Alaska.  Ecology,  47:  32-45. 

Jehl,  J.  R.,  Jr. 

1968.  Relationships  in  the  Charadrii  (shorebirds):  a  taxonomic  study  based  on  color  patterns  of  the  downy 
young.  San  Diego  Soc.  Nat.  Hist.,  Memoir  3. 

Kozlova,  E.  V. 

1961-1962.  Fauna  S.  S.  S.  R.,  Zool.  Inst.  Akad.  Nauk  S.  S.  S.  R.,  nov.  ser.  no.  81,  Ptitsy,  2,  no.  1,  pt.  3  [Birds, 
Charadriiformes.  Suborder  Limicolae]  Moscow-Leningrad.  Akad.  Nauk.  S.  S.  S.  R. 

Lowe,  P.  R. 

1931.  An  anatomical  review  of  the  "waders"  (Telmatomorphae).  Ibis,  1931:  721-771. 

Palmer,  R.  S. 

1967.  Species  accounts,  In  G.  D.  Stout  (ed.).  The  Shorebirds  of  North  America.  Viking  Press,  New  York. 

Peters,  J.  L. 

1934.  Check-list  of  birds  of  the  world.  Vol.  2.  Harvard  Univ.  Press,  Cambridge. 

Peterson,  R.  T. 

1947.  A  Field  Guide  to  the  Birds.  Houghton-Mifflin  Co.,  Boston. 

Recher,  H.  F. 

1966.  Some  aspects  of  the  ecology  of  migrant  shorebirds.  Ecology  47:  393-407. 

Streefkerk,  C.  J. 

1960.  Verslag  van  het  vergelijkend  onderzoek  naar  de  wijze  van  voedsel  zoeken  van  enige  soorten  steltlopers. 
Uitgegeven  door  de  Christelijke  Jeugdbond  van  Natuurvrienden,  Amsterdam. 


Sub-Department  of  Ornithology,  British  Museum  (Natural  History),  Tring,  Hertford- 
shire, England. 


cS4a/ 


UBRARY 


THORACIC  CIRRIPEDIA  FROM  GUYOTS 
OF  THE  MID-PACIFIC  MOUNTAINS 


M.  V.  LAKSHMANA  RAO  AND  WILLIAM  A.  NEWMAN 


TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 


VOL.  17,  NO.  6     31  AUGUST  1972 


THORACIC  CIRRIPEDIA  FROM  GUYOTS 
OF  THE  MID-PACIFIC  MOUNTAINS 


M.  V.  LAKSHMANA  RAO  AND  WILLIAM  A.  NEWMAN 


ABSTRACT.— Knowledge  of  the  fauna  of  oceanic  seamounts  is  meager.  To  determine  whether  seamounts 
serve  as  stepping  stones  for  the  distribution  and  dispersal  of  sedentary  faunas  across  oceanic  barriers,  and 
their  role  in  the  biogeography  and  speciation  of  deep-sea  faunas,  an  expedition  went  to  the  Mid-Pacific 
Mountains  in  the  summer  of  1968.  This  paper  reports  on  Thoracic  Cirripedia  from  six  guyots  located  there. 
Nine  species  were  identified,  of  which  four  are  new.  Three  of  the  new  species  are  allied  to  forms  from  the 
Indo-Pacific;  the  fourth  is  closely  related  to  a  Hawaiian  species.  Of  the  five  previously  known  species,  two  are 
widely  distributed  in  the  Indo-Pacific,  two  are  cosmopohtan  and  one  is  endemic  to  Hawaii.  Thus,  the  affinities 
of  the  cirripeds  are  predominantly  Indo-Pacific. 

Virtually  nothing  is  known  of  the  faunas  of  submarine  archipelagos.  One  would  like  to 
know  specifically  what  role  seamounts  serve  as  stepping  stones  at  bathyal  depths,  what  im- 
portance they  have  in  the  evolution  of  deep-sea  faunas,  and  to  what  degree  these  faunas 
tend  to  be  endemic.  In  the  summer  of  1968,  Wilham  A.  Newman  and  Richard  H.  Rosen- 
blatt, both  of  the  Scripps  Institution  of  Oceanography,  John  A.  Allen  of  the  Dove  Marine 
Laboratory,  England,  and  the  late  Edwin  C.  Allison  of  San  Diego  State  College,  and  Harry 
S.  Ladd  of  the  U.S.  Geological  Survey  staged  an  expedition  (Styx-Leg  7)  aboard  the  R/  V 
Alexander  Agassiz  to  investigate  both  recent  and  extmct  faunas  of  the  Mid-Pacific  Moun- 
tains. 

The  Mid-Pacific  Mountains  are  a  chain  of  seamounts  located  between  17°  and  23°N, 
with  the  main  axis  extending  between  165° W  and  170°E  for  some  2,780  km  (Fig.  1).  This 
chain  of  seamounts,  part  of  the  Marcus-Necker  Ridge,  forms  the  northeastern  portion  of 
the  region  designated  as  the  Darwin  Rise  (Menard,  1964).  Numerous  flat-topped 
seamounts  occur  along  this  chain  at  depths  ranging  between  approximately  1,000  and  1,700 
m.  Shallow  water  megafossils  taken  by  dredging  indicate  that  many  flat-topped  seamounts 
are  guyots,  land  forms  produced  by  subaerial  erosion  and  marine  planation  at  a  time  when 
they  broke  the  sea  surface.  The  fossils,  particularly  the  rudist  molluscs  and  associated  or- 
ganisms such  as  corals,  indicate  that  the  seamounts  persisted  as  shaflow  water  banks  and 
reefs  up  to  the  mid-Cretaceous  before  subsiding  more  than  a  kilometer  to  their  present 
depths  (Hamilton,  1956).  Prior  to  this  time,  the  Mid-Pacific  Mountains  formed  an  extensive 
island  chain,  comparable  to  the  present  Hawaiian  Archipelago.  Since  formation,  the  chain 
has  migrated  northwest  some  25°  to  its  present  position  (Lonsdale  et  ai,  1972).  Con- 
sequently the  chain  has  always  been  beneath  tropical  waters. 

The  tops  of  the  guyots  have  been  altered  to  varying  degrees  since  they  subsided  (Karig 
et  ai,  1970;  Lonsdale  et  ai,  1972).  Virtually  all  exposed  hard  surfaces,  such  as  rudist  reefs, 
limestone,  exhumed  chert  and  basalt  outcrops  are  covered  with  ferromanganese  oxides  of 
varying  thickness.  For  some  reason  fresh  manganese-coated  surfaces  appear  unfavorable 
for  attachment  of  benthic  organisms  and  the  numerous  large  slabs  and  nodules  dredged 
from  the  Mid-Pacific  Mountains  were  devoid  of  them.  Generally  pieces  of  pumice,  appar- 
ently of  recent  origin,  and  occasional  small  rocks  (cherts)  are  free  of  manganese  coatings. 
Otherwise,  uncoated  hard  surfaces  on  which  sedentary  organisms  might  be  expected  to 
settle  and  attach  are  hmited  to  the  hard  parts  of  living  organisms  such  as  spicules  of  si- 
liceous sponges,  shells  of  gastropods,  barnacles  and  corals  usually  occurring  on  soft  sedi- 
ments. In  the  present  collection  cirripeds  were  taken  from  all  these  with  the  exception  of  the 
corals. 

SampHng  methods  were  varied.  Pipe  and  chain-bag  dredges  were  employed  on  hard 
bottoms  and  outcrops;  otter  and  beam  trawls  over  soft  bottoms.  A  variety  of  benthic  in- 
vertebrates was  recovered  and  the  present  paper  reports  on  the  class  Cirripedia.  Only  mem- 

SAN  DIEGO  SOC.  NAT.  HIST,  TRANS.  17(6);  69-94,  31  AUGUST  1972 


70 


25° 


350CX 
1464 

Horizon  Guyot 


170°E 


180 


70°  W 


Figure  1.  Chart  indicating  the  location  of  the  guyots  sampled  on  the  Mid-Pacific  Mountains  during  the  Styx-7 
Expedition.  Depths  in  meters.  Numerous  other  guyots  in  the  region  not  indicated.  Of  the  guyots  indicated,  only 
Darwin,  Hess  and  Horizon  have  been  named  previously. 

bers  of  the  order  Thoracica  were  encountered  on  the  six  guyots  sampled  (Table  1).  (Station 
numbers  for  this  leg  of  the  expedition  were  numerical:  year,  month,  day,  1,  2  or  3  etc.). 

Of  the  nine  identifiable  species  found,  four  are  new,  the  relative  number  of  new  species 
being  comparable  to  that  of  the  Antarctic  (Newman  and  Ross,  1971),  One  cannot  assume 
that  all  four  new  species  from  the  Mid-Pacific  Mountains  are  endemic  to  the  region  because 
knowledge  of  deep-sea  cirripeds  is  meager  (see  Zevina,  1972).  While  one  of  the  new  species 
is  most  closely  related  to  a  species  known  previously  only  from  Hawaii,  the  remaining  three 
show  close  affinities  with  forms  widely  distributed  in  the  Indo-Pacific.  Of  the  five  previously 
known  species,  two  are  known  from  the  Indo-Pacific,  one  from  Hawaii  (and  thus  perhaps  an 
Indo-West  Pacific  derivative  endemic  to  Hawaii),  and  two  are  cosmopolitan.  Thus,  as  one 
might  have  anticipated,  the  affinities  of  the  cirriped  fauna  of  the  Mid-Pacific  Mountains  are 
primarily  with  the  Indo-Pacific.  The  fishes  show  comparable  affinities  (R.H.  Rosenblatt, 
pers.  comm.). 

While  virtually  nothing  is  known  of  the  faunas  of  other  submarine  archipelagos,  the 
fauna  of  Shoal  Guyot,  situated  at  approximately  25°  S,  85°  W,  some  1,300  km  west  of 
South  America  at  a  depth  of  288  m,  is  relatively  well  known  and  is  of  considerable  bio- 
geographic  interest.  Hubbs  ( 1959)  published  on  fishes,  and  information  on  echinoderms  and 
barnacles  was  given  by  Zullo  et  al.  (1964),  Zullo  and  Newman  (1964)  and  Allison  et  al. 
( 1967).  One  might  have  expected  the  fauna  of  Shoal  Guyot  to  be  strongly  Eastern  Pacific  in 
character  since  it  is  separated  from  the  Indo-Pacific  by  the  so-called  East  Pacific  Barrier.  To 
the  contrary  however,  it  proved  to  be  primarily  an  eastward  extension  of  the  Indo-West 
Pacific  and  thus  would  appear  to  be  the  eastern  terminus  of  a  series  of  submarine  stepping 
stones  at  bathyal  depths  across  the  East  Pacific  Barrier  (AUison  et  al,  1967).  The  situation  is 
more  complicated  than  this  however,  for  there  is  apparently  a  peculiar  extension  of  neretic 
plankton  toward  this  region  from  the  west  (A.  Fleminger,  pers.  comm.),  and  this  indicates 
that  the  eastward  extension  of  the  Indo-Pacific  fauna  is  not  simply  by  way  of  submarine 
stepping  stones  in  this  region,  as  previously  supposed. 

SYSTEMATIC  ACCOUNT 

Order  Thoracica  Darwin,  1854 

Suborder  Lepadomorpha  Pilsbry,  1916 

Family  Scalpellidae  Pilsbry,  1916 

Genus  A  rcoscalpelluni  Hoek,  1907 


Arcoscalpellum  alcockianum  (Annandale),  1905 
Figures  2  and  1 IG 

ScalpeUitm  alcockianum  ^x\nlm&^\Q.  1905:82;  1906a:  138;  1906b:392:  1913:229;  1916: 129,  pi.  vi, 
man,  1918a:115;  Nilsson-Cantell,  1928:6;  1931:2;  1938:7. 


fig.  5;  Cal- 


71 


Table  I.     Cirripedia  from  guyots  of  the  Mid-Pacific  Mountains 


Species 

Horizon 

Hess 

Allison 

Agassiz 

Sio       Darwin 

Previously  known  distributions 

Sources 

LEPADOMORPHA 

Family:   Scalpellidae 

l.'^Arcoscalpellum  sp. 

1718m 

Present  report 

l.ArcoscalpMum  alcockianum 

i.Arcoscatpellum  etegqntissimum 

n.  sp. 

1652- 
1670m 

1418- 
1664m 

1566m 

Indian  Ocean ;  Mozambique  Channel;  Gulf  of 

Manaar;  Bay  of  Bengal;  Malay  Archipelago, 

SW  Pacific  Ocean;  (109S-1800m) 

Lukunor  Atoll.  Caroline  Islands;  (972m) 
(present  report) 

Annandale  (1906); 
Caiman  (1918a); 
Nilsson^'antell 
(1938) 

Present  report 

A.Arcoscalpellum  hawaiiense 

1415- 
1557m 

Hawaii;  (1460m) 

Pilsbry  (1907a) 

S.Arcoscatpellum  michelottianum 

1584- 
1800m 

1692- 
1735m 

1413- 
1645  m 

1557m 

Cosmopolitan  -  Atlantic,  Indian,  Pacific 
and  Antarctic  Oceans;  (40-2900m) 

Nilsson-CanteU 
(1938);  Newman 
and  Ross  (1971) 

S.Arcoscalpellum  radiatum 

1584- 
1800m 

1413- 
1645m 

Present  report 

1  .Arcoscalpellum  rossi  n.  sp. 

1692- 
1735m 

1413- 
1645m 

Present  report 

S.Mesoscatpellum  gruvelii 
Family:    Poecilasmalidac 

1429- 
1663m 

Indian  Ocean;  Gulf  of  Aden;  Laccadives; 
Gulf  of  Manaar;  Andaman  Sea;  (794-2268m) 

Nilsson-Cantell 
(1938) 

9.Megalasma  (Glyptelasma) 
pihbryi 

1445- 
1557m 

Indian  Ocean;  Malay  Archipelago;  Pacific 
and  Atlantic  Oceans;  (1098-1647m) 

Nilsson-Cantell 
(1938) 

VERRUCOMORPHA 

10.  Verruca  {Altiverruca) 
allisoni   n.  sp. 

1718- 
1770m 

1413- 
1645  m 

1300- 
1353m 

Present  report 

Material. -Slyx-7,  680903-04  Sta.  1.  Allison  Guyot  ( 18°3rN,  179°36'W),  1418-1664  m  (otter  trawl).  One  her- 
maphrodite on  long  glassy  spicules  of  a  siliceous  sponge. 

Supplementary  description  (hermaphrodite).— The  capitular  plates  were  adequately  de- 
scribed by  Annandale  (1906b).  Evidently  the  capitulum  and  the  peduncle  are  subject  to 
considerable  variation  (Annandale,  1913;  Nilsson-Cantell,  1928).  In  the  present  specimen 
as  compared  to  those  shown  by  Annandale  (1916),  tergum  is  not  as  reduced,  the  scutal  mar- 
gin is  hollowed  out,  and  the  carinal  margin  is  angular  and  recedes  from  the  carina  both 
above  and  below.  The  scutum  is  not  fully  calcified,  and  the  calcified  portion  is  triangular, 
reaching  to  the  lower  extremity  of  the  occludent  margin  of  the  tergum;  the  apex  is  terminal. 
A  rostrum  is  present. 

The  peduncle  is  cylindrical,  almost  as  high  as  the  capitulum  and  armed  with  about  13 
rows  of  transversely  elongate  plates.  There  is  no  basal  disc  in  the  present  specimen,  appar- 
ently reflecting  the  substrate  to  which  the  specimen  is  attached. 

Of  the  arthropodal  structures  brief  descriptions  were  given  by  Annandale  and  Nilsson- 
Cantell  together  with  figures  of  the  mandible,  maxillae  and  caudal  appendages.  The  follow- 
ing account  and  accompanying  figures  are  supplementary. 

Labrum  bullate,  very  broad  distally  and  mottled  by  pigment  all  over  the  surface;  teeth 
very  small;  palp  elongate,  pointed  at  the  tip  and  covered  with  plumose  spines  along  the 
entire  margin  (Fig.  2D).  Maxilla  I  having  nearly  straight  cutting  edge  divided  into  two  steps 
with  about  13  strong  spines  above,  and  approximately  10  strong  and  18  weaker  spines  of 
about  equal  length  below  (Fig.  2E).  Maxilla  II  large,  with  superior  margin  long,  supporting 
a  continuous  row  of  spines;  a  medial  notch  as  noticed  by  Nilsson-Cantell  ( 1928);  maxillary 
lobe  short,  broad  and  truncate  apically  (Fig.  2F).  Mandible  with  four  teeth  including  infe- 
rior angle;  first  tooth  well  separated  from  second,  third  tooth  nearer  to  inferior  angle  than  to 
second  (Fig.  2B);  inferior  angle  supporting  30-31  short  blunt  subspatulate  spines  some  of 
which  are  bifid  (Fig.  2C). 

Cirrus  I  widely  separated  from  the  rest;  intermediate  segments  of  the  anterior  ramus 
strongly  protuberant,  those  of  the  posterior  ramus  cylindrical  and  2/3  as  wide;  both  rami 
prominently  hairy  (Fig.  2A).  Cirrus  II  nearly  Wi  times  as  long  as  cirrus  I  and  cirrus  III  a 
little  longer  than  the  second.  Cirri  V  and  VI  with  terminal  segments  missing.  Segment  18  of 
cirrus  VI  is  figured  (Fig.  2G);  articular  areas  along  greater  curvature  with  2-4  long  and  1-2 
short  setae;  lateral  faces  with  a  few  short  setae;  interarticular  areas  devoid  of  setae  and 
bristles.  Setation  ctenopod;  four  major  pairs  of  setae  along  lesser  curvature.  Between  each 


72 


Figure  2.  Arcoscalpellum  alcockianum  (Annandale),  Styx-7,  680903-04  Sta.  1.  A,  cirrus  I;  B,  mandible:  C,  third 
tooth  and  inferior  angle  of  mandible;  D,  palp;  E,  maxilla  I;  F,  maxilla  II;  G,  intermediate  articles  of  cirrus  VI;  H, 
caudal  appendage;  I,  penis. 

major  pair  there  are  1-2  long  bristles.  Caudal  appendage  of  29  segments  reaching  to  at  least 
half  the  length  of  cirrus  VI.  Each  segment  with  1-5  setae  along  outer  margin,  terminal  seg- 
ment with  a  tuft  of  5  short  setae  at  tip  (Fig.  2H). 

Penis  short,  moderately  stout,  covered  with  small  hairs  and  annulated  in  the  proximal 
part;  distal  end  narrow  and  covered  with  minute  hairs  (Fig.  21). 

Two  complemental  males  were  recovered,  one  from  each  pouch  near  the  tip  of  the  in- 


73 


side  of  each  scutum.  The  male  is  sac-like,  without  traces  of  valves  or  cirri,  but  the  mantle  is 
covered  with  rows  of  spines  and  supports  two  prehensile  antennae  at  the  middle  of  the  ven- 
tral margin. 

Remarks.— This  species  is  apparently  distributed  widely  in  the  Indian  Ocean,  having 
been  reported  several  times  from  the  Bay  of  Bengal  and  Malay  Archipelago.  The  only 
record  outside  of  this  area  is  in  an  unpublished  report  by  Caiman  on  specimens  taken  be- 
tween Australia  and  New  Zealand  (Nilsson-Cantell,  1928).  The  present  report  extends  its 
range  far  east  into  the  Pacific. 

Arcoscalpellum  giganteum  (Gruvel)  from  the  Atlantic  is  closely  related,  but  its  tergum 
is  much  hollowed  and  the  caudal  appendage  consists  of  only  four  segments. 

Arcoscalpellum  elegantissimum  n.  sp. 

Figure  3 

Malerial.-Styx-l.  680907  Sta.  1,  Agassiz  Guyot  (17°50.6'  N,  178°25.0'  W),  1566  m  (otter  trawl)  1  spec; 
680829  Sta.  3.  Horizon  Guyot  (19°28.0'  N.  168°52.3'  W).  1652-1670  m  (rock  dredge),  1  spec;  CARMARSEL 
Exped.  Sta.  815  (off  Lukunor  Atoll,  Caroline  Islands,  10  March  1967),  972  m,  2  spec. 

Depositorv.-V.S.NM.  no.  140943  (Holotype.  Stvx-7,  680907,  Sta.  1)  U.S.N.M.  no.  140944  (Paratype,  Styx-7, 
680829,  Sta.  3,'  1  spec.)  U.S.N.M.  nos.  140945,  140946  (CARMARSEL  Exped.  Sta.  815,  2  spec) 

Diagnosis.— Capitulum  with  14  fully  calcified  approximate  plates  ornamented  with 
strong  radial  ridges.  Carina  broad  basally;  carinal  roof  traversed  by  longitudinal  ridges; 
parietes  well  developed.  Carinal  latus  as  broad  as  high.  Rostral  latus  wider  than  high.  Ros- 
trum ovotriangular  and  fully  exposed.  Inframedian  latus  higher  than  rostral  latus  but 
shorter  than  carinal  latus.  Mandible  with  4  teeth  including  inferior  angle.  Maxilla  I  with 
straight  cutting  edge.  Intermediate  segments  of  cirrus  VI  with  2-3  major  and  1  minor  pair  of 
setae.  Caudal  appendages  with  4  partially  fused  segments  reaching  %  the  length  of  first  seg- 
ment of  pedicel  of  cirrus  VI. 

Description  (female).— Capitulum  globose,  ovally  elongate,  apically  pointed,  hirsute 
especially  on  the  carinal  side;  14  fully  calcified  plates,  white,  with  no  indication  of  a  per- 
sistent cuticle.  Plates  ornamented  with  prominent  ridges,  radiating  from  the  umbones,  in- 
tersected by  faint  growth  lines  (Fig.  3 A). 

Tergum  nearly  twice  the  area  of  the  scutum,  rhomboid,  twice  as  long  as  wide;  apex 
prominently  acute;  basicarinal  angle  reaching  about  %  the  distance  towards  the  base  of  the 
capitulum,  nearly  to  the  lower  whorl  of  the  plates;  lateral  margin  partly  overlapped  by  the 
upper  latus.  Scutum  subquadrate,  more  than  twice  as  long  as  broad;  surface  convex,  ap- 
pearing divided  into  halves  by  a  diagonal  angulation  running  from  the  umbo  to  the  basil- 
ateral  angle.  Carina  strongly  bowed,  broad  basally,  tapering  towards  apex;  roof  essentially 
fiat,  traversed  by  prominent  longitudinal  ridges;  parietes  well  developed  and  also  promi- 
nently ridged.  Carinal  latera  meet  for  a  short  distance  at  base  of  carina  forming  a  broad  V- 
shaped  margin  (Fig.3B);  each  as  broad  as  high,  with  an  inwardly  curved  apex  which  pro- 
jects slightly  beyond  the  surface  of  the  capitulum;  basal  and  lateral  margins  irregular;  two 
ledges  running  from  umbo  to  base  divide  plate  into  two  parts,  a  shallow  wing-like  expan- 
sion at  base  of  upper  latus  and  two  triangular  areas  (one  a  raised  carinal  part  adjoining  the 
carina  and  the  other  a  concave  middle  portion  between  this  and  the  wing-like  portion).  In- 
framedian latus  triangular,  slightly  higher  than  broad;  apex  raised  above  the  surface  of  ca- 
pitulum and  curved  inwards.  Rostral  latus  twice  as  broad  as  high;  scutal  and  basal  margins 
subparallel;  plate  diagonally  divided  into  halves  by  a  faint  ridge;  apices  of  both  sides  partly 
overlapped  by  rostrum  (Fig.  3C).  Rostrum  ovotriangular,  broad  at  anterior  end  and  narrow 
posteriorly  (Fig.  3C).  Peduncle  short,  Va  height  of  capitulum  and  armored  with  8-10  rows  of 
4-5  closely  packed,  narrow  and  elongate  scales.  Measurements  (in  mm)  of  the  holotype  fol- 
low: overall  height,  16.5:  height  of  capitulum,  13.0:  height  of  peduncle,  4.0. 

Labrum  bullate,  no  soft  setae  present;  crest  armed  with  about  45  teeth.  Palp  elongate, 
triangular,  somewhat  rounded  distally;  proximal  superior  margin  with  short  stiff  bristles; 
distal  border  with  long  setae  (Fig.  3H).  Mandible  with  four  teeth  including  inferior  angle; 
second  tooth  well  separated  from  first  (Fig.  3D);  inferior  angle  with  13-15  triangular  to  sub- 
spatulate  teeth  a  few  of  which  are  bifid  (Fig.  3E).  Maxilla  I  with  cutting  edge  feebly  concave 
above  and  convex  below;  concave  part  supports  2  long,  stout  and  3-4  shorter,  thinner  spines 
(Fig.  3F).  Maxilla  II  triangular  in  shape,  lobes  weakly  developed;  marginal  setae  dis- 


74 


^^^i!^-^^ 


,AC 

5.0  mm 

,B 
,DFHIJ 

SO 

0.5 

,E 

0.2 

,6 

_0.5 

,K 

0.5 

Figure  3.  Arcoscalpellum  elegantissimum  n.  sp.,  Holotype,  Styx-7  680907  Sta.  1,  Agassiz  Guyot.  A,  side  view  of 
female;  B,  carinal  view;  C,  rostral  view;  D,  mandible;  E,  third  tooth  and  inferior  angle  of  mandible;  F,  maxilla  I; 
G,  maxilla  11;  H,  palp;  1,  cirrus  I;  J,  intermediate  articles  of  cirrus  VI;  K.  caudal  appendage. 

tributed  in  three  clusters,  but  those  of  the  superior  and  distal  borders  are  contiguous;  max- 
illary lobe  short,  broad  and  truncate  apically  (Fig.  3G). 

Cirrus  I  well  separated  from  the  rest;  anterior  ramus  shorter  than  posterior;  inter- 
mediate segments  of  anterior  ramus  protuberant,  those  of  posterior  cylindrical  and  %  as 


75 


wide;  both  rami  clothed  with  long  setae  (Fig.  31).  Cirrus  II  normal.  Cirri  IV-VI  nearly  equal 
in  length  with  equal  or  sub-equal  rami.  Each  articulation  along  greater  curvature  of  inter- 
mediate articles  of  Cirrus  VI  supporting  4-5  short  slender  setae.  Interarticular  areas  along 
greater  curvature  and  lateral  faces  free  of  setae.  Setation  ctenopod;  2-3  major  pairs  and  1 
minor  pair  with  1-2  slender  spines  at  bases  of  major  setae  (Fig.  3J).  Caudal  appendage  com- 
posed of  4  stout,  partially  fused  segments,  extending  about  %  length  of  pedicel  of  Cirrus  VI; 
distal  article  with  tuft  of  3-5  setae,  2  or  3  being  longer  than  appendage  (Fig.  3K).  Cirral 
counts  of  the  four  specimens  follow: 

I  II  III  IV  V  VI  Ca 


Styx-7  680907 

8 

22 

22 

24 

25 

25 

Sta.  1 

Agassiz  Guyot 
(Holotype) 

12 

8 

13 

20 
21 
22 

21 

22 
24 

25 
21 
25 

17+ 

22 

24 

26 
26 
26 

Styx-7  680829 

7 

20 

23 

25 

15+ 

25 

Sta.  3  (Paratype) 

12 

20 

23 

22 

23 

23 

7 

20 

22 

13+ 

23 

23 

12 

20 

23 

23 

23 

23 

CARMARSELSta.  815 

8 

20 

22 

22 

24 

25 

(Paratype) 

12 

20 

21 

22 

24 

25 

7 

16+ 

21 

22 

16+ 

12+ 

12 

19 

21 

22 

23 

23 

CARMARSELSta.  815 

7 

15 

16 

16 

17 

17 

(Paratype) 

9 

15 

15 

15 

17 

17 

7 

14 

15 

16 

15 

17 

9 

14 

15 

18 

17 

18 

Remarks.— Arcoscalpellum  elegantissimum  is  closely  related  to  the  A.  michelottianum 
group  of  scalpeUids  (Newman  and  Ross,  1971).  Important  uniting  characters  are:  a  promi- 
nently hirsute  capitulum,  a  triangular  inframedian  latus  not  exceeding  the  height  of  the 
carinal  latus  and  with  an  apical  umbo,  carinal  lateral  plates  not  interdigitating  where  they 
meet  below  the  carina,  and  a  rostral  latus  almost  twice  as  wide  as  high.  The  new  species  is 
closely  alhed  to^.  hawaiiense  {ViX^ibry)  but  differs  in  having:  1)  apex  of  carinal  latus  curved 
inwards,  not  projecting  beyond  the  base  of  the  carina;  2)  posterior  ramus  of  cirrus  I  only 
slightly  longer  than  the  anterior  ramus  but  with  \Vi  times  as  many  segments;  3)  caudal  ap- 
pendage with  but  four  incompletely  fused  segments  and  less  than  the  height  of  the  first 
segment  of  the  pedicel;  4)  scales  on  the  peduncle  not  overlapping  or  imbricating  and  5)  a 
much  smaller  size.  In  its  small  size  and  the  general  nature  of  the  arthropodal  structures /i. 
elegantissimum  shows  some  resemblance  to  A.  hirsutum  (Hoek).  However,  in  the  latter  spe- 
cies the  roof  of  the  carina  does  not  possess  longitudinal  ridges  and  the  carinal  latus  is  less 
elaborately  developed. 

The  specific  name  refers  to  the  elegant  capitular  ornamentation. 

Arcoscalpellum  hawaiiense  (Pilsbry),  1907 
Figures  4  and  1 1  C-D 

Scalpellum  hawaiiense  Pilsbry  1907a:  181.  pi.  IV,  fig.  1-2. 

Material. -Styx-7,  680905  Sta.  2,  Allison  Guyot  ( 179°37.r  W,  18°35.4'  N)  1450-1557  m  (otter  trawl).  1  spec, 
attached  to  a  small  rock. 

Supplementary  description  (female).— The  capitular  structure  of  this  scalpellid  agrees 
well  with  the  description  of  the  type  from  Kauai,  Hawaii  (Pilsbry,  1907a).  The  present  spec- 
imen, larger  than  the  type,  has  the  following  dimensions  (in  mm):  overall  height,  44;  capitu- 
lar height,  3 1 ;  width  of  capitulum,  22;  height  of  peduncle,  14. 

The  arthropodal  structures  were  not  described  and  are  dealt  with  here.  Labrum  bul- 
late,  longer  than  broad,  apex  gently  curving,  surface  mottled  with  pigment.  Palp  long  and 
narrow,  superior  and  anterior  margins  clothed  with  thick,  short,  slightly  plumose  setae;  an- 


76 


terior  margin  and  lateral  faces  naked  (Fig.  4F).  Mandible  with  4  teeth  including  inferior 
angle;  teeth  more  or  less  equidistantly  spaced  (Fig.  4B);  inferior  angle  supporting  about  14 
teeth  many  of  which  are  worn  and  blunt  (Fig.  4C).  Maxilla  I  with  cutting  edge  nearly 
straight  without  evident  notch;  upper  half  supporting  7  spines,  the  uppermost  2  long  and 
stout,  the  rest  shorter  and  thinner;  lower  portion  supporting  17-18  long  and  short  spines 
(Fig.  4D).  Maxilla  II  triangular,  with  3  weakly  developed  lobes;  marginal  setae  distributed 
in  three  clusters,  those  of  superior  and  anterior  margins  being  longer;  lateral  faces  devoid  of 
setae  (Fig.  4E). 

Cirrus  I  widely  separated  from  the  rest;  posterior  ramus  IVi  times  longer  than  ante- 
rior ramus;  intermediate  segments  of  anterior  ramus  strongly  protuberant,  those  of  pos- 
terior ramus  cylindrical  and  %  as  wide  (Fig.  4A).  Cirrus  II  normal.  Cirri  III-IV  about  equal 
in  length  with  equal  or  subequal  rami.  Each  articulation  along  greater  curvature  of  inter- 
mediate segments  of  cirrus  VI  with  a  cluster  of  2-5  short  setae.  Setation  ctenopod;  3  major 
pairs  of  setae  along  lesser  curvature,  a  pair  of  long  slender  setae  at  base  of  distal  pair  and  2-3 
short  bristles  at  bases  of  all  major  pairs  (Fig.  4G).  Caudal  appendage  of  six  segments,  each 
with  1-3  spines,  reaching  to  about  Vi  length  of  second  segment  of  pedicel  of  cirrus  VI.  Ter- 
minal segment  with  a  tuft  of  4  long  and  2-3  short,  slender  setae  (Fig.  4H).  Cirral  counts  are 
as  follows: 


II  III  IV  V  VI  Ca 


Styx-7  680905  Sta.  2 


9 

28 

33 

32 

36 

40 

17 

27 

32 

32 

34 

40 

9 

28 

30 

33 

34 

40 

17 

28 

32 

34 

36 

40 

Eight  dwarf  males  were  recovered,  four  from  each  pouch  on  the  inside  of  the  distal  end 
of  the  scutal  plates.  The  males  are  sac-like,  devoid  of  plates  but  covered  with  rows  of  spines. 

Remarks.— This  is  the  second  report  of  A.  hawaiiense  which  was  originally  dredged  off 
Kauai,  Hawaii  at  a  depth  of  1460  m.  Though  Pilsbry  (1907a)  did  not  describe  the  mouth 
parts  and  cirri,  the  capitular  structure  of  our  specimen  agrees  almost  point  for  point  with  the 
description  of  the  type  specimen.  The  bathymetry  also  agrees.  Pilsbry  drew  attention  to  the 
relationship  between  A.  hawaiiense,  A.  rubrum  (Hoek)  and  A.  hirsutum  (Hoek).  In  a  later 
publication  he  (Pilsbry,  1911)  included  these  species  under  the  group  of  Scalpellum  vehiti- 
num.  With  this  we  concur.  However,  a  detailed  comparison  of  ^4.  hawaiiense  from  Allison 
Guyot  with  more  complete  descriptions  of^.  rubrum  (Pilsbry,  1911)  andv4.  hirsutum  (New- 
man and  Ross,  1971)  shows  that  the  resemblance  is  rather  superficial,  there  being  several 
differences  in  the  capitular  structure,  mouth  parts  and  cirri.  ArcoscalpeUum  hawaiiense 
shows  close  resemblance  to  A.  elegantissimum  n.  sp.  While  closely  related,  these  can  be 
distinguished  from  one  another  by  the  following  characters;  A.  hawaiiense  has  1)  the  apex 
of  the  carinal  latus  projecting,  though  slightly,  away  from  the  carina;  2)  close  and  imbricat- 
ing scales  of  the  peduncle;  3)  posterior  ramus  of  cirrus  I,  Wi  times  longer  than  the  anterior 
and  composed  of  nearly  double  the  number  of  segments,  4)  a  caudal  appendage  composed 
of  6  segments  and  reaching  to  about  Vi  the  length  of  second  segment  of  the  pedicel  of  cirrus 
VI,  and  5)  a  much  larger  overall  size. 

ArcoscalpeUum  wyethi  (Cornwall)  from  Guam  appears  to  be  a  related  form,  but  in  this 
species  the  carinal  latera  project  strongly  beyond  the  base  of  carina,  the  scales  on  the  pe- 
duncle do  not  overlap  and  are  widely  spaced,  and  the  intermediate  articles  of  cirrus  VI  sup- 
port 5  pairs  of  setae  instead  of  3  pairs  as  in  ^4.  hawaiiense. 

ArcoscalpeUum  michelottianum  (Seguenza),  1876 
Figures  5  and  1 1  A-B 

Scalpellum  michelotiianum  Seguenza,  1876:381,  pi.  6,  figs.  15-25:  464,  pi.  10,  fig.  26;  ArcoscalpeUum  mich- 
elottianum: Newman  and  Ross,  1971:  71,  pi.  IXB,  text-fig.  34  (see  this  reference  for  complete  synonymy  of  this 
species). 

A/a/ma/.-Styx-7,680901  Sta.  3,  Hess  Guyot  ( 174°24.8'  W;  17°53.2'  N),  1692-1735  m  (Sigsbeebeam  trawl),  1 
spec;  680903-04  Sta.  1,  Allison  Guyot  ( 179°36.0'  W;  18°31.0"  N);  1413-1645  m  (otter  trawl),  2  spec;  680905  Sta.  2, 
Allison  Guvot  (179°37.r  W;  18°35.4'  N),  1413-1449  m  (otter  trawl),  several  spec;  680907  Sta.  4.  Agassiz  Guyot 
(178°  14.2'  W;  17°58.5'  N),  1557  m  (Sigsbee  beam  trawl),  1  spec 

Supplementary  description  (female).— The  large  series  of  specimens  agree  closely  with 


77 


.S.Oinm 


iBEFBH 


.1.0 


_0.5 


.1.0 


Figure  4.  Arcoscalpellum  hawaiiense  (Pilsbry),  Styx-7,  680905.  Sta.  2,  Allison  Guyot.  A,  cirrus  I;  B,  mandible;  C. 
third  tooth  and  inferior  angle  of  mandible;  D,  maxilla  I;  E,  maxilla  II;  F,  palp;  G,  intermediate  articles  of  cirrus 
VI;  H,  caudal  appendage. 

the  descriptions  of  Scalpellum  eximium  Hoek  {=  Arcoscalpellum  michelottianum).  The 
lengthy  synonymy  under /i.  michelottianum  (see  Newman  and  Ross,  1971)  indicates  that 
this  species  is  not  only  variable  but  that  also  several  species  were  confused  with  and  in- 
cluded in  it.  Because  of  this  it  is  important  that  the  specimens  from  the  Mid-Pacific  be 
carefully  characterized,  for  the  synonymy  problem  will  undoubtedly  continue. 

Capitulum  robust,  thick  near  the  peduncle  and  flatter  towards  the  apex;  surface  cov- 
ered by  a  yellow  to  olive  colored  cuticle,  velvety  to  touch,  prominently  hairy  in  young  indi- 
viduals and  sparsely  so  in  older  ones;  14  fully  calcified  plates  usually  fully  approximate, 
but  in  some  specimens  carina  separated  from  others  by  a  narrow  chitinous  interspace 
(Fig.  1  IB).  Scutum  trapeziform;  \Vi-2  times  as  long  as  broad,  divided  into  two  parts  by  a 
faint  diagonal  ridge  running  from  umbo  to  basilateral  angle.  Carina  strongly  bowed,  nar- 
row apically  and  gradually  increasing  in  width  towards  base;  roof  gently  convex  and  trav- 
ersed by  an  indistinct  longitudinal  median  ridge  and  marked  by  V-shaped  growth  lines; 
parietes  well  developed  and  sculptured  with  4-6  distinct  longitudinal  ridges;  base  triangu- 
lar and  enters  as  a  wedge  between  carinal  latera  (Fig.  1 1  A).  Carinal  latus  irregular;  umbo 
at  recurved  apex  which,  in  some  specimens,  is  raised  above  surface  of  capitulum;  plate 
divided  into  three  parts  by  two  ridges  running  from  umbo  to  basal  margin.  Inframedian 
latus  as  high  or  slightly  higher  than  wide;  apex  usually  curved  downwards.  Form  of  plate 


78 


,ABCEGH 

10  mm 

.DiKLM 

■)  n 

.F 

SO 

.1 

Figure  5.  Arcoscalpellum  micheloiiianum  {Seguenzd).  Styx-7,  680901,  Sta.  3,  Hess  Guyot.  A,  palp;  B,  maxilla  I;  C, 
maxilla  11;  D.  mandible;  E,  inferior  angle  and  third  tooth  of  mandible;  F,  cirrus  I;  G,  dwarf  male;  H.  intermediate 
articles  of  cirrus  VI;  I.  caudal  appendage;  J-M,  side  view  of  females. 

changes  considerably  with  growth  (Fig.  5L-M).  Rostral  latus  shorter  than  height  of  infra- 
median  latus;  broad  and  divided  into  two  parts  by  a  ridge  running  from  umbo  to  lateral 
margin.  Rostrum  appearing  externally  in  young  individuals,  lanceolate  in  shape;  umbo 
apical.  With  growth,  rostrum  is  overlapped  by  rostral  latera  of  both  sides  and  becomes 
hidden.  Peduncle  long;  scales  closely  set  or  widely  spaced,  completely  covered  by  a  mem- 


79 


brane  or  partly  projecting  through  it. 

The  measurements  (in  mm)  of  four  dissected  individuals  are  given  below: 


Station  (Styx-7) 

680901 

680903-04 

680903-04 

680907 

Sta.  3 

Sta.  1 

Sta.  1 

Sta.  4 

Overall  height 

53 

58 

22 

50 

Height  of  capitulum 

30 

36 

16 

32 

Height  of  peduncle 

25 

25 

8 

20 

Labrum  buUate;  palp  triangular,  rather  broad  and  short,  superior  and  distal  margins 
covered  with  long  plumose  spines,  inferior  margin  naked  (Fig.  5A).  Maxilla  I  with  cutting 
edge  nearly  straight;  spines  distributed  in  three  indistinct  sets;  upper  margin  supporting  2 
long  stout  and  5  short  spines;  intermediate  set  consisting  of  1  long  and  2  short  spines;  lower 
margin  with  a  set  of  1  long  and  9-1 1  short  spines  (Fig.  5B).  Mandible  with  four  teeth  in- 
cluding inferior  angle;  teeth  spaced  nearly  equidistant  from  one  another  (Fig.  5D);  inferior 
angle  supporting  22-25  bluntly  pointed  teeth  (Fig.  5E).  Maxilla  II  broadly  triangular,  setae 
distributed  in  three  clusters,  those  of  superior  and  distal  lobes  contiguous;  maxillary  lobe 
broad,  short  and  truncate  apically  (Fig.  5C). 

Cirrus  I  widely  separated  from  others;  intermediate  segments  of  anterior  ramus 
strongly  protuberant,  those  of  posterior  ramus  moniliform  and  %  as  wide  (Fig.  5F).  Cirrus  II 
normal,  almost  twice  length  of  cirrus  I.  Cirri  III-VI  subequal  with  equal  or  subequal  rami; 
articular  areas  along  greater  curvature  have  3-4  long  plumose  setae;  lateral  faces  with  2-5 
rows  of  setae;  setation  ctenopod,  three  major  pairs  and  one  minor  pair  along  lesser  curva- 
ture; 2-3  short  setae  between  major  pairs  (Fig.  5H).  Caudal  appendage  of  4  incompletely 
fused  segments,  less  than  height  of  first  segment  of  pedicel  of  cirrus  VI;  distally,  articular 
areas  with  2-4  spines  on  the  outer  margin;  third  segment  with  one  long  and  one  short  seta  on 
distal  margin;  distal  segment  with  a  tuft  of  7-8  long  plumose  setae  (Fig.  51).  Cirral  counts  of 
four  dissected  specimens  follow: 

I  II  III  IV  V  VI  Ca 


Styx-7  680901 

8 

26 

31 

35 

18+ 

17+ 

Sta.  3 

13 

29 

30 

25+ 

20+ 

16+ 

8 

28 

30 

35 

18+ 

18+ 

13 

27+ 

31 

26+ 

18+ 

19+ 

Styx-7  680903-04 

8 

26 

30 

35 

19+ 

18+ 

Sta.  1  (spec.  1) 

13 

30 

31 

25+ 

22+ 

18+ 

8 

28 

30 

35 

36 

15+ 

13 

31 

31 

36 

40 

13+ 

Styx-7  680903-04 

8 

18 

23 

25 

25 

25 

Sta.  1  (spec.  2) 

11 

21 

22 

24 

23 

23 

8 

19 

23 

20 

24 

25 

11 

19 

22 

21 

25 

24 

Styx-7  680907 

7 

28 

35 

36 

34 

35 

Sta.  4 

13 

27 

28 

32 

35 

33 

7 

29 

32 

31 

37 

35 

13 

27 

32 

34 

35 

35 

tt:,  4 


:n  2 


Dwarf  males  recovered  from  three  large  specimens;  as  many  as  5-9  in  a  pouch  on  the 
inner  sides  of  the  scuta  (Fig.  5G).  They  resemble  those  figured  for  S.  eximium  by  Hoek 
(1883,  pi.  9.  fig.  10). 

Remarks.— Tht  Mid-Pacific  specimens  agree  closely  with  Hoek's  description  of  Scal- 
pellum  eximium  and  the  resemblance  is  particularly  striking  with  regard  to  the  character- 
istic shape  of  the  dwarf  males. 

The  specimens  from  the  Mid-Pacific  differ  from  the  examples  of  Newman  and  Ross 
( 197 1 ).  The  latter  have  a  supramedian  notch  in  the  cutting  edge  of  Maxilla  I  where  as  none 
is  apparent  in  the  present  specimens;  the  intermediate  segments  of  cirrus  VI  have  4  pairs  of 
setae  in  Pacific  specimens  as  opposed  to  3  pairs  in  the  North  Atlantic  individuals.  The  Elta- 


80 


nin  specimens,  which  are  relatively  small,  came  from  depths  exceeding  3000  meters  while 
the  specimens  from  the  Mid-Pacific  were  dredged  at  nearly  half  that  depth  and  are  large. 
However,  while  it  is  possible  that  allometry  and  bathymetry  account  for  the  observed 
differences,  it  is  also  possible  that  the  differences  are  genetic.  There  are  presently  in- 
sufficient data  to  resolve  this  problem. 

Arcoscalpellum  radiatum  n.  sp. 

Figure  6 

A/a/ma/.-Styx-7,  680903  Sta.  1   Allison  Guyot  ( 179°36.0' W,  18°31.0'N),  1413-1645  m  (otter  trawl).  2  spec. 

Depository.-V.S.NM.  no.  140947  (Holotype.  Styx-7.  680903  Sta.  1);  U.S.N.M.no.  140948  (Paratype,  Styx-7, 
680903.  Sta.  1). 

Diagnosis.— Capitulum  with  14  fully  calcified  approximate  plates  sculptured  with 
prominent  radial  ribs  emanating  from  the  umbones.  Carinal  latera  interdigitate  at  base  of 
carina.  Carinal  roof  flat,  parietes  well  developed.  Rostrum  exposed,  elongate  triangular. 
Mandible  with  4  teeth  including  a  strongly  denticulate  inferior  angle.  Maxilla  I  with  a  deep 
medial  notch  in  cutting  edge.  Caudal  appendage  uniarticulate  and  much  shorter  than  first 
segment  of  pedicel  of  cirrus  VI. 

Description  (female).— Capitulum  elongate,  oval,  almost  twice  as  long  as  broad;  occlu- 
dent  and  carinal  margins  moderately  arched,  covered  with  long  hairs;  14  fully  calcified  ap- 
proximate plates  sculptured  with  prominent,  evenly  spaced  ribs  which  extend  from  um- 
bones to  basal  margins;  ribs  intercepted  by  feeble  lines  of  growth  (Fig.  6A).  Scutum 
subquadrate;  twice  as  long  as  broad  and  broadest  in  the  middle;  occludent  and  carinal  mar- 
gins subparallel,  the  latter  %  as  long;  surface  slightly  convex  and  traversed  by  ribs  emanat- 
ing from  region  of  umbo;  ribs  more  conspicuous  in  lower  half  of  plate;  apical  umbo  partly 
overlapping  occludent  margin  of  tergum.  Tergum  triangular,  sculptured  with  longitudinal 
ribs  except  for  a  narrow  carinal  portion.  Upper  latus  appears  triangular  but  is  four  sided;  a 
faint  diagonal  angulation  runs  from  umbo  to  carinolateral  angle.  Carina  with  broad  base 
enclosed  between  carinal  latera;  roof  flat  and  marked  with  broad  'U'  shaped  lines  of 
growth;  parietes  well  developed,  smooth  (Fig.  6B).  Carinal  latus  higher  than  wide,  lateral 
margin  long  and  partly  overlapped  by  inframedian  latus;  ribs  radiate  from  umbo;  while  not 
shown  in  figure,  carinal  margins  broadly  interdigitating  (Fig.  6B-C).  Inframedian  latus 
more  than  4  times  as  long  as  broad;  traversed  by  transverse  striae;  umbo  at  truncate  apex. 
Rostral  latus  trapeziform;  divided  into  two  unequal  triangular  areas  by  a  faint  ridge  that 
runs  from  umbo  to  basilateral  angle.  Rostrum  well  developed,  triangular,  broad  above  and 
pointed  below  (Fig.  6D).  Peduncle  short,  covered  with  6  rows  of  strong  scales  with  pro- 
jecting edges. 

Labrum  bullate;  crest  armed  with  21  V-shaped  pointed  teeth  (Fig.  6H).  Palp  long  and 
narrow;  proximal  superior  and  distal  margins  covered  with  spines  (Fig.  61).  Mandible  with 
4  teeth  including  the  inferior  angle;  first  tooth  well  separated  from  second  (Fig.  6E);  inferior 
angle  strongly  denticulate  and  armed  with  8  pointed  teeth  (Fig.  6F).  Maxilla  I  with  a  deep 
notch  in  middle  of  cutting  edge;  2  long  and  2  short  spines  above  notch  and  2  long  and  2-3 
short  spines  below  notch;  surface  covered  with  long  setae  (Fig.  6G).  Maxilla  II  triangular 
and  covered  with  a  few  marginal  setae  on  superior,  distal  and  inferior  margins. 

Cirrus  I  separated  from  remaining  cirri;  posterior  ramus  slightly  longer;  segments 
moniliform;  covered  with  long  plumose  setae  (Fig.  6J).  Cirrus  II  not  modified.  Cirri  III-VI 
essentially  equal  in  length  with  subequal  rami.  Articular  areas  along  greater  curvature  with 
1-2  long  thin  setae;  interarticular  areas  and  lateral  faces  naked;  setation  ctenopod;  2  major 
and  1  minor  pair  of  setae  along  lesser  curvature;  a  few  short  bristles  at  bases  of  these  (Fig. 
6K).  Caudal  appendage  uniarticulate;  shorter  than  first  segment  of  pedicel  of  cirrus  VI;  an- 
terior and  posterior  borders  free  of  setae;  4-5  setae  distally.  Cirral  counts  of  the  holotype 
follow: 

I  II  III  rv  V  VI         Ca 


Styx-7  680903 

7 

11 

14 

15 

16 

17 

Sta.  1 
(Holotype) 

8 

7 

12 
12 

13 

14 

15 
15 

16 
15 

16 
15 

8 

10 

14 

14 

16 

17 

7^  1 


81 


O  M    V 


,ltBCD 

2.0mm 

,EGHI 

1  n 

■  F 

05 

JK 

OS 


T^;^ 


Figure  6.  Arcoscalpelliim  radiatum  n.  sp.,  Holotype,  Styx-7,  680903,  Sta.  I,  Allison  Guyot.  A,  side  views  of  fe- 
male; B,  carina!  view;  C,  base  of  carina  and  abutment  of  carinal  latera;  D.  rostrum  and  adjoining  plates;  E, 
mandible;  F,  third  tooth  and  inferior  angle  of  mandible;  G,  maxilla  I;  H,  crest  of  labrum;  I,  palp;  J,  cirrus  I;  K, 
intermediate  articles  of  cirrus  VI. 


Remarks.— Arcoscalpelliim  radiatum  is  related  most  closely  \o  A.  pacificum  ( Pilsbry),  A. 
c/2/7/eme(Pilsbry)  (new  name  for  A.  ^raf/7<?  (Pilsbry)  and /I.  semisculptum  (Pilsbry).  Charac- 
ters in  common  are  an  elongate  capitulum,  a  narrow  and  elongate  inframedian  latus  with 
an  apical  umbo  and  shorter  than  the  carinal  latus,  the  presence  of  a  narrow  and  elongate 
rostrum  and  the  possession  of  carinal  latera  which  are  higher  than  wide  and  interdigitating 
where  they  meet.  The  ornamentation  of  the  capitular  plates  o{  A.  radiatum  recalls  the  con- 
dition in  A.  pacificum.  However,  the  rostral  latera  of  A.  pacificum  are  wider  than  high,  the 


82 


umbones  of  the  carinal  latera  are  at  the  lower  '/4  of  the  carinal  margin  (Pilsbry,  1907a),  the 
cutting  edge  of  maxilla  I  is  nearly  straight  and  the  caudal  appendage  is  five-segmented 
(Annandale,  1913).  Arcoscalpellum  radiatum  differs  from  A.  pacificum  in  all  these  charac- 
ters. It  differs  from  A.  chiliense  in  the  possession  of  longitudinal  ribs  on  the  terga  and  scuta, 
in  the  carinal  roof  being  flat  rather  than  convex,  and  in  the  inframedian  latus  which  is  pro- 
portionately much  wider  and  decidedly  higher  than  the  adjoining  rostral  latus.  Arcoscal- 
pellum semisculptum  also  has  an  inframedian  latus  which  is  much  narrower  than  in  A. 
radiatum,  but  in  this  species  the  umbones  of  the  carinal  latera  are  placed  at  the  lower  'Z?  of 
the  carinal  margin  as  opposed  to  their  distinctly  medial  position  in  the  new  species.  The 
type  o^  A.  semisculptum  came  from  a  depth  of  512  meters  which  is  nearly  one-third  the 
depth  from  which  the  Mid-Pacific  specimens  were  taken.  Broch  ( 1953)  recorded  one  speci- 
men from  a  depth  of  1484  meters,  comparable  to  the  Pacific  station.  Unfortunately  neither 
Pilsbry  (1907c)  nor  Broch  gave  any  details  of  the  arthropodal  structures  of  this  species. 
Also  present  at  the  same  station  is  a  small  individual  which  has  not  yet  developed  the 
radial  sculpture,  but  is  in  all  other  respects  similar  to  the  one  described  above. 

Arcoscalpellum  rossi  n.  sp. 

Figure  7 

Ma/mfl/.-Styx-7,  680901  Sta.  3,  Hess  Guyot  (174°24.8' W,  17°53.2'N),  1692-1735  m  (Sigsbee  beam  trawl).  1 
spec.  Styx-7,  680903-04  Sta.  1,  Allison  Guyot  (' 179°36.0'  W,  18°31.0'  N),  1413-1645  m  (otter  trawl),  2  spec. 

Depositorv.-XJ.S.fiM.no.  140949  (Holotype,  Styx-7,  680901,  Sta.  1)  U.S.N. M.  no.  140950 (Paratypes,  Styx-7, 
680903-04  Sta.  1,2  spec). 

Diagnosis  (female).— Capitulum  long  and  narrow,  composed  of  14  fully  calcified 
plates.  Roof  of  carina  flat,  parietes  well  developed,  especially  towards  distal  half  of  plate. 
Rostrum  large,  ovotriangular  and  fully  exposed.  Maxilla  I  with  notch  in  middle  of  cutting 
edge.  Mandible  with  four  teeth  including  inferior  angle;  upper  margin  of  third  tooth  ser- 
rated. Caudal  appendage  of  4  segments  and  reaching  to  ^4  height  of  first  segment  of  pedicel 
of  cirrus  VI. 

Description  (female).— Capitulum  long  and  narrow,  composed  of  14  fully  calcified 
plates  and  sparsely  covered  with  hairs.  Plates  separated  by  narrow  chitinous  interspaces 
and  marked  with  faint  lines  of  growth.  Occludent  margin  strongly  convex;  carinal  margin 
irregularly  straight;  apex  slightly  retroverted  towards  the  carinal  side  (Fig.  7A). 

Tergum  triangular,  occludent  margin  short  and  convex,  scutal  and  basal  margins  al- 
most straight,  carinal  margin  concave  for  %  the  distance  towards  the  carinal  angle  and 
straight  thereafter.  Scutum  more  than  twice  as  long  as  broad;  lateral  margin  sinuate  just 
below  tergolateral  angle;  apex  of  upper  latus  projects  towards  this  sinuous  part;  umbo  api- 
cal, overlapping  occludent  margin  of  tergum.  Upper  latus  appearing  triangular  but  five 
sided.  Carinal  latus  fully  twice  as  long  as  broad;  carinal  margin  curving  out  at  base  of  ca- 
rina, beyond  which  umbones  bluntly  project.  Carinal  latera  meet  and  surround  base  of  ca- 
rina in  form  of  a  broad  'V  and  do  not  interdigitate  (Fig.  7C).  Carina  long  and  simply 
bowed;  roof  flat;  parietes  well  developed  towards  distal  half  of  plate  (Fig.  7B).  Inframedian 
latus  rectangular,  more  than  four  times  as  long  as  broad,  umbo  submedial  in  position, 
slightly  displaced  towards  distal  half  and  slightly  raised  above  surface  of  plate.  Rostral 
latus  nearly  rectangular  in  outline,  with  parallel  but  unequal  scutal  and  basal  margins  and 
sub-parallel  lateral  margins.  Rostrum  large,  fully  exposed,  elongate  triangular,  broad 
above  and  pointed  below  (Fig.  7D).  Peduncle  short,  bent  at  right  angles  to  capitulum  and 
covered  with  6-8  rows  of  narrow  elongate  plates  with  chitinous  interspaces. 

Labrum  bullate;  crest  armed  with  22  teeth.  Palp  narrow  and  elongate;  superior  and 
anterior  margins  armed  with  a  few  spines;  inferior  margin  with  proximal  short  stout  spine 
(Fig.  7E).  Maxilla  I  with  a  well  defined  notch  in  middle  of  cutting  edge,  2  long  and  1-2 
short  stout  spines  above  and  one  long  and  3-5  short  spines  below  notch  (Figs.  7H,  I).  Max- 
illa II  with  3  well  defined  lobes;  marginal  setae  long  and  setulose;  setae  distributed  in  3 
clusters,  those  of  inferior  margin  being  segregated;  lateral  margins  sparsely  setose;  max- 
illary lobe  moderately  long  and  cylindrical  (Fig.  7J).  Mandible  with  4  teeth  including  infe- 
rior angle;  second  tooth  twice  the  distance  from  the  first  than  from  the  third  tooth;  upper 
margin  of  third  tooth  serrate  (Fig.  7F);  inferior  angle  supporting  8  long,  narrow  and 
pointed  teeth  (Fig.  7G). 


83 


.AC 

2.0 

2.0 

1.0 

0.2 

,FI) 

,eH 

0.1 

0-5 

0.2 


05 


Figure  7.  Arcoscalpellum  rossi  n.  sp.,  Holotype,  Styx-7.  680901,  Sta.  3,  Hess  Guyot.  A,  right  side  view  of  female; 
B,  carina!  view;  C,  carinal  latera;  D,  rostrum;  E,  crest  of  labrum  and  left  palp;  F.  mandible;  G.  third  tooth  and 
inferior  angle  of  mandible;  H-I,  maxillae  I;  J.  maxilla  II;  K,  intermediate  articles  of  cirrus  VI;  L,  cirrus  I;  M, 
caudal  appendage. 


Cirrus  I  (Fig.  7L)  separated  from  the  rest:  cirrus  II  normal;  articular  areas  along 
greater  curvature  of  cirrus  VI  with  one  sharp  spine;  interarticular  areas  faintly  serrated  with 
5-6  spines;  lateral  faces  devoid  of  setae.  Setation  ctenopod;  2  major  and  1  minor  pair  along 


84 


lesser  curvature.  Caudal  appendage  composed  of  4  segments;  reaching  to  %  length  of  first 
segment  of  pedicel  of  cirrus  VI;  distal  segment  with  2  long  and  1  short  setae  (Fig.  7M).  Cir- 
ral  counts  follow: 

I  II  III  IV  V  VI  Ca 


Styx-7,  680901 

7 

12 

15 

18 

18 

18 

Sta.  3 

8 

13 

18 

18 

19 

19 

(Holotype) 

1  + 

12 

17 

19 

19 

20 

8 

14 

15 

17 

19 

18+ 

tt:  4 


Remarks.— Arcoscalpellum  rossi  is  related  ioA.flavum  (Hoek,  1883: 127),  yl.  novae-Zea- 
landeae {Hoek,  1883: 124),^.  a^v^^/co/a  (Hoek,  1883:114),/J.  m/>7w/wm  (Hoek,  1883:113),^!. 
perlongum  (Pilsbry,  1907b:  198),  A.  albatrossianum  (Pilsbry,  1907c:54).  Characters  that 
unite  all  these  species  are:  1)  the  capitulum  is  elongate  and  narrow,  the  lower  whorl  of 
latera  contributing  in  part  to  its  lengthening;  2)  a  long  and  narrow  inframedian  latus  with 
an  umbo  that  is  medial  to  basal. 

With  the  exception  o^  A.  rossi  all  these  species  have  an  inframedian  latus  which  is  ei- 
ther hour-glass  shaped  or  has  at  least  a  narrow  constriction  in  the  middle.  Arcoscalpellum 
rossi  can  be  separated  readily  from  these  by  its  rectangular  inframedian  latus  which  is  not 
at  all  constricted.  Further,  it  has  a  well  developed  rostrum  that  is  fully  exposed  whereas  in 
the  others  a  rostrum  has  not  been  described  or,  if  present,  is  of  a  smaller  size. 

In  the  relative  proportions  of  the  capitular  plates  and  in  the  general  nature  of  the  ar- 
thropodal  structures  A.  rossi  shows  a  close  resemblance  to  A.  albatrossianum  and  A.  per- 
longum. All  three  species  have  a  mandible  in  which  the  upper  margin  of  the  third  tooth  is 
serrate  and  maxilla  I  has  a  deep  notch  in  the  middle  of  the  cutting  edge  (Nilsson-Cantell, 
1925;  MacDonald,  1929).  However,  in  A.  albatrossianum  and  A.  perlongum  the  caudal  ap- 
pendages reach  beyond  the  pedicel  of  cirrus  VI  and  respectively  have  8  and  6  segments 
whereas  in  A.  rossi  the  caudal  appendage  is  shorter  than  the  first  segment  of  the  pedicel  of 
cirrus  VI  and  has  4  segments. 

The  species  is  named  for  Arnold  Ross,  Natural  History  Museum,  San  Diego,  student  of 
barnacles,  and  friend. 

1  Arcoscalpellum  sp. 
Figure  1  lE-F 

Material.-Sly\-1,  680910  Sta.  5.  Sio  Guyot  (171°05.r  E,  18°17.7  N).  1692  m  (pipe  dredge),  broken  shells. 

The  shell  fragments  from  Sio  Guyot,  while  undoubtedly  belonging  to  a  scalpellid, 
are  too  incomplete  to  allow  positive  identification  and  are  tentatively  assigned  to  Arcoscal- 
pellum on  the  basis  of  a  carina  (Fig.  1  IF)  and  a  scutum  (Fig.  1  IE). 

Genus  Mesoscalpellum  Hoek,  1907 

Mesoscalpellum  gruvelii  (Annandale),  1906 

Figures  8  and  1 IH-I 

Scalpellum  gruvelii  Annandale  1906b:390;  1906a:  141,  text-fig.  4;  1907-1908,  pi.  1,  fig.  1.  pi.  11,  figs.  1,  la,  3; 
1913:232;  Scalpellum  gruvelii  var.  quadralum  Annandale,  1906b:391;  1907,  pi.  II,  fig.  3:  Annandaleum  gruvelii; 
Newman  and  Ross,  1971 :  122;  Scalpellum  chitinosum  Hoek.  1907:73  pl.  VII,  fig.  4;  Scalpellum  imperfectum  Pilsbry, 
1907c:75.  pl.  IV,  figs.  15-18,  text-fig.  30;  Barnard,  1924:46;  1925:3;  MacDonald,  1929:537,  pl.  2,  fig.  3;  Broch, 
1953:9;  Stubbings,  1961:11,  fig.  2;  Zevina;  1969:67;  Mesoscalpellum  imperfectum:  Newman  and  Ross,  1971:119; 
fig.  62. 

Malerial.~Slyx-l.  680903-04  Sta.  1.  Allison  Guyot  ( 179°36.0'  W,  18°31.0'  N),  1429-1663  m  (otter  trawl),  1 
spec;  680905,  Allison  Guyot  ( 179°37.r  W,  18°35.4'  N),  1449-1557  m  (otter  trawl),  several  spec. 

Supplementary  description  (female).— There  is  considerable  variation  in  the  external 
morphology  of  the  large  series  of  specimens  from  the  Mid-Pacific  (see  fig.  8A-C,  1 IH-I). 
However,  specimens  comparable  in  size  to  Annandale's  types  appear  identical  with  his  de- 
scriptions. Some  clarification  is  needed  as  regards  the  vase-shaped  nature  of  the  in- 
framedian latus,  supposedly  characteristic  of  the  genus  Annandaleum  (Newman  and  Ross, 
1971).  In  A.  gruvelii,  both  in  the  original  description  and  in  several  of  the  specimens  in  the 
present  collection  the  outline  of  this  plate  has  the  shape  of  an  hourglass.  A  club-shaped 
ridge,  with  its  expanded  extremity,  projects  outwards,  and  it  is  this  ridge  that  gives  the  plate 
its  vase-like  appearance,  especially  when  seen  through  the  semi-transparent  membrane. 


85 


.^>>%.-^ 


,*c 

,B 

4n 

,D 

in 

,EN 

in 

JIIM 

ni 

,Sl 

ns 

,H 

D1 

,R                                           7 

2.0 


Figure  8.  Mesoscalpellum  gruvelii  (Annandale),  Styx-7,  680903-04  Sta.  1,  Allison  Guyot  (A  and  D-N);  Styx-7, 
680905,  Sta.  2.  Allison  Guvot  (B-C).  A-C,  side  views  of  females;  D,  male  cyprid;  E.  crest  of  labrum;  F,  palp:  G, 
mandible;  H,  third  tooth  and  inferior  angle  of  mandible;  I.  maxilla  I;  J,  maxilla  II;  K,  cirrus  I;  L.  intermediate 
article  of  outer  ramus  of  cirrus  VI;  M,  intermediate  article  of  inner  ramus  of  cirrus  VI;  N,  caudal  appendage. 


The  bearing  this  problem  has  on  the  distribution  of  the  genus  will  be  taken  up  below,  under 
remarks. 

Annandale's  descriptions  of  mouth  parts  and  cirri  are  incomplete  and  are  elaborated 
on  as  follows.  Labrum  long,  triangular  rather  than  buUate;  crest  armed  with  40-50  bluntly 


86 


pointed  teeth  (Fig.  8E).  Palp  elongate,  bluntly  triangular  distally;  superior  proximal  margin 
with  a  few  stout  spines;  distal  extremity  strongly  spinose,  lateral  faces  with  a  few  spines 
(Fig.  8F).  Maxilla  I  with  straight  cutting  edge;  upper  margin  with  1  short  and,2  long  spines; 
lower  margin  supporting  2  long  and  4  short,  stout  spines  (Fig.  81).  Mandible  with  4  teeth 
including  a  slightly  receding  inferior  angle;  first  tooth  well  separated  from  second;  third 
tooth  proximal  to  inferior  angle  (Fig.  8G);  inferior  angle  supporting  12-13  moderately  long, 
somewhat  pointed  teeth  (Fig.  8H).  Maxilla  II  triangular,  lobes  feebly  developed;  marginal 
setae  distributed  in  3  clusters;  lateral  faces  setose;  maxillary  lobe  elongate,  broad  near  the 
base  and  narrow  apically  (Fig.  8J). 

The  cirri,  as  noted  by  Annandale,  are  devoid  of  pigment.  Cirrus  I  widely  separated 
from  the  rest;  intermediate  segments  of  anterior  ramus  protuberant,  those  of  posterior 
ramus  cyhndrical  and  Va  as  wide  (Fig.  8K).  Cirri  II-VI  increasing  progressively  in  length  and 
with  equal  or  subequal  rami.  Greater  curvatures  of  cirri  II-VI  with  2-3  rows  of  stiff  bristles; 
lateral  faces  with  1-4  rows  of  setae;  articular  areas  with  a  cluster  of  3-5  setae;  interarticular 
areas  with  1-4  setae.  Distal  cluster  of  setae  along  lesser  curvature  of  intermediate  segments 
hypolasiopod  in  outer  ramus  and  ctenopod  (3  major  pairs  and  I  minor  pair)  in  inner  ramus; 
2-3  pairs  at  bases  of  all  major  setae  (Figs.  8L,  M).  Caudal  appendages  as  long  as  pedicels  of 
cirrus  VI;  each  consisting  of  8  segments;  distal  segment  supporting  6  long  setae  of  equal 
length  (Fig.  8N).  Cirral  counts  of  one  specimen  are: 

I  II  III  IV  V  VI  Ca 


Styx-7,  680903-04 
Sta.  1 

8 

11 

19 

22 

23 
24 

27 
28 

28 
26 

30 
28 

8 

20 

24 

26 

28 

29 

10 

22 

26 

26 

28 

29 

A  male  cyprid  was  found  in  a  pouch  on  the  inner  side  of  the  right  scutum  of  one  speci- 
men. It  resembles  (Fig. 8D)  the  male  cyprid  of  5".  gruvelii  (  =  M.  gruvelii)  by  Stewart  (1911). 

Remarks.—Several  authors  expressed  concern  over  the  similarities  between  Annanda- 
leiim  gruvelii  (Annandale),  Mesoscalpellum  imperfectum  (Pilsbry),  and  M.  sanctaebarbarae 
(Pilsbry),  (see  Pilsbry  1907c;  Annandale,  1913;  Barnard,  1924;  Stubbings,  1961;  Newman 
and  Ross,  1971).  This  and  related  problems  need  clarification  here.  The  first  is  with  regard 
to  the  inclusion  of  gruvelii,  in  the  genus  Annandaleum  proposed  by  Newman  and  Ross 
(1971)  for  the  reception  of  this  and  three  other  Indo-West  Pacific  species.  The  most  diagnos- 
tic characteristic  of  Annandaleum  is  the  large  and  vase-shaped  inframedian  latus,  which  as 
far  as  can  be  judged  from  published  figures,  is  present  in  A.  japonicum,  A.  lambda  and  A. 
flavum.  In  A.  gruvelii,  it  is  not  well  developed  and  the  inclusion  of  this  species  '\n  Annanda- 
leum greatly  weakens  the  definition  of  this  genus.  Actually /I.  gruvelii  has  a  facies  similarity 
with  members  of  the  genus  Mesoscalpellum  and  we  propose  that  it  be  transferred  to  this 
genus.  This  greatly  sharpens  the  distinction  between  the  two  genera. 

The  second  problem  concerns  M.  gruvelii  and  M.  imperfectum.  Several  specimens  from 
the  Mid-Pacific  collection  show  a  point  to  point  similarity  with  the  description  of  M.  gruvelii 
(Annandale,  1906b)  and  there  is  no  doubt  that  the  specimens  before  us  belong  to  this  spe- 
cies. However,  the  arthropodal  structures  of  the  Mid-Pacific  specimens  are  almost  identical 
with  those  of  the  paratypes  of  M.  imperfectum  figured  by  Newman  and  Ross  (1971,  text-fig. 
62).  Therefore  we  believe  that  imperfectum  and  gruvelii  are  the  same  species,  a  synonymy 
that  has  been  suggested  before. 

This  leaves  us  the  question  as  to  the  status  of  M.  sanctaebarbarae.  Newman  and  Ross 
pointed  out  several  diff'erences  in  the  anatomy  of  A/,  imperfectum  (  =  M.  gruvelii)  and  M. 
sanctaebarbarae.  We  have  had  the  opportunity  to  examine  closely  the  soft  parts  of  the  latter 
species  from  the  San  Diego  Trough,  which  confirm  that  the  diff'erences  are  consistent,  with 
one  exception.  The  setation  of  the  distal  cluster  of  the  intermediate  articles  of  the  outer 
rami  of  cirrus  VI  is  said  to  be  ctenopod  in  the  paratypes  whereas  it  is  distinctly  hypolasio- 
pod in  the  specimen  from  the  San  Diego  Trough.  The  importance  of  this  diff'erence  remains 
to  be  determined  and  in  the  light  of  other  dilTerences  we  are  inclined  to  continue  to  recog- 
nize the  two  species. 


87 


With  the  transfer  of  gruvelii  to  Mesoscalpellum  and  the  recognition  of  A/,  imperfectum 
as  a  synonym  of  M.  gruvelii,  the  genus  and  the  species  take  on  a  world  wide  distribution 
(Indian,  Atlantic  and  Pacific  oceans). 

Family  Poecilasmatidae  Annandale,  1909 

Genus  Megalasma  Pilsbry,  1907c 

Subgenus  Glyptelasma  Pilsbry,  1907c 

Megalasma  (Glvptelasma)  pilsbryi  Caiman,  1919 

Figure  9 

Megalasma  (Glvptelasma)  pilsbryi  Caiman,  1919:363.  fig.  lA-C.  fig.  2;  Nilsson-Cantell,  1928:20,  fig.  9A-E; 
1938:10. 

MaleriaL^Sxy\-l.  680905  Sta.  2,  Allison  Guyot  ( 179°37.r  W.  18°35.4'  N),  1445-1557  m  (otter  trawl).  10  spec, 
all  attached  to  Anoscalpellum  michelollianum  and  Mesoscalpellum  gruvelii.. 

5'i//7/7/£'me/7/arvfife5cr//7//6>n.— The  external  morphology  ofour  Specimens  agrees  with  the 
description  given  by  Caiman  (1919).  The  base  of  the  carina  is  produced  into  two  teeth  on 
the  inner  side  (Fig.  9C).  In  the  present  specimens  the  basal  margin  of  the  scutum  and  carina 
meet  at  an  angle  of  more  than  90°  whereas  in  Caiman's  specimens  these  are  shown  to  meet 
at  right  angles.  This  character  varies  with  growth. 

Nilsson-Cantell  (1928)  gave  brief  descriptions  of  the  mouth  parts.  More  detail  of  the 
trophi  and  cirri  is  in  order.  Labrum  buUate,  slightly  broader  than  long  and  bluntly  triangu- 
lar anteriorly;  anterior  margin  and  surface  covered  with  tufts  of  2-6  fine,  short  hairs;  crest 
armed  with  35  small,  stout  and  somewhat  pointed  teeth  (Fig.  9F).  Palp  broad  proximally 
and  bluntly  conical  distally;  superior  margin  free  of  setae;  inferior  and  distal  margins  bor- 
dered by  long  plumose  setae;  lateral  faces  setose  (Fig.  9F).  Mandible  with  5  teeth  including 
inferior  angle;  upper  margin  of  fourth  tooth  serrate;  surface  profusely  covered  with  long 
thin  spinules  some  of  which  cross  the  cutting  edge;  superior  and  inferior  margins  bordered 
by  short,  thin  hairs  along  the  entire  length  (Fig.  9G);  inferior  angle  tridentate,  the  teeth 
being  short  and  pointed  (Fig.  9H).  Maxilla  I  with  cutting  edge  concave  above  and  strongly 
convex  below,  without  a  well  defined  notch,  2  long  and  1  short  spine  above;  the  convex 
lower  portion  supports  a  set  of  14-15  short  and  long  spines,  and  superior  and  inferior  mar- 
gins as  well  as  surface  covered  with  short  hairs  (Fig.  91).  Maxilla  II  triangular,  higher  than 
wide;  superior  lobe  well  developed,  distal  and  inferior  lobes  feebly  so;  marginal  setae  dis- 
tributed in  three  clusters,  those  of  superior  and  distal  lobes  separated  by  a  naked  superior 
margin  (Fig.  9J). 

Cirrus  I  widely  separated  from  cirrus  II;  anterior  and  posterior  rami  equal  in  length 
and  composed  of  9  and  10  segments  respectively;  segments  of  anterior  ramus  l'/4-iy2  times 
broader  than  those  of  posterior  (Fig.  9E).  Cirrus  II,  Wi  times  longer  than  cirrus  I;  cirri  III- 
VI  equal  in  length  with  equal  rami  and  composed  of  a  rather  constant  number  of  segments. 
Articular  areas  along  greater  curvatures  with  2-3  long  and  1-2  short  setae;  interarticular 
areas  and  lateral  faces  devoid  of  setae;  setation  ctenopod;  3  major  pairs  and  1  minor  pair 
along  lesser  curvature;  1-3  short  bristles  at  bases  of  major  pairs  (Fig.  9K).  Caudal  appen- 
dage uniarticulate,  short,  about  Vy  height  of  first  segment  of  pedicel  of  cirrus  VI;  anterior 
and  posterior  margins  bordered  with  small  and  inconspicuous  spinules;  distal  end  broad, 
with  8  short  to  long  plumose  setae  (Fig.  9L).  Penis  large,  proximally  broad,  gradually  taper- 
ing to  a  blunt  apex;  surface  covered  with  long  thin  setae  which  are  sparsely  distributed  for  a 
greater  length  of  the  organ  but  are  more  profuse  and  conspicuous  towards  the  distal  end.  A 
single  pair  of  rather  short,  slender  filamentary  appendages  are  present  on  dorsum  of  pro- 
soma  near  its  posterior  margin  (Fig.  9M),  as  described  and  figured  by  Caiman  (1919).  Cirral 
counts  of  the  dissected  specimen  follow; 

I  II  III  IV  V  VI  Ca 


Styx-7,  680905 

9 

15 

19 

18 

19 

19 

Sta.  2 

10 

16 

19 

19 

19 

20 

9 

17 

19 

19 

19 

20 

10 

16 

19 

19 

19 

18 

88 


<AD 

50 

|B 

10 

.CM 

2  0 

,E 

05 

iFGIJKL 

0 

,H 

02 

Figure  9.  Me^alasma  (Glvptelasma) pilsbryi  Caiman,  Styx-7, 680905,  Sta.  2,  Allison  Guyot.  A-B,  right  side  view  of 
hermaphrodites;  C,  inner  view  of  disarticulated  shells;  D,  outer  view  of  carina;  E,  cirrus  I;  F,  labrum  and  palps;  G. 
mandible;  H,  third  and  fourth  teeth  and  inferior  angle  of  mandible;  I,  maxilla  I;  J,  maxilla  II;  K,  intermediate 
articles  of  cirrus  VI;  L,  caudal  appendage;  M,  prosoma  and  filamentary  appendages. 

Remarks.— ?'\hhxy  ( 1 907c,d)  and  Caiman  (1918b,  1919)  discussed  the  status  and  defini- 
tions of  the  genus  Megalasma  and  its  subgenera  Megalasma  s.s.  and  Glvptelasma.  The  speci- 
mens from  the  Mid-Pacific  are  clearly  referable  to  Glvptelasma  because  the  basal  margin  of 


89 


the  scutum  forms  a  distinct  angle  with  the  occludent  margin,  and  also  by  the  weak  sculpture 
of  the  plates.  The  configuration  and  relative  proportions  of  the  capitular  plates  and  the  gen- 
eral structure  of  the  trophi  and  cirri  are  in  agreement  with  the  description  of  Megalasma 
(Glvptelasma) pilsbrvi.  The  bathymetry  is  also  similar.  The  figures  and  descriptions  given  by 
Nilsson-Cantell  show  that  the  crest  of  the  labrum  has  about  50  teeth,  the  rami  of  cirrus  I 
have  9  and  1 1  segments  and  the  posterior  cirri  have  23-24  segments.  In  contrast,  the  crest  of 
the  labrum  of  the  Mid-Pacific  specimen  supports  35  teeth,  the  rami  of  cirrus  I  have  9  and  10 
segments  respectively  and  the  posterior  cirri  are  composed  of  18-20  segments.  It  is  likely 
that  allometry  may  account  for  these  differences,  Nilsson-Cantell's  specimens  being  larger 
(capitular  height:  20  mm)  than  the  Mid-Pacific  example  (capitular  height:  1 1  mm). 

Megalasma  pilsbrvi  is  closely  related  to  M.  annandalei  Pilsbry.  Caiman  (1919)  and 
Barnard  (1925)  recognized  this,  but  both  authors  advocated  their  retention  as  good  species. 
Caiman  stated  that  A/,  pilsbrvi  diflfers  from  M.  annandalei  "in  having  no  sudden  widening  of 
the  sides  of  the  carina  and  no  excavation  of  the  adjacent  sides  of  the  scutum,  as  well  as  in  the 
thick  cuticle  covering  the  valves  ..."  Furthermore  the  intermediate  segments  of  cirrus  VI  of 
M.  pilsbrvi  support  3  major  pairs  and  one  minor  pair  of  setae  along  the  lesser  curvature 
whereas  in  M.  annandalei  there  are  4  major  pairs  and  1  minor  pair  (Pilsbry,  1907,  pi.  V, 
Fig.  14). 

The  Mid-Pacific  specimens  were  found  attached  to  Arcoscalpellum  michelottianum  and 
Mesoscalpellum  gruvelii.  Nilsson-Cantell  (1928)  collected  this  species  from  Scalpellum  ve- 
lutinum  (  =  A.  michelottianum)  and  S.  alcockianum  (  =  A.  alcockianum). 

Suborder  Verrucomorpha  Pilsbry,  1916 

Family  Verrucidae  Darwin,  1854 

Genus  Verruca  Schumacher,  1817 

Subgenus /I ///vernvcfl  Pilsbry,  1916 

Verruca  (Altlverruca)  allisoni  n.  sp. 

Figure  10 

Material.-Sty\-1,  680901,  Hess  Guyot  (174°24.8'  W,  17°53.2'  N),  1,718-1,770  m  (Sigsbee  beam  trawl),  3 
spec,  on  trochid  gastropods:  Styx-7  680915  Sta.  1.  Darwin  Guyot,  (171°16.5'  E.  21°53.3'  N),  1.300-1.353  m  (rock 
dredge),  2  spec,  on  manganese  fragment. 

De'/70.5//o/;v.-HolotypeU.S.N.M.no.  140951  (Styx-7, 6809 15);  Paratypes  U.S.N. M.  no.  140952 (Styx-7, 680901). 

Z)/ag«o5/5.— Distinguished  from  all  other  A  Itiverruca  in  having  7  rather  than  3  or  4  in- 
terlocking teeth  forming  the  suture  between  the  carina  and  rostrum. 

Description.—ShQll  white,  without  persistent  yellow  cuticle.  Suture  between  rostrum 
and  carina  formed  by  numerous  interdigitating  ribs  (Fig.  lOH).  It  can  be  deduced  from 
successive  growth  lines  that  the  number  of  ribs  increases  throughout  life.  In  the  holotype 
this  number  has  increased  from  as  few  as  3  or  4  to  7.  Sutures  formed  by  rostrum  overlap- 
ping fixed  scutum  and  by  carina  overlapping  fixed  tergum;  sutures  simple  except  carinal 
margin  of  fixed  tergum  is  ala-like.  Suture  between  fixed  tergum  and  fixed  scutum  formed 
by  an  ala-like  margin  on  former  and  radius-like  margin  on  latter  (Fig.  lOG).  Parietal  por- 
tion of  fixed  tergum  interdigitates  between  radius-like  and  parietal  portions  of  fixed  scutum. 

Movable  tergum  and  scutum  articulated  by  the  interdigitation  of  proximal  portions 
of  their  apico-basal  ridges  (Fig.  lOH).  Supplemental  ridges  parallel  main  ridges  on  scutal 
side  of  the  movable  tergum,  and  the  rostral  portion  of  the  movable  scutum  is  ornamented 
by  longitudinal  lines  (Figs.  lOK,  L). 

Crest  of  labrum  supports  numerous  teeth,  nearly  80  in  the  paratype;  palps  pointed, 
sparsely  covered  with  short  strong  setae  (Fig.  lOA).  Mandible  with  3  teeth,  not  including 
lower  cutting  edge  and  inferior  angle;  upper  margin  of  third  tooth  and  lower  cutting  edge 
serrate;  inferior  angle  of  several  stout  spines  (Fig.  lOB).  First  maxilla  with  group  of  long 
strong  spines  above  and  below  well  developed  notch;  inferior  angle  supporting  few  short, 
bifid  spines  (Fig.  IOC).  Second  maxilla  notched  and  sparsely  covered  with  setae  (Fig.  lOD). 

Second  cirri  resemble  more  the  first  than  the  succeeding  pairs.  Their  rami  are  rela- 
tively short,  subequal  and  uncoiled.  Intermediate  articles  of  posterior  pairs  ctenopod,  each 
supporting  1  long  plumose  and  1  short  simple  pair  of  setae,  and  often  1  minute  simple  seta, 
along  the  lesser  curvature  (Fig.  lOF).  Caudal  appendages  of  6  to  8  segments  and  less  than 


90 


Figure  10.  Verruca  (Altiverruca)  allisoni  n.  sp.  A-F  and  I-L,  Holotype.  A,  labrum  and  palp,  left  palp  removed;  B, 
mandible;  C,  maxilla  I;  D,  maxilla  II;  E,  pedicle  of  cirrus  VI  supporting  penis  and  caudal  appendage;  F,  inter- 
mediate article  of  cirrus  VI;  G,  lateral  view  of  entire  specimen  illustrating  relationship  of  carina,  fixed  tergum 
and  fixed  scutum;  H,  lateral  view  of  entire  specimen  illustrating  relationship  of  movable  terga  and  scuta  with  the 
carina  and  rostrum;  I  and  L,  interior  and  exterior  views  of  the  movable  scutum  respectively;  J  and  K,  interior  and 
exterior  views  of  the  movable  tergum  respectively. 


91 


Figure  1 1.  A.  Arcoscalpellum  michelottianum  (Seguenza),  carinal  view  of  female  X  1.42;  B,  right  side  view  of  the 
same  individual  X  1.6;  C,  A.  hawaiiense  (Pilsbry),  right  side  view  of  female  X  1.7;  D,  carinal  view  of  the  same 
individual  X  1.4;  E,  ?  Arcoscalpellum  sp.  scutum  X  4.2;  F.  ?  Arcoscalpellum  sp.,  carina  X  3.4;  G,  A.  alcockianum 
(Annandale).  right  side  view  of  hermaphrodite  X  1.57;  H,  Mesoscalpellum  gruvelii  (Anr\2LX\da.\t),  rightside  view  of 
female  X  1.4;  Mesoscalpellum  gruvelii  (Annandale),  right  side  view  of  female  X  3.9  (A-D  and  H  from  Styx-7, 
680905  Sta.  2,  Allison  Guyot;  E-F  from  Styx-7,  680910  Sta.  5,  Sio  Guyot;  G  and  I  from  Styx-7,  680903-04  Sta. 
1,  Allison  Guyot). 


92 


the  length  of  the  basal  segment  of  the  pedicel  of  cirrus  VI;  the  penis  relatively  short,  pro- 
vided distally  with  a  few  short  seta  (Fig.  lOE).  Cirral  counts  for  three  specimens  follow: 

I  II  III  IV  V  VI       * 


Hess  Guyot 
Styx-7,  680901 
(Paratype) 

9 
8 

7 
8 

13 
16 

22 
23 

22 

24 
26 

Hess  Guyot 
Styx-7.  680901 
(Paratype) 

11 
9 

9 

11 

13 
16 

17 
21 

20 
15 

17 
19 

Darwin  Guyot 
Styx-7,  680915 
Sta.  7 
( Paratype ) 

6 
6 

9 

10 

13 

14 

17 
16 

17 

19 

Remarks.  —  Verruca  (Altiverruca)  allisoni  is  similar  to  V.  (A.)  cristallina  Gruvel,  1907 
from  the  East  Indies,  V.  (A.)  gibbosa  Hoek,  1883  which  is  nearly  cosmopolitan,  and  V.  (A.) 
regularis  Nilsson-Cantell  1929  from  the  Nicobar  Islands.  It  differs  from  the  first  in  having  7 
rather  than  3  or  4  interlocking  teeth  between  the  rostrum  and  carina,  in  lacking  multiple 
interlocking  ridges  between  the  suture  of  the  fixed  tergum  and  scutum  and  in  lacking  the 
small  beaded  ridges  along  the  scutal  margin  of  the  rostrum.  It  differs  from  the  second  and 
third  in  having  7  rather  than  4  interlocking  teeth  between  the  rostrum  and  the  carina,  in 
having  the  movable  scutum  with  longitudinal  markings  and  in  having  a  fixed  scutum  lack- 
ing an  ala-like  rostral  margin. 

The  major  difference  between  this  and  other  species  of  Altiverruca  is  the  large  number 
of  interlocking  teeth  between  the  rostrum  and  carina.  In  fact  this  same  difference  separates 
this  species  from  all  other  Verruca  except  V.  (?Rostratoverruca)  dens  Broch  1931,  V.  (R.) 
intexta  Pilsbry  1912,  V.  (?R.)  koehleri  Gruvel  1907,  V.  (?R.)  nexa  Darwin  1854  and  V.  (Ver- 
ruca) scrippsae  Zullo  1964.  Broch's  illustration  indicates  there  are  about  6  interlocking  teeth 
between  the  rostrum  and  carina,  essentially  as  in  the  present  species.  Pilsbr)'  (1907c),  dis- 
cussing Darwin's  species,  says  that  there  are  7  ribs  on  the  carina  interlocking  with  the  ros- 
trum. It  is  curious  that  the  ne'w  Altiverruca  should  be  so  similar  in  this  regard  to  these  mem- 
bers of  Rostratoverruca.  One  might  suspect  that  the  subgeneric  diagnosis  was  wrong. 
However  there  is  no  question  that  the  apex  of  the  rostrum  is  not  separated  from  the  scutal 
margin  of  the  plate,  as  it  is  in  Rostratoverruca.  The  similarity  to  V.  (V)  scrippsae  is  only  with 
regard  to  the  carino-rostral  suture;  the  complex  interlocking  sutures  between  the  fixed  scu- 
tum and  the  rostrum  and  the  fixed  tergum  and  the  carina  are  wholly  lacking  in  V.  (A.)  alii-' 
soni. 

ACKNOWLEDGMENTS, 

This  paper  is  a  contribution  from  the  Scripps  Institution  of  Oceanography  and  was  supported  in  part  by 
National  Science  Foundation  grants  GB-7596  and  GB-30908X. 

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1938.  Cirripedes  from  the  Indian  Ocean  in  the  collection  of  the  Indian  Museum,  Calcutta.  Mem.  Indian 

Mus.  13(1):  1-81. 
Pilsbry,  H.A. 

1907a.  Hawaiian  Cirripedia.  Bull.  U.S.  Bureau  Fish.  26:  181-190. 

1907b.  Cirripedia  from  the  Pacific  Coast  of  North  America.  Bull.  U.S.  Bureau  Fish.  26:  193-204. 

1907c.  The  barnacles  (Cirripedia)  contained  in  the  collections  of  the  U.S.  National  Museum.  U.S.  Natl.  Mus. 

Bull.  60:  1-122. 
1907d.  Notes  on  the  cirriped  genus  Megalasma.  Proc.  Acad.  Nat.  Sci.  Philadelphia  59:  408-416. 

1911.  Barnacles  of  Japan  and  Bering  Sea.  Bull.  U.S.  Bureau  of  Fish.  29:  59-84. 

1912.  Diagnoses  of  new  barnacles  from  Philippine  Archipelago  and  China  Sea.  Proc.  U.S.  Natl.  Mus.  42: 
291-294. 

1916.  The  sessile  barnacles  (Cirripedia)  contained  in  the  collections  of  the  U.S.  National  Museum,  including 
a  monograph  of  the  American  species.  U.S.  Natl.  Mus.  Bull.  93:  1-366. 
Schumacher,  C.F. 

1817.  Essai  d'un  nouveau  Systeme  des  habitations  des  Vers  testaces.  Schultz,  Copenhagen. 
Seguenza,  G. 

1873-1876.  Ricerche  palaentologiche  intorno  ai  Cirripedi  Terziarii  della  provincia  di  Messina.  Con  appen- 
dice  intorno  ai  Cirripedi  viventi  nel  Mediterraneo,  e  sui  fossili  terziarii  dell  "Italia  meridionale.  Pt.  1, 
Balanidi  e  Verrucidi,  1873:  Pt.  II,  Lepadidi,  1876.  Atti  Accad.  Pontaniana,  Napoli.  10:  265-481. 
Stewart,  F.H. 


94 


1911.  Studies  in  postlarval  development  and  minute  anatomy  in  the  genera  Scalpellum  and  Ibla.  Mem.  In- 
dian Mus.  3(2):  33-51. 
Stubbings,  H.G. 

1961.  Cirripedia  from  the  Tropical  West  Africa.  Atlantide  Rept.  6:  7-41.  " 

Zevina,  G.B. 

1969.  Cirripedia  Thoracica.  The  Biology  of  the  Pacific  Ocean,  book  II  pt.  I,  Deep-Sea  Bottom  Fauna,  V.G. 
Kort.  ed.,  Inst.  Okeanol.  Akad.  Sci.  U.S.S.R.:  66-68. 

1972.  Benthic  Lepadomorpha  (Cirripedia  Thoracica)  from  the  Southeast  Pacific.  Crustaceana  22(  1):  39-63. 
Zullo,  V.A.  R.F.  Kaar,  J.W.  Durham  and  E.C.  Allison. 

1964.  The  echinoid  genus  Salenia  in  the  Eastern  Pacific.  Palaeont.  7(2):  331-349. 
Zullo,  V.A.  and  W.A.  Newman. 

1964.  Thoracic  Cirripedia  from  a  Southeast  Pacific  Guyot.  Pacific  Sci.  18(4):  355-372. 


Scripps  Institution  of  Oceanography,  LaJolla,  California  92037 


sa^  ^6/r 


MUS.  COMF.  ZOOi:. 
LIBRARY 

DEC    71976 

HARVARD 


A  NEW  MITRID  FROM  THE  WESTERN  ATLANTIC 


GEORGE  E.  RADWIN  AND  LOYAL  J.  BIBBEY 


TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  7  31  AUGUST  1972 


A  NEW  MITRID  FROM  THE  WESTERN  ATLANTIC 


GEORGE  E.  RADWIN  AND  LOYAL  J.  BIBBEY 


ABSTRACT.— Mitra  helenae  n.  sp.  from  Cay  Sal  Bank,  between  the  Florida  Keys  and  Cuba,  is  assigned  to 
the  subgenus  Pleioptvgma  Conrad,  1863,  which  was  previously  known  to  contain  only  species  of  Miocene  and 
Pliocene  age.  The  large  size  of  this  gastropod,  its  distinctive  clouded  color  pattern  and  its  threaded  sculpture  are 
unique  among  Recent  western  Atlantic  mitrids.  Although  assigned  to  the  Mitridae,  the  true  familial  affin- 
ities oi  Pleioptvgma  and  this  new  species  must  await  study  of  the  radular  dentition. 

Malacology  has  long  benefitted  from  the  cooperation  of  commercial  fishermen  whose 
constant  searching  of  the  ocean  for  the  objects  of  their  commerce  has  led  them  to  discover 
many  new  forms  of  marine  life. 

Recently,  through  the  kindness  of  Mr.  Ivan  Thompson  of  El  Cajon,  California,  we  ex- 
amined two  specimens  of  a  remarkable  and  apparently  undescribed  mitrid.  Mr.  Thompson 
received  these  gastropods  from  Captain  Jack  Casey  of  Marathon,  Florida,  who  reported 
collecting  them  in  lobster  pots  set  at  a  depth  of  2 1 .5  m  on  Cay  Sal  Bank,  between  the  Florida 
Keys  and  Cuba,  in  December,  1971.  Both  shells  were  inhabited  by  hermit  crabs  whose  well- 
known  carnivorous  and  scavenging  feeding  habits  almost  certainly  account  for  their  pres- 
ence in  the  pots. 

Family  Mitridae  Swainson,  1831 

Genus  Mitra  Lamarck,  1798 

Subgenus  Pleioptvgma  Conrad,  1863 

Type  species.  — Valuta  carolinensis  Conrad,  1840,  by  monotypy;  Miocene;  North  Caro- 
lina. 

Definition.— ^hoW  large,  up  to  125  mm  in  length,  fusiform,  elongate,  inflated,  and  mod- 
erately light  in  weight;  protoconch  of  2  or  IVi  smooth  whorls,  teloconch  of  6  to  8  slightly 
convex  whorls.  Sculpture  consisting  of  moderate  to  very  sharp  spiral  threads  or  cords, 
spaced  irregularly  and  becoming  obsolete  medially  on  the  whorl.  Aperture  elongate, 
slightly  longer  than  spire,  smooth  within,  lip  edge  thin,  with  or  without  a  slight  swelling 
anteriorly  just  below  the  shoulder  slope  on  the  inner  surface  of  the  lip;  columella  with  6-9 
irregular,  moderately  thin,  simple  folds.  Siphonal  notch  weak  to  moderately  strong;  a  thin 
columellar  callus  extends  along  the  entire  inner  margin.  (Modified  after  Cernohorsky, 
1970:60). 

/^ema/'A.y.— Cernohorsky  (1970)  indicated  that  Pleioptvgma  probably  belongs  in  the 
Volutidae.  He  based  his  opinion  on  "large  size,  inflated  and  light  shell,  large  columellar  cal- 
lus, absence  of  a  siphonal  notch  and  thin,  irregular,  often  intercalate  columellar  folds." 

We  disagree  with  this  placement  for  several  reasons,  but  primarily  as  a  result  of  our 
examination  of  two  Recent  specimens  of  a  species  referable  to  Pleioptvgma.  It  seems  to  us 
that  the  extremely  diverse  nature  of  mitrid  and  volutid  shell  form  greatly  weakens  Cerno- 
horsky's  arguments.  Although  volutids  average  larger  than  mitrids,  the  existence  of  such 
species  as  Mitra  swainsoni,  M.  belcheri  and  M.  mitra,  all  of  which  reach  120- 150mm,  clearly 
shows  the  potential  for  large  size  in  this  family.  Cernohorsky's  other  arguments  are  equally 
difficult  to  accept  as  criteria  for  excluding  Pleioptvgma  from  the  Mitridae.  A  siphonal  notch 
is  apparent  on  our  specimens  of  the  type  species  and  on  our  new  species.  Irregularities  in  the 
plication  and  extent  of  callus  development  are  certainly  no  greater  here  than  in  Dibaphus 
Philippi,  1847,  an  unquestioned  mitrid  with  no  plaits  or  callus. 

Our  contention  for  a  mitrid  assignment  is  based  on  the  general  form  of  the  shell  and,  in 
particular,  on  the  unusual  clouded  color  pattern  and  the  threaded  sculpture.  Also,  the  pro- 
toconchs  of  both  fossil  and  Recent  species  (Figs.  7,  8)  are  different  from  any  known  type  of 
volutid  protoconch  (see  Pilsbry  and  Olsson,  1954).  We  are  thus  tentatively  placing  the  sub- 
genus Pleioptvgma  in  the  Mitridae,  pending  examination  of  the  radular  dentition  of  A/.  (P.) 

SAN  DIEGO  SOC.  NAT.  HIST..  TRANS.  17  (7):  95-100,  31  AUGUST  1972 


96 


helenae. 

As  Cernohorsky  noted,  no  Recent  representatives  have  been  found. 

Mitra  (Pleioptygma)  helenae  n.  sp. 

Type  locality.— Cay  Sal  Bank  (between  the  Florida  Keys  and  Cuba),  ca.  23°45'N., 
80°20''W.,  21.5  m.  Captain  Jack  Casey  coll.,  December,  1971  (holotype:  Figs.  2,  5). 

Type  depository.— Hololype,  San  Diego  Soc.  Nat.  Hist.,  Mar.  Invert,  no.  61863;  para- 
type,  collection  of  Ivan  Thompson. 

Diagnosis.— Mitra  helenae  is  comparable  to  two  fossil  species  from  the  southeastern 
United  States.  It  is  similar  in  size  and  shape  to  M.  carolinensis  (Conrad,  1840),  a  species  that 
is  probably  identical  to  M.  heilprini  Cossmann,  1899  (  =  A/.  lineolata  Heilprin,  1887,  not 
Bellardi,  1885).  It  differs  from  M.  carolinensis  in  its  more  poorly  marked  columellar  callus, 
its  less  sharp-crested  more  closely  spaced  spiral  threads,  its  broader  more  inflated  penulti- 
mate nuclear  whorl,  its  more  strongly  impressed  suture,  its  more  apparent  siphonal  fasciole, 
its  slopingly  shouldered  body  whorl  and  its  possession  of  small  intermediate  plaits  between 
the  anterior  columellar  plaits. 

The  other  fossil  species,  M.  prodroma  Gardner,  is  probably  the  ancestor  of  A/,  heilprini 
(see  Gardner,  1937:406).  It  is  generally  much  smaller  than  M.  helenae  (avg.  length  69mm. 
vs.  112mm.),  has  fewer  (3-5)  columellar  plaits,  which  are  of  regularly  increasing  promi- 
nence, and  has  a  proportionately  smaller  body  whorl  that  makes  up  about  three-fifths  of  the 
total  shell  length  compared  to  two-thirds  or  more  of  the  total  shell  length  in  M.  helenae. 

Chronologically,  M.  prodroma  was  the  first  to  appear,  followed  by  M.  carolinensis  and 
then  by  M.  helenae.  Morphologically,  as  well  as  chronologically,  M.  carolinensis  apparently 
is  closer  to  M.  helenae. 

No  other  western  Atlantic  mitrid  has  been  reported  to  reach  the  size  of  M.  helenae. 
Another  Floridian  member  of  the  family,  M.  (Dibaphimitra)  florida  Gould,  1856,  reaches  a 
relatively  large  size  (38-50mm)  but  has  a  more  convex  whorl  profile,  a  shorter  spire,  a  more 
ventricose  body  whorl  and  a  white  shell  with  spiral  rows  of  brown  dots  and  some  nebulous 
brown  blotches. 

Species  to  which  M.  helenae  could  be  compared  in  its  color  pattern  and  sculpture  in- 
clude M.  versicolor  Reeve,  1844,  M.  nebulosa  Reeve,  1844,  M.  lamarcki  Reeve,  1844  and  M. 
serpentina  Lamarck,  1822.  None  of  these  reach  the  size  of  M.  helenae,  none  have  its  almost 
volute-like  form  and  all  are  apparently  hmited  to  the  Indo-west  Pacific. 

Description.— J\iQ  shell  is  large  for  the  genus  (98- 123mm  in  length).  It  is  moderately 
heavy,  fusiform,  and  has  a  moderately  high  spire  (about  2/5  of  total  shell  length).  The  shell 
surface  is  smooth  and  polished.  The  spire  whorls  are  demarcated  by  an  impressed  suture. 
The  spire  consists  of  2y4  smooth,  polished,  tightly  wound  nuclear  whorls  and  7  or  8  weakly 
convex  postnuclear  whorls. 

The  body  whorl  is  large  (about  3/5  of  total  shell  length)  and  fusoid;  it  is  weakly  shoul- 
dered a  short  distance  anterior  to  the  suture  and  tapers  gradually  toward  the  anterior  end. 
The  aperture  is  long,  moderately  narrow,  and  almost  rectangular,  except  at  its  posterior 
end.  The  outer  apertural  lip  is  thin  and  even  in  a  mature  specimen.  Just  below  the  shoulder 
margin,  on  the  inner  surface  of  the  outer  apertural  lip,  there  is  a  slight  swelling  extending 
anteriorly  for  about  25  mm.  The  inner  lip  is  oblique  and  is  coated  with  a  thin  callus  of  minor 
extent.  The  inner  lip  bears  a  series  of  9  plaits  of  various  strengths.  The  two  posterior-most 
are  strongest  and  of  these  the  first  is  stronger  and  thicker  than  the  second.  These  are  fol- 
lowed anteriorly  by  1  weak  and  6  moderately  weak  plaits  that  diminish  in  strength  and  ex- 
tent of  projection  from  the  aperture  proceeding  anteriorly.  The  siphonal  fasciole  is  well- 
defined,  originating  as  a  white  raised  ridge  at  the  fifth  plait  from  the  upper  end  of  the  series. 
The  siphonal  notch  is  well-defined  and  moderately  deep. 

Axial  sculpture  is  lacking  except  for  fine  growth  lines,  and  erratically  occurring 
stronger  lines  representing  major  growth  stoppages.  Spiral  sculpture  on  the  spire  whorls 
consists  of  numerous  fine  erratically  spaced  cords;  2  or  3  immediately  below  the  suture  are 
bunched  more  closely  than  the  others.  The  stronger  primary  cords  are  sharply  raised  and 
bear  an  interrupted  brown  and  white  spotted  color  pattern  that  is  distinct  from  the  back- 
ground. Weaker  secondary  cords  are  ephemeral  and,  as  such,  are  not  visible  uniformly  over 


Figure  1,  4.  Mitra  ( Pleioptvgma)  helenae  n.  sp.,  paratype.  Cay  Sal  Bank,  21.5  m,  in  lobster  pots,  length- 123  mm, 
maximum  diameter— 41.1  mm.  collection  of  Ivan  Thompson.  2,  5,  M.  (P.)  helenae  n.sp.,  holotype.  Cay  Sal  Bank, 
21.5  m,  in  lobster  pots,  length— 98.4  mm,  maximum  diameter— 32. 1  mm,  SDSNH  Mar.  Invert,  no.  61863.  3,  6,  M. 
(P.)  carolinensis  (Conrad,  1840),  Pliocene,  Clewiston,  Florida,  length- 103  mm,  maximum  diameter— 34.8  mm, 
SDSNH  Paleo.  no.  07248. 


98 


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<£■  .;i  ./A'f''^  .vi-a"*!'        ^-'^^f^" 


ilip^"" 


a 


Figure  7.  M/Vra  fP.j  carolinensis  (Conrad,  1840), 
protoconch,  locality  data  as  in  Figure  3. 


Figure  8.  Mitra  (P.)  helenae  n.sp.,  protoconch,  lo- 
cality data  as  in  Figure  2. 


the  shell.  This  spiral  sculpture  becomes  partially  obsolete  on  the  periphery  of  the  body 
whorl. 

The  shell  is  white  with  numerous  irregular  diffuse  flammules  of  reddish  chestnut 
brown.  The  interior  of  the  aperture  is  porcellaneous  white. 

Mea5wreweA7/5.—Holotype— length,  98.4mm;  greatest  diameter,  32.1mm;  paratype— 
length,  123mm  (lacking  protoconch);  greatest  diameter,  41.1  mm. 

Remarks.— J\\Q  holotype  and  the  single  paratype  were  inhabited  by  hermit  crabs  at  the 
time  they  were  collected.  The  holotype  is  in  fresh  condition  and  apparently  is  not  full- 
grown;  it  has  a  thin,  immature,  outer  apertural  hp.  The  paratype  has  apparently  attained 
full  size  but  it  lacks  a  protoconch;  its  surface  is  more  worn  and  the  color  pattern  is  com- 
paratively faded. 

Mitra  helenae  is  here  considered  a  living  representative  of  Pleioptygma,  a  genus  that  is 
known  otherwise  only  from  Miocene  and  Pliocene  species.  Other  species  previously  as- 
signed to  this  group  include  M.  carolinensis,  M.  heilprini  and  M.  prodroma. 

Etymologv.— This  patronym  honors  the  late  Mrs.  Helen  Thompson  of  El  Cajon,  Cah- 
fornia. 

ACKNOWLEDGMENTS 

We  thank  Captain  Jack  Casey  and  Mr.  Ivan  Thompson  for  their  interest  and  courtesy  in  providing  us  with  the 
only  known  specimens  of  M.  helenae.  Mr.  David  K.  Muiliner  photographed  the  specimens  and  Mr.  Clifton  Martin 
supplied  references.  Mr.  Anthony  D'Attilio  illustrated  the  protoconchs. 

LITERATURE  CITED 

Bellardi,  Luigi 

1850.  Monografia  delle  Mitre  fossili  del  Piemonte.  Mem.  Real.  Accad.  Sci.  Torino,  ser.  2,  11:1-34,  pis.  1-2. 
(not  seen) 

Cernohorsky,  W.  O. 

1970.  Systematics  of  families  Mitridae  and  Volutomitridae  (Mollusca,  Gastropoda).  Bull.  Auckland  Inst. 
Mus.,  194p. 

Conrad,  T.  A. 

1840.  New  fossil  shells  from  North  Carolina.  Amer.  J.  Sci.  Arts  39:  387-388. 

Cossmann,  A.  E.  M. 

1899.  Essais  de  Paleoconchologie  Comparee,  Paris,  3:  1-201,  pis.  1-8. 

Gardner,  J. 

1937.  The  molluscan  fauna  of  the  Alum  Bluff"  Group  of  Florida,  pt.  VI,  Pteropoda,  Opisthobranchia  and 
Ctenobranchia  (in  part).  U.S.  Geol.  Surv.  Prof  Paper  142F,  187p. 

Gould,  A.  A. 

1856.  Descriptions  of  new  shells.  Proc.  Boston  Soc.  Nat.  Hist.  6:  11-16. 


99 


Lamarck,  J.  B.  P.  A. 

1798.  Tableau  encyclopedique  et  methodique  des  trois  regnes  de  la  nature.  Paris,  pis.  287-390.  (not  seen) 

Philippi,  R.  A. 

1847.  Beschreibung  zweier  neuer  conchyliengeschlechter,  Dihaphus  und  Amphichaena  nebst  einigen  be- 
merkungen  uber  Cyamium,  Ervilia  und  Entodesma.  Arch.  Naturg.  13(  1):  61-66,  pi.  3.  (not  seen) 

Pilsbry,  H.  A.  and  A.  A.  Olsson 

1954.  Systems  of  the  Volutidae.  Bull.  Amer.  Paleont.  35(152):  1-36,  pis.  1-4  (25-28). 


Department  of  Marine  Invertebrates,  Natural  History  Museum,  P.O.  Box  1390,  San 
Diego,  California  92112  and  San  Diego  Shell  Club,  P.O.  Box  1390,  San  Diego,  California 
92112. 


■^**-^L.» 


.  ■■■^■i.  J*., 


:n. :, ^    ' 


5-4  a/ 


DIAGNOSES  OF  NEW  CYPRINID  FISHES  ^UfivS^**^' 

OF  ISOLATED  WATERS  IN  THE  GREAT  BASIN 
OF  WESTERN  NORTH  AMERICA 


CARL  L.  HUBBS  AND  ROBERT  RUSH  MILLER 


TRANSACTIONS 

OF  THE   SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  8  29  SEPTEMBER  1972 


DIAGNOSES  OF  NEW  CYPRINID  FISHES 

OF  ISOLATED  WATERS  IN  THE  GREAT  BASIN 

OF  WESTERN  NORTH  AMERICA 

CARL  L.  HUBBS  AND  ROBERT  RUSH  MILLER 


ABSTRACT.— One  new  genus,  two  new  species,  and  six  new  subspecies  are  diagnosed  from  highly  restricted 
endorheicbasinsof  the  western  United  States— Relictussolitarius  n.  gen.  andn.  sp.,  from  the  basins  of  pluvial 
lakes  Gale,  Franklin,  Steptoe,  and  Waring;  Gila  alvordensis  n.  sp.,  from  the  basin  of  Lake  Alvord;  and  the 
following  new  subspecies:  Gila  bicolor  newarkensls  and  G.  b.  euchila  ( Lake  Newark),  and  G.  b.  isolata  ( Lake 
Clover),  Rhlnichthys  osculus  reliquus  ( Lake  Gilbert),  R.  o.  ollgoporus  and  R.  o.  lethoponis  ( Lake  Clover). 

In  amplification  of  our  general  summary  (Hubbs  and  Miller,  1948),  we  are  now  docu- 
menting in  detail  the  correlations  between  the  hydrographic  history  of  the  endorheic  waters 
of  the  Great  Basin  and  the  differentiation  of  the  remnant  fish  fauna  that  somehow  has  man- 
aged to  survive  in  the  pitiful  remnants  of  the  pluvial  lakes  and  streams  that  in  late  Pleisto- 
cene time  covered  about  one-fifth  of  the  now  arid  area.  One  of  the  species  described  herein, 
and  the  post-Pleistocene  desiccation  of  the  Alvord  basin  to  which  it  is  rigidly  confined,  are 
under  intensive  study;  all  of  the  other  taxa  are  integral  parts  of  a  major  treatise  now  in  final 
processing  (Hubbs  and  Miller,  in  press). 

The  type  specimens  are  deposited  in  the  University  of  Michigan  Museum  of  Zoology 
(UMMZ). 

Relictus  n.  gen. 

Tvpe  species.— Relictus  solitarius. 

A  cyprinid  of  moderate  size  (larger  than  Rhinichthvs),  with  some  distinctive  os- 
teological  characters:  dorsal  crest  of  maxilla  greatly  expanded  upward  and  backward: 
cleithrum  slender;  supraethmoid  elongate,  slender  medially  but  notably  expanded  laterally 
at  front  (resembling  that  of  Rhinichthvs );  urohyal  long  and  narrow.  Vertebrae  35-39.  Phary- 
ngeal arch  moderately  strong  and  heavy,  but  rather  thin  and  somewhat  lacy  on  the  strongly 
expanded  median  section;  not  strongly  elevated  at  the  posterior  end  of  the  tooth  row;  with- 
out a  flattened  shelf  on  which  a  second  tooth  row  might  develop;  teeth  4—4  (rarely  5—4  or 
4— 3).  Gill-rakers  small  and  few  (7-12,  usually  8-11,  on  first  arch).  Mouth  oblique  and  termi- 
nal, completely  lacking  horny  cutting  edges;  no  frenum  or  barbel.  Lateral  line  obsolescent, 
rarely  extending  to  below  origin  of  dorsal  fin,  commonly  disrupted;  total  pores  3-29.  Supra- 
temporal  canal  seldom  complete  (only  4  of  76  specimens  have  the  commissure  closed),  with 
usually  3  or  4  (0-5)  pores  in  each  lateral  segment;  preoperculomandibular  pores  11-19; 
mandibular  pores  3-8.  Scales  rather  small  (50-70  transverse  rows),  poorly  imbricated  and 
markedly  irregular;  each  usually  vertically  oval  but  sometimes  becoming  rectangular  with 
age;  with  numerous  radii  on  all  fields  (much  as  in  Rhinichthvs  and  some  other  Western  gen- 
era). Fins  small  and  strongly  rounded;  the  pelvic  especially  and  uniquely  paddlelike;  dorsal 
and  pelvic  both  displaced  backward,  and  both  beginning  at  approximately  the  same  verti- 
cal (as  in  the  subgenus  Siphateles  of  the  genus  Gila  and  in  many  species  of  the  typical  sub- 
genus Gila);  dorsal  and  pelvic  rays  typically  8,  anal  7.  Nuptial  tubercles  form  a  highly  dis- 
tinctive pattern  on  head;  the  largest  uniserially  line  the  infraorbital  sensory  canal  and 
suborbital  margin;  large  uniserial  caducous  cones  (much  stronger  than  in  Gila)  line  the  up- 
per edge  of  the  first  pectoral  ray;  smaller  cones,  also  strictly  uniserial  (not  forking  once  as 
they  do  in  Rhinichthvs)  occur  along  one  to  several  following  rays;  in  high  males  some  tu- 
bercles develop  along  outer  pelvic  rays  and  along  first  anal  rays.  Head  and  body  turgid. 
Coloration  much  as  in  Siphateles.  rather  even,  and  often  with  large  melanophores  on  lower 
side;  lacking  the  two  lateral  bands,  the  head  stripe,  the  paired  light  spots  at  caudal  base,  and 

SAN  DIEGO  SOC.  NAT.  HIST.  TRANS.  17(8);  101-106,  29  SEPTEMBER  1972 


102 


Other  features  characteristic  of  Rhinicluhvs.  Intestine  forming  a  single,  simple,  compressed- 
S  loop,  as  in  Rhinichthvs  and  many  other  American  cyprinids.  Karyotype  distinguished  by  a 
relatively  large  number  (2  large  and  8  small)  of  acrocentric  chromosomes  but  many  (12) 
metacentrics;  remaining  28  are  subtelocentric  and  submetocentric  (total  50  as  in  other 
American  cyprinids  examined). 

Relictus  solitarius  n.  sp. 

Holotvpe.—VMMZ  186904,  a  nuptial  male  60.3  mm  in  standard  length,  from  upper, 
hillside  spring  on  Kirkpatrick  Ranch  (earlier  called  "Atwood  Ranch,"  later  called  "Don 
Phalan  Ranch")  on  east  side  of  Butte  Valley  north  of  the  narrows,  in  east  part  of  T.29  N., 
R.62  E.,  Elko  County,  Nevada,  21  km  northwest  of  Currie;  collected  by  the  Hubbs  family 
June  27,  1942  (collection  H42-47). 

The  characters  of  the  species  are  essentially  those  of  the  genus.  Counts  for  the  holotype 
and  the  paratypes  (UMMZ  141518)  from  the  same  collection  follow.  Rays:  dorsal  7-8 
(mean  7.40),  anal  6-7  (6.95),  caudal  18-21  (19.17),  pectoral  13-16  (14.17),  pelvic  7-9  (7.95). 
Vertebrae:  35-37  (36.05).  Scale-row  counts:  lateral-line  50-57  (54.6),  predorsal  30-33  (31.4), 
dorsal  to  anal  origins  2 1  -23  (22.4),  around  body  55-58(55.8),  around  peduncle  30-3 1  (30.2). 
Pores:  lateral-line  13-26  (18.4),  supratemporal  2-4  (3.0),  mandibular  4-7  (5.33).  Gill-rakers 
7-1 1  (8.90).  Measurements  of  holotype  in  thousandths  of  standard  length:  predorsal  length 
579,  anal  origin  to  caudal  base  318,  body  depth  295,  caudal-peduncle  depth  158,  head 
length  282,  head  depth  207,  head  width  170,  snout  length  76,  orbit  length  59,  upper-jaw 
length  82,  mandible  length  1 02,  interorbital  width  88,  suborbital  width  4 1 ,  depressed-dorsal 
length  223,  caudal  length  238,  pectoral  length  208,  pelvic  length  160. 

Gila  alvordensis  n.  sp. 

Holotvpe.— UMMZ  130495,  an  adult  female  70.7  mm  in  standard  length, from  Trout 
Creek,  tributary  to  Alvord  Desert,  in  Harney  County,  Oregon;  just  below  the  canyon  and 
just  below  bridge  where  roads  to  Denio,  Jordan  Valley,  and  Fields  meet,  in  southeast  part  of 
T.39  S.,  R.36  E.;  collected  by  the  Hubbs  family  July  26,  1934  (collection  M34-87). 

A  chub  of  moderate  size  (though  usually  greatly  dwarfed  in  Borax  Lake),  agreeing 
most  closely  with  Siphateles  (now  regarded  as  a  subgenus  of  Gila),  but  with  scales  much 
reduced  in  size  and  more  embedded,  and  with  radii  all  around,  much  as  in  Rhinichthvs  and 
Relictus.  Pharyngeal  teeth  uniserial,  normally  5—4  (rarely  5—5, 4—5, 4—4,  or  4—3),  with  the 
first  tooth  on  a  moderately  elevated  base.  Nuptial  tubercles  strong  on  the  flattened  and 
moderately  twisted  pectoral  fin  of  nuptial  males;  developed  on  the  outer  half  (by  number) 
of  the  rays,  over  at  least  two-thirds  of  the  width  of  the  fin,  covering  nearly  the  full  length  of 
each  thickened  ray;  uniserial  and  small  on  the  only  moderately  thickened  outermost  ray; 
the  row  branching  once  on  each  of  the  following  rays:  very  strong  on  rays  2  and  3  (on  the 
ridge  of  the  distorted  fin),  then  decreasing  inward  in  number  and  size;  each  tubercle  set  on  a 
single  ray  segment  and  rising  from  a  large  rounded  base  to  end  in  a  rather  narrow  and 
sharply  pointed,  essentially  erect,  tip  (with  only  a  slight  cant  basad);  in  high  males  similar 
but  weaker  tubercles  discernible  on  the  pelvic  fin,  but  not  on  other  fins;  minute  excres- 
cences, simulating  tubercles,  over  the  top  and  sides  of  head  in  high  males.  General  color 
dusky  with  a  continuous  file  of  large  melanophores,  usually  uniserial  or  nearly  so,  ahgned 
on  either  side  of  the  back. 

Fin  rays:  dorsal  7-10  (normally  7),  anal  6-9  (normally  7),  caudal  17-20  (normally  19), 
pectoral  12-17,  pelvic  7-9  (normally  8).  Gill-rakers:  16-22,  usually  short,  especially  forward. 
Measurements  of  holotype  in  thousandths  of  standard  length:  predorsal  length  568,  anal 
origin  to  caudal  base  312,  body  depth  250,  caudal-peduncle  depth  129,  head  length  267, 
head  depth  163,  head  width  139,  snout  length  77,  orbit  length  48.  upper-jaw  length  72,  man- 
dible length  100,  interorbital  width  85,  suborbital  width  33,  depressed-dorsal  length  204, 
caudal  length  238,  pectoral  length  191,  pelvic  length  141,  pelvic  insertion  to  anal  origin  185. 

Gila  bicolor  newarkensis  n.  subsp. 
Holotype. -UMMZ  188893,  a  nuptial  male  68.0  mm  in  standard  length,  from  spring  in 


103 


Newark  Valley  on  west  side  near  Diamond  Peak  (called  South  Peak  in  1934),  on  alluvial 
slope  about  opposite  south  end  of  Newark  Dry  Lake,  near  middle  of  T.IO  N.,  R.55  E.,  in 
northwestern  White  Pine  County,  Nevada;  collected  by  the  Hubbs  family  September  11, 
1934  (collection  M34-206). 

A  medium  to  rather  small-sized  chub  (largest  of  many  specimens  97  mm  long).  Gen- 
eral color  tone  darker  and  more  uniform  than  in  G.  b.  ohesa,  not  closely  approaching  the 
bicolored  pattern  of  that  subspecies;  dark  pigmentation  of  sides  less  uniform  than  in  other 
forms,  because  of  the  thick  and  broad  concentration  of  melanophores  around  margins  of 
scale  pockets,  leaving  the  rounded  central  area  of  pockets  largely  clear,  to  form  rather  con- 
spicuous stripes  along  the  horizontal  scale  rows;  the  dark  pigment  extending  farther  down, 
usually  more  or  less  completely  rounding  caudal  peduncle;  basicaudal  spot  replaced  by  a 
thin  blackish  streak  along  curving  posterior  border  of  squamation.  Head  and  body  strongly 
turgid,  rounded  in  all  aspects.  Muzzle  broadly  rounded;  mouth  generally  low,  curved,  and 
less  oblique  than  usual,  becoming  more  nearly  horizontal  forward;  mandible  slightly  in- 
cluded at  tip.  Nuchal  region  more  humped  than  in  most  forms;  dorsal  contour  scarcely  ele- 
vated at  front  of  dorsal  tin.  Fins  distinctively  rounded,  without  any  falcation;  unusually 
large;  sexual  dimorphism  in  fin  lengths  extreme.  Anal-ray  count  averaging  low,  modally  7; 
pelvic  rays  averaging  8.10  and  8.66  in  two  races.  Vertebral  and  scale  counts  averaging  low 
(scale  counts  around  body  averaging  fewer  than  47:  those  around  peduncle  fewer  than  27). 
Gill-rakers  outstandingly  few  (modally  12),  short,  soft,  and  swollen.  Pharyngeal  teeth  usu- 
ally 5-4. 

Gila  bicolor  euchila  n.  subsp. 

Holotype.—VMMZ  124938,  an  adult  female  141  mm  in  standard  length,  from  Fish 
Creek  Springs  in  northwestern  part  of  Fish  Creek  (Little  Smoky)  Valley,  in  main  ditch 
about  0.5  km  below  junction  of  two  main  spring-fed  branches,  in  Sec.  8,  T.  16  N..  R.53  E.; 
near  southwest  corner  of  Eureka  County,  Nevada;  collected  by  the  Hubbs  family  August 
17,  1938  (collection  M38-134). 

An  outstandingly  large  chub  (for  an  isolated  population),  males  reaching  114  mm  and 
females  149  mm;  the  distinction  in  bulk  is  even  more  striking  than  in  length.  Agreeing  with 
G.  b.  newarkensis  in  color  pattern  (as  described  above),  but  differing  in  color:  females  deep 
moss-green  on  back,  with  scale  borders  tending  to  converge  backward,  with  sides  usually 
mottled  or  speckled  on  individual  scales,  with  lower  fins  deep-olive,  grading  to  blackish  on 
rays  and  to  yellowish  on  membranes,  and  with  dorsal  and  caudal  fins  very  dark  olive;  adult 
males  with  much  more  gilt  than  females  on  cheeks,  opercles,  and  sides,  and  with  gilt  on 
body  somewhat  rosy,  with  blue  reflections  rather  strong  on  lower  sides,  with  scale  margins 
ventrally  orange-red,  with  a  considerable  wash  of  lemon-orange  on  dorsal  and  caudal  fins, 
with  axils  of  paired  fins  rather  bright  orange,  with  this  color  rather  strong  on  interradial 
membranes,  and  with  rays  of  lower  fins  deep-olive.  Body  contours  typically  much  less  tur- 
gid than  in  G.  b.  newarkensis,  and  head  much  more  pointed  in  side  view,  with  the  tip  much 
nearer  horizontal  midline  of  head;  anterodorsal  profile  much  straighter  and  less  decurved; 
head  much  larger:  suborbital  and  muzzle  wide  and  flat:  mouth  much  larger,  straighter,  and 
more  oblique,  with  particularly  massive  lips  and  mandible  (yet  tip  of  mandible  is  also 
slightly  included).  Fins  hardly  falcate,  but  less  rounded  than  in  G.  b.  newarkemis.  Supra- 
temporal  canal,  as  also  only  in  G.  b.  newarkensis,  but  in  contrast  with  other  forms  of  G.  bico- 
lor, more  often  complete  than  incomplete.  Dorsal  fin  more  posteriorly  inserted  than  in  other 
subspecies,  even  more  than  in  G.  b.  newarkensis.  Paired  fins  in  males  larger  than  in  nearly  all 
other  populations  studied.  Scale-row  counts,  as  in  G.  b.  newarkensis,  average  lower  than  in 
other  forms,  with  little  overlap  in  most  categories.  Gill-rakers  average  few  and  generally 
shorter  and  less  hard  than  usual  in  G.  b.  obesa. 

Gila  bicolor  isolata  n.  subsp. 

Holotype.-VMMZ  186906,  an  adult  female  85.8  mm  in  standard  length,  from  Warm 
Springs  of  Independence  Valley  (also  known  as  Ralph's  Warm  Springs),  just  off  base  of  Pe- 
quop  Mountains,  approximately  on  edge  of  bed  of  pluvial  Lake  Clover,  on  either  side  of 


104 


T. 35-36  line  near  middle  of  R.66  E.,  in  east-central  Elko  County,  Nevada;  collected  by 
Miller  and  Hubbs  August  25,  1965  (collection  M65-33). 

A  somewhat  dwarfed  chub  (largest  male  73  mm  and  largest  female  91  mm  long).  Un- 
pigmented  ventral  band  wider  than  in  G.  b.  newarkensis  and  G.  b.  eiichila\  the  pigment 
almost  never  rounding  peduncle  below;  however,  almost  all  specimens  have  a  highly  dis- 
tinctive black  speck  on  midventral  line  at  the  very  outset  of  the  lower  procurrent  caudal 
rays.  Anterodorsal  profile  less  rounded  and  decurved  than  usual  in  G.  b.  newarkensis.  As  in 
G.  b.  obesa,  contrasting  with  G.  b.  newarkensis  and  G.  b.  euchila  front  tips  of  mandible  and 
upper  lip  about  even;  in  contrast  with  G.  b.  newarkensis,  mouth  nearly  straight,  and  suffi- 
ciently oblique  to  rise  nearly  to  lateral  midline  of  head.  Lateral  line,  even  in  larger  adults, 
usually  incomplete  posteriorly,  lacking  at  least  on  peduncle,  usually  throughout  that  region 
and  in  some  farther  forward,  where  it  may  be  either  lacking  or  interrupted.  Supratemporal 
canal  regularly  complete,  as  in  none  of  the  other  subspecies  studied.  Dorsal  fin,  with  little 
overlap,  farther  back  than  in  any  of  the  other  forms  considered  except  G.  b.  newarkensis  and 
G.  b.  euchila.  Distance  from  anal  origin  to  caudal  base  averaging  shorter  than  in  the  other 
subspecies  considered,  including  G.  b.  newarkensis  but  not  G.  b.  euchila.  Mandible  aver- 
aging larger  than  in  other  forms  considered,  except  G.  b.  euchila  and  the  variant  form  of  G. 
b.  obesa  in  Sulphur  Spring  (Diamond  Valley).  Sexual  dimorphism  of  pectoral  fin  about  as  in 
G.  b.  obesa,  much  less  than  in  G.  b.  newarkensis  and  G.  b.  euchila.  Anal  rays  predominantly  7 
rather  than  8— as  also  in  G.  b.  euchila  and  two  of  the  three  populations  of  G.  b.  newarkensis 
studied.  Pelvic  rays  predominantly  8  instead  of  9  (as  in  two  G.  b.  newarkensis  populations). 
Numbers  of  vertebrae  and  scale  rows  low.  Gill-rakers  also  few  (8-14,  averaging  11.14). 
Rakers  essentially  like  those  of  G.  b.  obesa. 

Rhinichthys  osculus  reliquus  n.  subsp. 

Holotvpe.—VMMZ  124906,  an  adult  female  67  mm  in  standard  length,  from  spring- 
fed  creek  in  a  grassy  meadow  in  the  partly  enclosed  southwestern  arm  of  Grass  Valley,  13 
km  east  of  Mt.  Callaghan,  in  course  of  Callaghan  (Woodward)  Creek,  on  Grass  Valley 
Ranch,  in  SW  1/4,  Sec.  10,  T.21  N.,  R.46  E.,  in  eastern  Lander  County,  Nevada;  collected 
by  Hubbs  family  and  Miller,  August  9,  1938  (collection  M38-116). 

A  relatively  large  dace,  despite  its  occurrence  (now  apparently  extinct)  in  a  restricted 
habitat;  largest  size  82  mm.  Quite  different  in  appearance  from  R.  o.  robustus:  body  less 
speckled;  blackened  regenerated  scales  rather  fewer  and  less  emphasized;  underlying  main 
dark  lateral  band  generally  broader,  more  solid,  more  even-edged.  Pattern  further  in- 
tensified by  more  definitely  lightened  ground  color  between  this  lateral  band  and  the  dark, 
broad  predorsal  stripe.  Deep-lying  giant  melanophores  often  formed  on  the  lower  sides, 
especially  posteriorly,  much  more  conspicuous  than  in  R.  o.  robustus,  forming  punc- 
ticulations  somewhat  similar  to  those  on  this  region  in  subgenus  Siphateles  of  Gila.  Lower 
dark  lateral  line,  usually  rather  well  developed  in  R.  o.  robustus,  obsolescent.  A  very  dis- 
tinctive dark  streak  or  wedge  developed  along  lower  border  of  caudal  peduncle.  Head  char- 
acteristically darkened  from  the  dark  area  of  the  suborbital  region,  and  from  front  of 
mouth,  upward  and  backward  over  front  and  top  of  head:  horizontal  dark  stripe  on  snout, 
characteristic  of  R.  o.  robustus,  barely  even  suggested.  Vertical  fins  also  more  uniformly 
darkened,  and  less  speckled,  than  in  R.  o.  robustus,  with  hardly  a  trace  of  the  especial  black- 
ening at  the  bifurcation  of  the  rays.  Lower  lip,  even  in  specimens  with  lower  surface  of  head 
elsewhere  devoid  of  pigment,  heavily  punctate  all  around.  Red  color  often  apparent  in  Rhi- 
nichthys osculus,  in  axils  of  paired  fins,  about  mouth,  and  on  preopercle,  scarcely  evident  in 
life.  Body  more  turgid  in  nuchal  region,  and  snout  more  rounded,  more  declivous,  and 
broader  (overall  width  of  mouth,  in  consequence,  approximately  equally  as  long  as,  rather 
than  shorter  than,  the  snout).  Mouth  as  seen  from  below  broadly  U-shaped,  instead  of 
being  narrower  approaching  a  V.  Barbel  almost  invariably  absent.  Lateral  line  on  both 
body  and  head  greatly  reduced;  supratemporal  canal  commissure  consistently  interrupted 
medially,  typically  very  widely.  Body  averaging  slenderer  than  in  other  forms;  caudal-pe- 
duncle depth  is  less  than  in  R.  o.  lethoporus,  with  slight  overlap.  Pelvic-fin  insertion  more 
posterior  than  in  any  other  form  considered.  Sexual  dimorphism  in  pectoral-fin  length  most 
extreme.  Dorsal  fin  more  posteriorly  inserted  in  males  than  in  females,  on  the  average,  con- 


105 


trary  to  findings  for  other  forms  oi'  Rhinichthvs  (and  for  cyprinids  in  general).  Pectoral-ray 
counts  on  the  average  lower  than  in  other  subspecies  treated.  Caudal  vertebrae  definitely 
averaging  fewer.  Scale  counts  averaging  consistently  higher  than  in  the  two  forms  described 
below. 

Rhinichthys  osculus  oligoporus  n.  subsp. 

Holotvpe.—VMMZ  186902,  an  adult  female  55.2  mm  in  standard  length,  from  Warm 
Springs  in  Clover  Valley,  at  Warm  Creek  (formerly  Clover)  Ranch,  near  southeastern  cor- 
ner of  Clover  Valley,  near  foot  of  bajada  just  above  the  ancient  bed  of  Lake  Clover,  in  Sec. 
7,  T.33  N.,  R.61  E.,  in  southeastern  Elko  County,  Nevada;  collected  by  James  E.  Deacon 
and  Mary  Beth  Rheuben  September  14,  1964. 

A  dace  of  about  average  size.  Body  more  extensively  speckled  with  black  than  in  R.  o. 
robustus;  lower  lateral  band  as  a  rule  much  less  or  not  at  all  evident;  dark  pigmentation 
around  snout  generally  diffused,  with  no  evidence  of  the  usual  horizontal  black  streak  in 
front  of  eye.  but  retaining  a  tendency  for  its  continuation  across  opercle.  Jet-black  basicau- 
dal  wedge  much  reduced  in  size  and  intensity,  more  disrupted,  occasionally  hardly  evident. 
Dusky  dashes  on  dorsal  and  caudal  fins  tending  to  be  more  numerous,  but  barely  evident  on 
anal  fin  (where  often  evident  in  R.  o.  robustus);  these  marks  much  less  apt  to  form  in,  and  to 
be  largely  restricted  to,  the  crotches  of  the  bifurcating  rays.  Main  lateral  band  bordered 
above  by  a  light  streak,  barely  evident  in  R.  o.  robustus.  Life  color  on  back  bright-olive  or 
golden-green  and  below  silvery,  with  an  intervening  bright-gilt  stripe  and  with  dusky  mot- 
tling. Axils  of  paired  fins  and  base  of  anal  in  adult  male  clear  red  (contrasting  with  males  of 
R.  o.  reliquus).  Differing  from  R.  o.  robustus  in  general  form:  outlines  of  body,  and  espe- 
cially of  head,  more  curved;  head  in  particular  more  rounded,  in  both  dorsal  and  lateral 
aspects.  Mouth  tending  to  be  more  definitely  lower  than  lower  border  of  eye,  and  generally 
more  curved;  whole  aspect  bulkier.  Barbel  invariably  absent  (51  specimens).  Reduction  of 
lateral  line  on  body  extreme  (as  in  R.  o.  reliquus  and  R.  o.  lethoporus).  Suborbital  averaging 
slightly  narrower  than  in  other  forms  studied,  R.  o.  lethoporus  excepted.  Pelvic-fin  insertion 
averaging  farther  back  than  in  typical  R.  o.  robustus  or  in  R.  o.  lethoporus,  but  farther  for- 
ward than  in  R.  o.  reliquus.  Number  of  rays  in  paired  fins  somewhat  reduced  in  average 
number.  Scale  counts  averaging  definitely  lower  than  in  R.  o.  reliquus,  about  the  same  or 
not  quite  so  low  as  in  R.  o.  lethoporus,  and  somewhat  lower  than  in  more  typical  races  of  R. 
o.  robustus. 

Rhinichthys  osculus  lethoporus  n.  subsp. 

Holotvpe.—VMMZ  186905,  an  adult  female  35.3  mm  in  standard  length,  from  Warm 
Springs  in  Independence  Valley  (the  same  collection  from  which  the  type  of  Gila  bicolor 
isolata  was  taken;  see  above). 

Apparently  the  most  dwarfed  dace  of  any  in  the  general  area  under  consideration:  the 
largest  male  measures  34  mm  and  the  largest  female  39  mm  in  standard  length,  among  the 
101  specimens  collected  (not  much  larger  than  young-of-the-year  of  some  of  the  other 
forms).  Dark  speckling  usually  very  fine,  and  tending  to  extend  downward  across  the  caudal 
peduncle;  lower  edge  of  peduncle  often  with  a  blackish  wedge  or  streak.  Horizontal  stripe 
on  head  restricted  largely  to  snout  and  upper  part  of  opercle,  usually  developed,  at  least  as  a 
trace  (much  as  in  R.  o.  robustus,  contrasting  with  R.  o.  oligoporus).  Blackening  of  crotches  at 
bifurcation  of  dorsal  and  caudal  rays,  and  occasionally  of  anal  rays,  more  as  in  R.  o.  robustus 
than  in  R.  o.  oligoporus.  Light  streak  above  main  lateral  band,  frequent  in  R.  o.  oligoporus, 
obvious  in  only  a  few  of  the  preserved  specimens.  Form  particularly  distinctive,  unusually 
compressed  for  a  Rhinichthys:  greatest  body  width  steps  over  the  curve  of  the  sides  about 
2.0  times,  rather  than  about  1 .5  times  in  R.  o.  robustus  {R.  o.  oligoporus  approximately  inter- 
mediate). Anterior  profile  less  flattened  than  in  R.  o.  robustus  and  less  arched  than  in  R.  o. 
oligoporus.  Anterior  part  of  head  more  foreshortened  than  in  R.  o.  robustus,  but  rather  more 
pointed  (less  rounded)  than  in  R.  o.  oligoporus.  Mouth  definitely  straighter  than  in  R.  o. 
oligoporus,  but  more  oblique,  rising  forward  to  a  horizontal  through  the  lower  edge  of  the 
eye.  Barbel  almost  invariably  absent,  as  in  the  two  other  subspecies  here  named.  As  in  the 
other  two,  development  of  lateral  line  greatly  reduced— even  more  than  in  R.  o.  oligoporus. 


106 


less  extreme  than  in  R.  o.  reliquus.  The  body  proper,  and  more  strikingly  the  caudal  pe- 
duncle, averaging  deeper  than  in  the  two  other  forms  here  described.  Dorsal  and  anal  fins 
are  inserted  farther  back  than  in  R.  o.  robustus.  The  mouth,  strikingly,  is  strongly  oblique 
and  nearly  straight,  the  upper  jaw  rising  to  about  level  with  middle  of  eye.  Pectoral  rays 
average  few  (12.72).  Vertebrae  and  scale  rows  somewhat  reduced  in  number. 

LITERATURE  CITED 

Hubbs,  C.  L..  and  R.  R.  Miller 

1948.  11.  The  zoological  evidence/Correlation  between  fish  distribution  and  hydrographic  history  in  the 
desert  basins  of  western  United  States.  In,  The  Great  Basin,  with  emphasis  on  Glacial  and  Postglacial 
times.  Bull.  Univ.  Utah,  39(20).  Biol.  Ser.  10(7):  17-166,  figs.  10-29,  map  1. 

In  press.  Hydrographic  history  and  relict  fishes  of  the  north-central  Great  Basin.  Mem.,  California  Acad.  Sci. 


Scripps  Institution  of  Oceanography,  La  Jolla,  California  92037  and  The  University  of 
Michigan,  Museum  of  Zoology,  Ann  Arbor,  Michigan  48104. 


-Wfi-S 


MUS.  CCMP.  ^OOU 
UBRARV 

JUN22t973 

HARVARD 
UNlVERSITYi 


PATTERNS  OF  LARVAL  DEVELOPMENT 
IN  STENOGLOSSAN  GASTROPODS 


GEORGE  E.  RADWIN  AND  J.  LOCKWOOD  CHAMBERLIN 


TRANSACTIONS 

OF  THE   SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  9        12  MARCH  1973 


PATTERNS  OF  LARVAL  DEVELOPMENT 
IN  STENOGLOSSAN  GASTROPODS 

GEORGE  E.  RADWIN  AND  J.  LOCKWOOD  CHAMBERLIN 


ABSTRACT.— Studies  of  egg  capsules  and  the  mode  of  development  in  certain  species  of  stenoglossan  pro- 
sobranchs  tVom  the  northeastern  Gulf  of  Mexico  revealed  an  apparent  disproportionate  number  of  species 
with  non-pelagic  larval  development.  Thorson  ( 1950)  suggested  that  among  shallow-water  marine  in- 
vertebrates incidence  of  pelagic  development  increased  from  the  arctic  to  the  tropics  and  predominated  in 
the  subtropics  and  tropics.  His  conclusions  were  based  largely  on  prosobranch  moUusks.  We  suggest  that 
the  mode  of  early  development  in  the  Stenoglossa  tends  to  follow  phyletic  lines,  regardless  of  latitude  or 
climatic  conditions. 

Many  prosobranch  gastropods  lay  their  eggs  in  parchment-Hke  capsules,  separately 
or  in  clusters,  attached  to  firm  substrata.  Unequivocal  species  identification  is  possible 
when  the  capsules  are  deposited  in  an  aquarium  containing  individuals  of  a  single  species 
or  when  observations  on  ovipositing  snails  are  made.  The  young  of  most  higher  pro- 
sobranchs  pass  the  veliger  stage  within  the  capsule  and  may  be  sufficiently  developed  be- 
fore emergence  to  be  identifiable  either  by  the  sculpture  of  the  early  teloconch  sculpture 
or  by  the  radular  dentition.  In  other  groups  the  young  are  released  as  veligers  and  are  car- 
ried in  the  plankton  until  they  settle  and  metamorphose.  Pearse  (1969)  described  a  third 
mode  of  development  which  seems  to  be  intermediate  between  these  two.  In  this  type  a 
modified  veliger  (called  by  some  authors  a  veUconcha)  emerges  from  the  egg  capsule  and 
swims  feebly  for  a  short  time  in  the  bottom-most  layer  of  water  before  settling.  He  has 
called  this  a  demersal  mode  of  development.  The  only  stenoglossan  species  we  know  to 
exhibit  this  kind  of  development  is  Olivella  verreauxi  (Duclos). 

Identification  of  capsules  of  marine  gastropod  species  can  contribute  to  distributional 
data  which  may  be  useful  in  zoogeographic  studies  and  may  serve  as  an  ecological  tool  in 
determining  the  reproductive  range  of  a  species.  In  addition,  their  use  as  taxonomic  char- 
acters at  the  generic  level  should  be  considered. 

In  this  paper  the  spawning  conditions  and  egg  capsules  of  nine  species  of  stenoglos- 
san mollusks  from  the  northeastern  Gulf  of  Mexico  are  described.  These  observations 
were  made  from  March  1963  to  July  1964.  The  species  treated  are:  Phyllonotus  pomum 
(Gmelin),  Muricanthus  fulvescens  (Sowerby),  Calotrophon  ostrearum  (Conrad),  JJrosal- 
pinx  tampaensis  (Conrad),  Urosalpinx perrugata  (Conrad),  Thais  floridana  (Conrad),  Can- 
tharus  cancellarius  (Conrad),  Cantharus  multangulus  (Philippi),  and  Pollia  tincta  (Con- 
rad). 

SPAWNING  SITES,  EGG  CAPSULES,  AND  LARVAL  DEVELOPMENT 

Phyllonotus  pomum  (Gmelin,  1791)  (Fig.  1,  la).  Localities:  St.  Teresa  and  Bay  Mouth 
Bar,  Franklin  Co.,  Fla.,  attached  to  large,  empty  bivalve  shells.  Period:  May-July.  The 
capsules  are  deposited  in  irregular  compact  masses  up  to  30  cm  across;  individual  cap- 
sules are  superficially  similar  to  those  of  Buccinum  and  Neptunea.  From  two  to  five  larvae 
develop  in  each. 

Tryon  (1880),  Webb  (1942),  and  Perry  and  Schwengel  (1955)  described  and  figured 
the  capsule  mass  of  P.  pomum,  and  Webb  reported  communal  spawning  by  as  many  as 
twenty-five  females.  This  egg  mass  is  similar  to  that  reported  for  Murex  senegalensis  (see 
Knudsen,  1950).  D'Asaro  (1970b)  reported  non-pelagic  development  for  P.  pomum. 

Muricanthus  fulvescens  (Sowerby,  1834)  (Fig.  5).  Locality:  St.  Andrews  State  Park, 
Bay  Co.,  Fla.,  attached  to  rocks  of  the  breakwater.  Period:  June-August.  Capsules  depos- 
ited in  clusters  with  their  bases  fused.  Each  capsule  is  a  flattened  cylinder  about  25  mm 

SAN  DIEGOSOC.  NAT.  HIST..  TRANS.  17(9):  107-118.  12  MARCH  1973 


108 


high,  with  the  top  broader  than  the  base.  All  were  empty  when  collected.  Identification 
was  made  on  the  basis  of  a  laying  female  and  an  egg  mass  (catalogue  no.  599643)  in  the 
collection  of  the  Division  of  Mollusks,  U.S.  National  Museum  of  Natural  History.  Moore 
(1961:  26)  figured  a  similar  capsule  collected  off  Mississippi  as  M.  fiilvescens.  He  gave  the 
height  of  "one  typical  specimen"  as  14  mm  which,  from  the  examples  we  have  seen, 
seems  to  be  too  small.  He  also  reported  that  there  are  over  one  hundred  eggs  in  each  cap- 
sule. 

Calotrophon  ostrearum  (Conrad,  1846)  (Fig.  7,7a).  Localities:  1)  St.  Teresa.  Franklin 
Co.,  Fla.,  on  the  blades  of  turtle  grass;  2)  dredged  in  13  m  off"  Dog  Island,  Franklin  Co., 
Fla.,  attached  to  the  sides  of  egg  capsules  of  Ficus  communis  (Roding);  and  3)  attached  to 
the  walls  of  aquaria  in  which  specimens  of  C  ostrearum  were  isolated  (see  Radwin  and 
Wells,  1968).  Period:  early  May  to  mid-June.  Numerous  capsules  are  laid  individually, 
their  bases  separated;  they  are  roughly  semicircular,  average  about  4  mm  high,  and  when 
first  deposited  usually  contain  3  to  5  large,  spherical,  reddish  eggs.  Emergence  is  in  the 
crawling  stage  (pelagic  stage  absent).  Egg  capsules  apparently  of  this  species  were  attrib- 
uted by  Perry  and  Schwengel  (1955)  to  both  Urosalpinx  perrugata  and  Cantharus  florid- 
anus. 

Urosalpinx perrugata  (Conrad,  1846)  (Fig.  2).  Localities:  1)  Bay  Mouth  Bar.  Alligator 
Harbor,  Franklin  Co.,  Fla.,  attached  to  empty  mollusk  shells;  2)  attached  to  the  sides  and 
bottoms  of  aquaria  in  which  adults  were  isolated  (see  Radwin  and  Wells,  1968).  Period: 
late  April  to  mid-June.  Numerous  erect  capsules,  with  fused  bases,  are  deposited  in  a  mat; 
the  capsules,  about  10  mm  high,  are  inversely  pyramidal  and  have  two  lateral  alae  and 
apical  protuberances.  An  egg  mass  may  contain  as  many  as  200  capsules.  A  large  but  un- 
determined number  of  eggs  is  initially  deposited;  the  majority  are  apparently  nurse-eggs, 
since  only  5  to  15  larvae  develop  fully.  Larvae  emerge  in  the  crawling  stage  (pelagic  stage 
absent).  Egg  capsules  of  this  species  are  misidentified  in  Perry  and  Schwengel  (1955)  as 
the  product  of  Nassarius  vibex. 

Urosalpinx  tampaensis  (Conrad,  1846)  (Fig.  3).  Locality:  Attached  to  the  floor  of  an 
aquarium  in  which  individuals  of  this  species  were  isolated  (see  Radwin  and  Wells,  1968). 
Period:  March  (in  aquarium).  The  erect  egg  capsules,  about  8  mm  high,  are  deposited 
singly.  They  resemble  plump  fingers  on  stalks  and  are  more  similar  to  those  of  Eupleura 
sulcidentata  (see  Perry  and  Schwengel,  1955)  than  to  those  of  the  other  two  western  Atlan- 
tic species  of  Urosalpinx  (cinerea  and  perrugata).  Each  capsule  contains  numerous  eggs 
which,  in  our  material,  did  not  develop. 

Thais  floridana  (Conrad,  1837)  (Fig.  4,4a).  (For  characters  distinguishing  this  species 
from  T.  haemastoma,  see  Radwin  and  Wells,  1968.)  Locality:  St.  Andrews  State  Park,  Bay 
Co.,  Fla.,  attached  to  empty  bivalve  shells  and  rocks  of  the  breakwater.  Period:  July- 
August.  The  elongate,  trough-shaped  capsules  are  about  12  mm  high,  have  apical  escape 
pores,  and  are  deposited  in  large  masses.  The  capsules  at  the  base  of  a  mass  tend  to  be 
nearly  erect  and  are  attached  side  by  side  to  the  substratum,  with  their  bases  fused.  Other 
capsules  are  attached  to  those  beneath  in  an  arborescent  pattern. 

Burkenroad  (1931)  figured  a  capsule  mass  and  commented  on  the  hatching  process. 
D'Asaro  (1966),  who  figured  the  capsule  and  described  the  spawning  and  embryology  in 
detail,  reported  communal  spawning  occurring  from  February  through  November  at 
Miami,  Fla.  He  suggested  that  spawning  "probably  occurs  also  in  December  and  January 
when  the  temperature  is  above  average."  A  shorter  spawning  season  in  the  northeastern 
Gulf  of  Mexico  is  consistent  with  the  shorter  period  of  warm  water  temperature  there. 
Large  numbers  of  veligers  emerge  and  have  a  prolonged  pelagic  development  (D'Asaro, 
1966).  This  mode  of  development  (also  reported  by  other  workers  for  this  species  in  North 
American  waters)  contrasts  with  that  of  most  stenoglossans  treated  in  this  paper. 

Thorson  ( 1946,  1950)  cited  T.  floridana  as  having  pelagic  development  in  some  partSi 
of  its  range  and  direct,  non-pelagic  development  in  others.  This  may  be  correct,  but  hisj 
evidence  is  apparently  inferred  from  Lamy  (1928),  who  referred,  in  turn,  to  Korschelt  and! 
Heider  (1900),  which  reference  we  have  not  seen.  Lamy  reported  only  that  many  of  the! 
larvae  die  after  cleavage  and  are  then  eaten  by  the  others  in  the  capsule.  Although  this! 
"nurse-egg"  type  of  feeding  is  usually  associated  with  non-pelagic  larval  development,  itj 


109 


Figure  1.  PhvUonmus  pomiim—^even  capsules  from  an  egg  mass.  la.  P.  pomtimsmaW  egg  mass.  2.  Urosalpinx 
perrugaiiisingk  egg  capsule.  3,  Urosalpinx  iampaensis—^\n^\c  egg  capsule.  4,  Thais  fhridana—single  egg  cap- 
sule. 4a,  Thais  fJoridana-ponion  of  an  egg  mass.  5,  Muricanihi/s  fulvescens-single  egg  capsule.  6.  Cantharus 
nnilranguhis— single  egg  capsule.  6a.  Cantharus  multanguhis~\.op  view  of  a  single  egg  capsule.  7.  Caloirophon  os- 
/rertn///;— single  egg  capsule.  7a.  Caloirophon  osirearum—side  view  of  a  single  egg  capsule.  8,  Cantharus  cancel- 
/ani/.s-single  egg  capsule.  8a.  Cantharus  cancellarins-lop  view  of  a  single  egg  ca'psule. 


110 


is  not  proof  of  such  development,  as  Thorson  ( 1950)  pointed  out  for  Natica  catena. 

Caniharus  cancellarius  (Conrad,  1846)  (Fig.  8,  8a).  Localities:  1)  Bay  Mouth  Bar,  Al- 
ligator Harbor,  Franklin  Co.,  Fla.,  on  empty  mollusk  shells;  2)  Seahorse  Key,  Cedar 
Keys,  Levy  Co.,  Fla.,  on  stones  and  empty  mollusk  shells;  and  3)  attached  to  the  sides  of 
aquaria  in  which  adults  were  isolated  (Radwin  and  Wells,  1968).  Period:  early  May  to  late 
June.  The  erect  capsules  are  deposited  in  a  mat  with  their  bases  confluent.  In  nature  the 
mats  contained  15-20  capsules;  the  number  of  capsules  laid  in  aquaria  was  smaller.  Indi- 
vidual capsules  are  roughly  rectangular,  have  four  distinctive  spinose  projections  at  the 
top,  and  are  about  4  mm  high.  In  each  capsule  approximately  10-20  larvae  develop  to  the 
crawling  stage. 

Moore  (1961:26)  figured  a  capsule  of  this  species  as  Cantharus  reticidatus.  He  also 
noted  that  on  the  Mississippi  coast  "these  capsules  are  rather  common  objects  during 
March,  April,  and  May,"  and  that  from  one  capsule  "a  dozen  or  more  eggs  hatch  out 
while  still  in  the  veliger  stage."  We  have  seen  no  other  report  of  pelagic  development  in 
this  species  or  elsewhere  in  the  entire  family  Buccinidae. 

Cantharus  tmdtanguhis  (Philippi,  1849)  (Fig.  6.  6a).  Capsules  illustrated  in  Perry  and 
Schwengel  (1955),  fig.  340.  Localities:  1)  Bay  Mouth  Bar,  Alligator  Harbor,  Franklin  Co., 
Fla.;  2)  St.  Teresa,  Franklin  Co.,  Fla.;  and  3)  deposited  on  the  floor  of  aquaria.  The  cap- 
sules collected  in  the  field  were  on  shells  and  turtle  grass.  Period:  May-July.  Each  capsule 
is  inversely  pyramidal  and  about  4  mm  high;  the  top  surface  bears  four  spine-like  projec- 
tions. The  capsule  mass  is  a  mat  formed  by  the  confluent  bases  of  the  capsules.  When  first 
deposited  each  capsule  contains  8-20  flesh-colored  eggs,  a  number  of  which  apparently 
serve  as  nurse-eggs,  as  only  a  few  crawling-stage  larvae  eventually  emerge  from  each  cap- 
sule. 

Pollia  tincta  (Conrad,  1846)  (see  Perry  and  Schwengel,  1955;  Lebour,  1945).  Local- 
ities: St.  Teresa,  Franklin  Co.,  Fla.,  and  Seahorse  Key,  Cedar  Keys,  Levy  Co.,  Fla.,  on 
shells  and  small  rocks.  Period:  June-July.  Clusters  of  several  capsules  are  deposited,  each 
about  5  mm  high,  broadly  goblet-shaped  and  basally  pedunculate.  Each  capsule  contains 
5  to  15  eggs,  which,  in  our  material,  did  not  hatch.  Lebour  (1945)  described  the  larval  de- 
velopment as  non-pelagic.  Generic  distinction  o^  Cantharus  and  Pollia  (as  Pisania),  based 
on  radular  dentition  (see  Troschel,  1866),  is  corroborated  by  differences  in  egg  capsule 
morphology.  Cantharus  capsules  are  four-sided  and  rectangular  or  inversely  pyramidal, 
with  a  flat  top.  Pollia  capsules  are  goblet-shaped. 

DISCUSSION 

The  nine  species  studied  belong  in  either  the  family  Muricidae  (six  species)  or  the 
Buccinidae  (three  species),  and  constitute  a  majority  of  these  families  reported  to  live  in 
the  area  of  field  work  (Perry  and  Schwengel,  1955).  The  two  families  are  both  in  the  sub- 
order Stenoglossa,  order  Neogastropoda. 

Among  shallow-water,  benthic,  marine  invertebrates,  Thorson  ( 1950)  found  that  spe- 
cies with  pelagic  larval  stages  were  rare  in  polar  regions  but  increased,  and  indeed  pre- 
dominated toward  the  tropics.  This  conclusion  was  based  primarily  on  samples  of  pro- 
sobranch  mollusks  from  several  widely  separated  areas.  However,  our  data  and  those  of 
D'Asaro  (1970)  indicate  that  at  least  in  the  stenoglossans,  non-pelagic  forms  of  devel- 
opment may  be  more  common  in  tropical  waters  than  is  generally  recognized.  Thorson's 
data  demonstrate  a  substantial  increase  in  the  percentage  of  species  with  pelagic  devel- 
opment from  arctic  to  temperate  waters  (0%  in  East  Greenland  to  63.5%  in  southern  Eng- 
land) but  they  show  a  much  smaller  increase  in  percentage  from  temperate  to  tropical  wa- 
ters (e.g.  southern  England  to  a)  Canary  Islands,  4.5%;  b)  Persian  Gulf,  11.5%;  c) 
Bermuda,  21.5%).  These  facts  have  led  us  to  question  whether  the  proportional  increase 
implied  by  Thorson  ( 1950)  is  demonstrable  in  lower  latitudes. 

A  review  of  the  literature  on  modes  of  larval  development  among  marine  pro- 
sobranchs  shows  that  in  the  Archaeogastropoda  there  is  no  clear  predominance  of  either 
pelagic  or  non-pelagic  development.  In  the  Mesogastropoda,  however,  pelagic  devel- 
opment predominates.  Within  the  Neogastropoda  the  suborder  Toxoglossa  exhibits  pela- 
gic larval  development,  whereas  the  suborder  Stenoglossa  is  the  only  major  prosobranch 


Ill 


group  in  which  non-pelagic  larval  development  seems  to  clearly  predominate  (Table  1). 

The  apparent  predominance  of  non-pelagic  development  in  the  Stenoglossa.  regard- 
less of  latitude,  as  well  as  the  abundance  of  species  of  this  suborder  in  lower  latitudes  sug- 
gests that  the  Stenoglossa  were  under-represented  in  at  least  some  of  the  areas  discussed 
by  Thorson.  The  Bermudas,  the  Canaries,  and  the  Persian  Gulf  are  not  typical  of  the 
main  tropical  and  subtropical  shelf  regions  of  the  world.  The  first  two  are  small  island 
groups,  separated  from  the  adjacent  mainland  by  deep  water  (over  1.000  m),  and  the 
third  is  a  hypersaline  body  of  water  with  excessively  high  water  temperatures  (Mohr, 
1929)  and  a  restricted  outlet  to  the  Indian  Ocean. 

Bermuda— Lehoufs  (1945)  data,  on  which  Thorson  (1950)  based  his  estimate  of  85% 
of  Bermudan  species  having  pelagic  development,  are  biased  toward  species  with  pelagic 
development,  as  her  study  was  based  principally  on  plankton  samples.  Only  29  of  her 
prosobranch  species  were  sufficiently  identified  to  be  used  in  a  calculation.  Of  these,  only 
three  (10%)  are  stenoglossans;  two  have  non-pelagic  development.  All  26  of  the  non-sten- 
oglossans  have  pelagic  development. 

The  actual  percentage  of  Bermudan  prosobranchs  with  pelagic  development,  though 
apparently  less  than  85%  may,  nevertheless,  be  higher  than  is  typical  of  tropical  and  sub- 
tropical western  Atlantic  areas.  Evidence  for  this  supposition  stems  from  the  fact  that 
stenoglossans  make  up  a  smaller  percentage  of  total  prosobranchs  at  Bermuda  than  is 
typical  of  other  similar  areas.  Peile  (1927)  listed  215  Bermudan  species  of  marine  pro- 
sobranchs, excluding  abyssal  species,  of  which  21%  are  stenoglossans.  In  comparison,  fau- 
nal  lists  for  the  adjacent  mainland  and  Caribbean  island  areas  give  the  following  percent- 
ages of  stenoglossans:  western  Florida,  28%  (Perry  and  Schwengel,  1955);  West  Indies,  29- 
32%  (Arango,  1878;  Dall  and  Simpson,  1901;  Morch,  1878);  Brazil,  32%  (Lange  de  Mor- 
retes,  1949). 

Canary  Islands— Thorson  (1950)  reported  that  68%  of  the  Canary  Islands  marine 
prosobranchs  exhibit  pelagic  development.  Faunal  lists  for  these  islands  and  for  the  adja- 
cent coast  of  western  Africa  indicate  a  situation  parallel  to  that  in  Bermuda,  with  fewer 
stenoglossans  among  marine  prosobranchs  at  the  islands  than  at  the  mainland  areas:  Ca- 
naries, 30%  (Dautzenberg,  1890,  1891);  western  Africa,  37%  (Nickles,  1950).  Sao  Thome, 
in  a  more  tropical  position  off' the  western  coast  of  Africa,  has  an  essentially  similar  situa- 
tion; 28%  of  the  marine  prosobranchs  are  stenoglossan  (Tomlin  and  Shakleford,  1923). 

Evidence  of  a  lower  percentage  of  prosobranchs  with  non-pelagic  larval  development 
at  Bermuda,  the  Canaries,  and  Sao  Thome  is,  in  itself,  of  biogeographical  and  ecological 
interest.  The  faunal  Hsts  cited  above  show  that  the  marine  moUusks  of  these  islands  in- 
clude few  endemics.  Such  low  endemism  is  evidence  of  recent  faunal  origin  by  immigra- 
tion. The  marine  molluscan  fauna  of  Bermuda  is  considered  a  depauperate  Antillean 
fauna  (Warmke  and  Abbott,  1961),  and  the  prosobranchs  of  the  Canaries  and  Sao  Thome 
are  just  as  clearly  depauperate  western  African.  The  colonization  of  these  islands  largely 
by  species  with  pelagic  larvae  could  be  attributed  to  their  ability,  as  larvae,  to  traverse  the 
geographical  and  bathymetric  barriers  isolating  the  islands  from  the  mainland. 

Persian  Gulf.— Thorson  (1940a,  1950)  found  that  75%  of  the  prosobranch  species 
studied  from  the  Persian  Gulf  had  pelagic  development.  His  data  seems  moderately 
biased  toward  such  species  as  only  24%  of  them  (5  of  21  species)  were  stenoglossans.  Mel- 
vill  and  Standen  (1901)  and  Melvill  (1928)  indicate  that  just  over  30%  of  the  marine  pro- 
sobranchs from  this  area  are  stenoglossans. 

In  view  of  Thorson's  original  data  showing  only  a  small  increase  in  the  percentages 
of  prosobranch  species  with  pelagic  larval  development  from  temperate  to  tropical  wa- 
ters the  question  arises  whether  any  significant  increase  exists.  Regardless  of  the  answer  to 
this  question— and  our  evidence  is  not  enough  to  resolve  it— there  remams  the  question  of 
why  a  steep  gradient  exists  in  higher  latitudes  but  only  a  weak  one  (if,  indeed,  any  exists) 
in  lower  latitudes.  Of  course,  data  on  larval  ecology  and  distribution  must  include  other 
invertebrate  groups  as  well. 

After  a  draft  of  this  paper  was  sent  to  Thorson  in  1968,  he  informed  us  (in  litt.)  that 
the  data  he  had  compiled  on  stenoglossan  early  development,  more  extensive  than  the 
data  in  Table  I,  suggest  an  appreciably  lower  percentage  of  species  with  non-pelagic  de- 


112 


TABLE  I 
OCCURRENCE  OF  PELAGIC  AND  NON-PELAGIC  LARVAL  DEVELOPMENT  WITHIN  THE  STENOGLOSSA 


Superfamily 

Family 

No.  Species  With 

Genus 

Pelagic  Larvae 

Muricacea 

Rapanidae 

Rapana 

3 

Muricidae 

Murex 

4 

Chicoreus 

— 

Phvllonotus 

3 

Boreotruphon 

— 

Caloirophon 

— 

Bedevina 

1 

Bedeva          \ 
Favartia         \ 

— 

— 

Viiularia 

1 

Ceratosioma 

— 

Ocenehra               \ 

— 

Vrosalpinx              \ 

— 

Eupleura 

\                    — 

Thaididae 

\ 

Purpura 

\               1 

Neptunea 

\            — 

Siphonalia 

\         — 

Pollia 

\       — 

"Cantharus" 

\    — 

Buccinum 

\  — 

Volutharpa 

V- 

Macron 

—V 

Chauvetia 

— \ 

Melongenidae 

\ 

Melongena 

—    \ 

Syrinx 

—       \ 

Busycon 

\ 

Hemifusus 

— 

Fascioiariidae 

Leucozonia 

— 

Peristernia 

— 

Fasciolaria 

— 

Pleuroploca 

— 

Fusinus 

— 

Troschelia 

— 

Volutacea 

Volutidae 

Valuta 

— 

Thais 

9 

Nucella 

— 

Buccinacea 

Columbellidae 

Pyrene 

— 

Milrella 

1 

Anachis 

5 

Zafrona 

1 

Astvris 

— 

Amphissa 

1            / 

Columhella 

2         / 

Nassariidae 

/ 

Nassarius 

II     / 

Trilia 

2  / 

llyanassa 

1/ 

Buccinidae 

/ 

Beriiii^ius  (Jumahil 

/— 

Volutopsius 

/ — 

Pyrulofusus 

/  — 

Colus 

/     — 

Plicifusus 

/       — 

A  lei  1  hoe 

/         — 

Melo 

/           — 

Cyniha 

/              — 

Cymhiola 

/                — 

Marginellidae 

/ 

Persicula                     . 

/                    — 

Mar^inella              / 

— 

Prunum                 / 

— 

Cancellariidae            / 

( aneellaria       / 

— 

Admete          / 

— 

Vasidae        / 

Vasum        / 

— 

Mitridae          / 

Siri^atella 

3 

Atriniilra 

1 

Turbincllidae 

lurhinella  (XancusI 

— 

Olividae 

Ancilla 

— 

Olivella 

— 

Oliva 

1 

X  — Jt.  Allison  Kay,  personal 

communication 

XX  -/this  paper 

No.  Species  With 
Non-pelagic  Larvae 


Reference 


18,37.83 


45,80 

6,15.24.26,43.45,49,63,83 

30,39,47 


3,17.30,41,42,45,63.79.83.92.93 

3.5 


'^ 


21,30,50,82 

30,85 

22,32    . 

30.82,84,xx 

50  \ 

33  \ 

6,20.38,88 

51 

9,45 

43 
43 
26 

43 
82 

26 

26.66 
16 

26 

63 

6,30,53,67 

65 


12 


TABLhI 
OCCURRENCE  OF  PELAGIC  AND  NON-PELAGIC  LARVAL  DEVELOPMENT  WITHIN  THE  STENOGLOSSA 


Superlamily 
lamily 
(ienus 

Muricacea 

Rapanidac 

Rapana 
Muricidae 

Murex 

Chicoreus 

Phvllonoius 

Boreoirophon 

Cahlrophon 

Bedevina 

Bedeva 

Favartia 

Vitularia 

Ceratostoma 

Ocenehra 

Urosalpinx 

Eupleura 
Thaididae 

Purpura 

Thais 

Nucella 
Buccinacea 

Columbellidae 

Pyrene 

Miirella 

Anachis 

Zafrona 

Aslyris 

Amphissa 

Columbella 
Nassariidae 

Nassarius 

Tritia 

llyanassa 
Buccinidae 

Benn^ius  (Jumalaj 

Voluiopsius 

Pyrulofusus 

Colus 

Plicifusus 

Neptunea 

Siphonalia 

Pallia 

"Canlharus" 

Buccinum 

Volulharpa 

Macron 

Chau  vetia 
Meiongenidae 

Melon^ena 

Syrinx 

Busycon 

Hemifusus 
Fasciolariidae 

Leucozonia 

Peristernia 

Fasciolaria 

Pleuropluca 

Fusinus 

Troschelia 
Voiutacea 

Voiulidae 

Valuta 

Alciihae 

Mela 

Cvniha 

Cymbiala 
Margineliidae 

Persicula 

Marginella 

Prunum 
Cancellariidae 

Cancellaria 

Admeie 
Vasidae 

Vasurri 
Mitridae 

Sirigaiella 

A  trim  it  ra 
Turbineilidae 

Turbinella  IXancus  I 
Olividae 

Ancilla 

Olivella 

Oliva 


No.  Species  With 
Pelagic  Larvae 


No.  Species  With 
Non-pelagic  Larvae 


I 

5 
I 

I 

2 

II 


2 
9 
1 

4 
I 
I 

2 
8 
I 
I 
I 

I 
I 

3 
1 

2 
I 

3 
2 
2 
I 


Reference 


18.37,83 

15.43,63,75,83 
26,62,76 
26,83 
30,50.55.85 

XX 

61 

7 

74 

25 

6 

30 

ll,34,xx 

II 

49 

6,15,24,26,43,45,49,63,83 

30,39,47 


72 
6,63 

6,25.54.78 

4.6 

82.85 

70 

43,72 

3. 1  7.30.4 1 .42.45.63.79.83.92.93 

3,5 

79 

21.30,50,82 

30,85 

22,32 

30,82,84,xx 

50 

6,30,40 

81 

48 

XX 

30,84 
30 
14 
30 

35.45,71 
36,6! 
45.7l,xx 
6 

26,49,75 

X 

26,45.xx 
45.75 
6.14.43 
30 


45,80 

33 

6,20,38.88 

51 

9,45 

43 
43 
26 

43 
82 

26 

26.66 
16 

26 

63 

6.30,53,67 

65 


X  —  E.  Allison  Kay,  personal  communication 


113 


velopment  (62%  compared  to  our  72%).  Thorson's  reasons  for  believing  "that  the  species 
with  a  non-pelagic  development  predominate  more  in  available  data  than  they  do  in  na- 
ture" are  1)  these  species  have  egg  capsules  which  are  large,  conspicuous,  and  easy  to  dis- 
cover; 2)  they  tend  to  be  discovered  more  often  with  their  capsules  than  do  species  with 
pelagic  development  because  they  have  a  longer  spawning  season;  3)  the  capsules  are  eas- 
ier to  identify  to  species  and  4)  his  experience  at  the  Canary  Islands  and  in  Thailand  in- 
dicates "that  most  species  with  pelagic  development  there  will  reproduce  in  the  hottest 
season  of  the  year,"  whereas  "biologists  tend  to  make  expeditions  to  such  places  at  the 
cooler  times  of  the  year."  Correction  for  these  biases  would  lower  Thorson's  entire 
gradient  of  pelagic  vs.  non-pelagic  development,  except  for  the  Arctic,  where  we  have 
seen  no  evidence  to  indicate  the  existence  of  pelagic  development  among  stenoglossans; 
thus  the  slope  of  the  gradient  probably  would  be  increased  from  high  to  mid-latitudes. 
We  would  not,  however,  expect  the  slope  to  be  changed  much  from  mid-  to  low  latitudes 
by  corrections  for  any  of  the  sources  of  bias  suggested  by  Thorson,  except  his  last  one, 
which  would,  in  theory,  result  in  some  steepening. 

SELECTIVE  ADVANTAGE  OF  NON-PELAGIC  LARVAL  DEVELOPMENT 

The  apparent  predominance  of  non-pelagic  development  in  the  Stenoglossa  has  ne- 
cessitated a  more  detailed  review  of  early  development  in  this  group  (Table  1).  The  mode 
of  larval  development  in  the  Stenoglossa  seems  generally  to  follow  phyletic  lines,  regard- 
less of  latitude  or  climatic  conditions  (beginning  with  the  Buccinidae  pelagic  development 
is  almost  unknown).  Exceptions  include  the  Nassariidae,  in  which  pelagic  development  is 
clearly  predominant  and  the  Mitridae,  whose  wide  distribution  in  the  Indo-west  Pacific 
(Cernohorsky.  1965)  suggests  that  the  pelagic  mode  of  development  predominates.  We 
cannot  explain  these  apparent  inconsistencies  on  the  basis  of  our  data. 

Thorson  (1950)  argued  that  pelagic  development  is  disadvantageous  in  the  Arctic  be- 
cause the  period  of  rich  plankton  production  on  which  most  pelagic  larvae  depend  for 
food  is  too  short.  For  the  lower  latitudes,  where  both  modes  of  development  are  practical, 
the  problem  remains. 

Garstang  (1928)  and  Thorson  (1950)  showed  that  pelagic  development  permits  rapid 
dispersal,  repopulation  of  depleted  areas,  and  establishment  of  dense  populations  when 
the  larvae  encounter  optimal  conditions.  By  contrast,  non-pelagic  larvae  tend  to  remain  in 
established  optimal  situations,  are  not  as  numerous  as  pelagic  larvae,  and  are  provided 
with  protection  and  a  large  food  supply  by  parental  brooding.  This  mode  inhibits  rapid 
dispersal,  repopulation  of  depleted  areas,  and  short-term  establishment  of  dense  popu- 
lations. 

There  is  little  information  on  the  advantages  of  the  various  modes  of  larval  devel- 
opment to  marine  prosobranchs  and  other  marine  invertebrates  of  shallow  waters.  Thus, 
the  selective  advantage  of  non-pelagic  larval  development  in  the  stenoglossans  is  not 
clearly  understood.  However,  most  stenoglossans  are  carnivorous  and,  therefore,  occupy 
relatively  high  trophic  levels  in  their  ecosystems.  It  seems  reasonable  to  suggest  that  these 
animals  are  probably  food-limited.  Thus,  it  may  be  more  advantageous  for  stenoglossans 
to  use  their  energy  in  producing  relatively  few,  non-pelagic  young  that  can  utilize 
"proved"  local  food  resources,  than  to  adopt  the  alternative  strategy  of  producing  vast 
numbers  of  highly  vagile  young  that  must  find  suitable  conditions  to  insure  survival. 

ACKNOWLEDGMENTS 

This  report  is  based  in  part  on  a  thesis  submitted  by  the  senior  author  to  Florida  State  University  in  partial 
fultiliment  of  the  requirements  for  the  Master  of  Science  degree.  Dr.  Harry  W.  Weils  directed  the  thesis  research. 
Drs.  Joseph  Rosewater  and  Harold  A.  Rehder.  Division  of  Mollusks.  U.  S.  National  Museum  of  Natural  His- 
tory, offered  constructive  ideas  and  suggestions.  Mr.  Anthony  D'Attilio  re-drafted  the  original  illustrations. 

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61.  Murray,  F.V. 

1964.  The  spawn  of  some  Australian  marine  prosobranch  mollusks.  Austr.  Nat.  Hist.  14:  405-408. 

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1963.  Some  observations  on  the  egg  capsules  and  embryos  of  Torvamiirex  teniius  (Reeve,  1845).  J.  Ma- 
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1957.  Studies  on  the  egg  masses  and  larval  development  of  some  prosobranchs  from  the  Gulf  of  Man- 
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64.  Nickles,  M. 

1950.  Mollusques  testaces  marins  de  la  cote  occidentale  d'Afrique.  Manuels  Ouest  Africains.  Vol.  2. 
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65.  Olsson,  A. A.,  and  M.E.  Crovo 

1968.  Observations  on  aquarium  specimens  of  Oliva  sayana  Ravenel.  Veliger  1 1 :  3 1-32. 

66.  Ostergaard,  J.M. 

1950.  Spawning  and  development  of  some  Hawaiian  marine  gastropods.  Pacific  Sci.  4:  75-1 15. 

67.  Paine.  R.T. 

1962.  Reproduction  of  0//vt^//(/ /?;;///ra.  Nautilus  75:  139-142. 

68.  Pearse.  J.S. 

1969.  Slow  developing  demersal  embryos  and  larvae  of  the  antarctic  sea  star  Odouiaster  validtis.  Mar. 
Biol.  3:  110-116. 

69.  Peile,  A.J. 

1927.  The  Mollusca  of  Bermuda.  Proc.  Malac.  Soc.  London  17:  71-98. 

70.  Pelseneer,  P. 

1906.  Biscayan  plankton  collected  during  the  cruise  of  H.M.S.  "Research"  1900.  VII-MoUusca.  Trans. 
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71.  Perry,  L.M.,  and  J.S.  Schwengel 

1955.  Marine  Shells  of  the  Western  Coast  of  Florida.  Paleont.  Res.  Inst.,  Ithaca,  N.Y.  318p. 

72.  Petit,  G.,  and  J.  Risbec 

1929.  Sur  la  ponte  de  quelques  gastropodes  prosobranches.  Bull.  Soc.  Zool.  France  54:  564-570. 

73.  Radwin,G.E.,  and  H.W.Wells 

1968.  Comparative  radular  morphology  and  feeding  habits  of  muricid  gastropods  from  the  Gulf  of 
Mexico.  Bull.  Mar.  Sci.  18:  72-85. 

74.  Raeihle,  D. 

1966.  An  ob.servation  on  captive  Murex  cellulosus  Conrad.  Amer.  Malac.  Union,  Ann.  Rept.  for  1966, 
p.  28. 

75.  Risbec,  J. 

1931.   Note  sur  le  reproduction  de  quelques  pro.sobranches  neo-Caledoniens.  Ann.  Inst.  Oceanog.  10: 
23-33. 

76.  Risbec,  J. 


17 


1932.  Notes  sur  la  ponte  et  le  developpement  de  mollusques  gasteropodes  de  Nouvelle-Caledonie. 
Bull.  Soc.  Zool.  France  57:  358-374. 

77.  Scheltema.  R.S. 

1965.  The  relationship  of  salinity  to  larval  survival  and  development  in  Nassarius  obsoletus  (Gastro- 
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78.  Scheltema,  R.  and  A.H.  Scheltema 

1963.  Pelagic  larvae  of  New  England  intertidal  gastropods  W—Anachis  avara.  Hydrobiologia  22:  85-91. 

79.  Scheltema,  R.,  and  A.H.  Scheltema 

1964.  Pelagic  larvae  of  New  England  intertidal  gastropoda  \\\— Nassarius  triviitaius.  Hydrobiologia  25: 
321-329. 

80.  Strebel.  H. 

1905.  Beitrage  zur  kenntnis  der  molluskenfauna  der  Magalhaenprovinz.  Zool.  Jahrb.  24:  91-174. 

81.  Taki,  Iw. 

1934.  Notes  on  the  shells  and  egg-capsules  oi  Siphonalia  fusoides  (Reeve).  Venus  4:  331-334. 

82.  Thorson,  G, 

1935.  Studies  on  the  egg-capsules  and  development  of  arctic  marine  prosobranchs.  Medd.  om  Gron- 
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83.  Thorson,  G. 

1940a.  Studies  on  the  egg  masses  and  larval  development  of  Gastropoda  from  the  Iranian  Gulf  Danish 
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84.  Thorson,  G. 

1940b.  Notes  on  the  egg-capsules  of  some  north  Atlantic  prosobranchs  of  genus  Troschelia.  Chryso- 
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1944.  Marine  Gastropoda  Prosobranchiata.  Medd.  om  Gronland  121:  1-181. 

86.  Thorson,  G. 

1946.   Reproduction  and  larval  development  of  Danish  marine  bottom  invertebrates.  Medd.  Komm. 
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1950.  Reproductive  and  larval  ecology  of  marine  bottom  invertebrates.  Biol.  Rev.  25:  1-45. 

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Dept.  of  Marine  Invertebrates,  Natural  History  Museum,  P.O.  Box  1390,  San  Diego, 
California,  92112,  and  Environmental  Oceanographic  Research  Program,  National  Marine 
Fisheries  Service,  Washington,  D.  C. 


I 


S-Aifi'S 


MUS.  COMP.  200L. 
LIBRARY 

HARVARD 
UNIVERSITY. 


A  MARINE  INVERTEBRATE  FAUNULE 
FROM  THE  LINDAVISTA  FORMATION, 
SAN  DIEGO,  CALIFORNIA 


GEORGE  L.  KENNEDY 


TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 


VOL.  17,  NO.  10  28  MARCH  1973 


I 


A  MARINE  INVERTEBRATE  FAUNULE 
FROM  THE  LINDAVISTA  FORMATION, 
SAN  DIEGO,  CALIFORNIA 

GEORGE  L.  KENNEDY 


ABSTRACT.— A  small  mainly  molluscan  invertebrate  fauna,  dominated  by  the  Pismo  clam  Tivela  stulto- 
rum  (Mawe),  occurs  in  the  reportedly  unfossiliferous  Lindavista  Formation  on  the  Linda  Vista  Terrace 
(at  an  altitude  of  130-140  m)  east  of  Murphy  Canyon,  city  of  San  Diego,  San  Diego  County,  California. 
The  faunule  is  suggestive  of  two  habitats,  an  exposed  open  coast  sandy  beach,  and  a  cobble  or  rocky-bot- 
tom, both  at  littoral  or  shallow  adlittoral  depths.  The  age  of  the  Lindavista  Formation  may  either  be  late 
Pliocene  or  early  Pleistocene  on  the  basis  of  the  fauna,  which  contains  the  extinct  species  Area  sisquo- 
censis  Reinhart  and  Pecten  helliis  (Conrad).  Because  of  the  fewer  tectonic-related  events  experienced  by 
the  Lindavista  Formation  than  by  the  unconformably  underlying  late  Pliocene  sediments,  the  formation 
may  actually  be  early  Pleistocene  in  age. 

The  presence  of  late  Pleistocene  marine  fossils  from  the  San  Diego  area  has  been 
well  documented  by  numerous  authors  (see  references  in  Kern,  1971;  also  Ellis,  in  Ellis 
and  Lee,  1919;  Berry,  1922;  Valentine  and  Meade,  1961;  Moore,  1968;  Kern,  Stump,  and 
Dowlen,  1971;  Bishop  and  Bishop,  1972).  Fossils  from  the  older  Pleistocene(?)  Lindavista 
Formation  (called  the  Sweitzer  Formation  by  some  authors)  are  unknown  from  the  San 
Diego  area,  although  Minch  (1967:  1 170)  has  reported  finding  "poorly  preserved  casts"  at 
one  locality  in  the  Lindavista  Formation  in  the  Tijuana-Rosarito  Beach  area  of  north- 
westernmost  Baja  California,  Mexico.  In  August,  1971,  Richard  C.  Schwenkmeyer  of  San 
Diego  Mesa  College  located  an  exposure  of  fossiliferous  beach  sand  containing  numerous 
fragments  and  a  few  complete  single  valves  of  the  Pismo  clam  Tivela  stultorum  (Mawe)  in 
a  new  housing  development  east  of  Murphy  Canyon  in  San  Diego  (Fig.  1).  Mr.  Schwenk- 
meyer kindly  brought  this  discovery  to  my  attention,  and  the  results  of  the  ensuing  in- 
vestigation form  the  basis  for  this  note. 

THE  LINDAVISTA  FORMATION 

The  Lindavista  Formation,  named  for  exposures  near  the  Lindavista  railroad  siding 
(Hanna,  1926:  218),  consists  of  several  meters  of  iron-red,  moderately  indurated  dirty  sand 
and  pebble-cobble  conglomerate.  Along  the  eastward  extent  of  the  formation,  the  sandy 
facies  interfingers  with  terrestrial  gravels  which  are  probably  deltaic  in  origin.  In  addition, 
the  formation  is  commonly  characterized  by  pea-sized  hematitic  concretions  on  weath- 
ered surfaces  (Hanna,  1926:  pi.  23;  Emery,  1950).  The  lithology  at  the  fossil  localities  (see 
also  Register  of  Localities)  varies  from  a  very  modern-looking  clean  gray  laminated  beach 
sand  to  a  fossiliferous  conglomerate  rich  in  heavy  minerals  (Fig.  2). 

The  Lindavista  Formation  blankets  the  Linda  Vista  Terrace,  a  broad  and  essentially 
planar,  slightly  westward  sloping  wave-cut  surface  extending  from  the  present  coastline 
nearly  fifteen  kilometers  inland,  where  it  terminates  at  the  base  of  the  foothills.  Remnants 
of  this  formation  are  exposed  on  terraces  from  northernmost  Baja  California  (Minch, 
1967:  1157,  1170)  to  areas  near  Oceanside  in  San  Diego  County  (Emery,  1950:  214,  and 
pi.  29).  The  most  prominent  features  of  the  Linda  Vista  Terrace  are  the  three  ancient 
beach  ridges  which  approximately  parallel  the  present  coastline.  These  have  been  inter- 
preted as  stillstands  during  the  marine  regression  which  followed  cutting  of  the  terrace 
(Peterson.  1970:  122).  Marine  sediment  along  the  eastern  margin  of  this  wave-cut  surface 
was  deposited  earlier  than  that  toward  the  coast. 

The  history  of  Pliocene  and  Pleistocene  sedimentation  of  the  San  Diego  coastal  plain 
has  been  summarized  by  Hertlein  and  Grant  (1944)  and  by  Peterson  ( 1970).  Two  possible 
sea  level  stands  have  been  postulated  for  the  events  in  the  formation  of  the  Linda  Vista 


SAN  DIEGO  SOC.  NAT.  HIST,  TRANS.  17(10);  119-128.28  MARCH  1973 


120 


Figure  1.  Index  map  of  San  Diego  area  showing  general  position  of  fossil  localities  on  east  side  of  Murphy 
Canyon. 

Terrace  (Hertlein  and  Grant,  1944:  64-65).  One  is  that  after  deposition  of  the  uppermost 
San  Diego  beds,  the  region  was  elevated  but  remained  sufficiently  below  wave  base  for 
wave  erosion  or  sea  floor  scour  to  truncate  the  marine  Pliocene  and  Eocene  beds.  The 
Lindavista  Formation  therefore  represents  distribution  by  ocean  waves  and  near-shore 
currents  of  coarse  material  derived  from  the  local  clastic  formations,  or  by  stream  erosion 
on  older  rocks  in  the  mountainous  areas  to  the  east.  The  second  possibility  is  that  parts  of 
the  San  Diego  Formation  were  elevated  slightly  above  sea  level  at  the  close  of  the  dia- 
strophic  movements  which  "inaugurated  Sweitzer  time."  The  soft  nature  of  the  San  Diego 
beds  resulted  in  their  quick  destruction  by  waves  and  subsequent  reduction  to  a  shallow 
submarine  platform.  However,  formation  of  the  Linda  Vista  Terrace  may  also  have  been 
the  result  of  a  relative  subsidence  of  the  coastal  plain  (or  rise  in  sea  level)  with  con- 


121 


comitant  transgression  of  a  shallow  sea.  then  followed  by  submarine  erosion.  Subsequent 
retreat  of  the  sea  (as  evidenced  by  the  beach  ridges)  and  deposition  of  the  offlap  facies 
(deltaic  and  terrestrial  clastic  sediments)  to  the  east  culminated  deposition  of  the  terrace 
material. 

Despite  problems  of  reconstructing  these  earlier  events,  at  least  150  meters  of  relative 
sea-level  change  and  only  minor  deformation  has  occurred  during  and  since  the  creation 
of  the  Linda  Vista  platform  (Peterson,  1970:  122).  West  of  the  Rose  Canyon  Fault  consid- 
erable tilting  and  uplift  has  occurred,  although  not  to  the  extent  as  affects  the  late  Plio- 
cene San  Diego  Formation  underlying  it  (Moore,  1972:  1 16,  fig.  3  [Structure  contours  on 
the  base  of  the  Lindavista  Formation]).  The  Lindavista  Formation  to  the  south  on  San 
Diego  Mesa  is  flat-lying  and  in  slight  angular  unconformity  with  the  underlying  San 
Diego  Formation  which  dips  6°  to  8°  to  the  south-southwest  (Hertlein  and  Grant,  1944: 
63  [as  the  Sweitzer  Formation]). 

AGE  OF  THE  LINDAVISTA  FORMATION 

The  age  of  the  Lindavista  Formation  has  been  variously  interpreted  as  late  Pliocene 
to  late  Pleistocene.  Originally  Hanna  (1926:  218)  simply  assigned  his  "Lindavista  terrace 
material"  to  the  Quaternary.  Hertlein  and  Grant  (1939:  71)  considered  their  Sweitzer 
Formation  (which  equals  the  Lindavista  Formation)  to  be  younger  than  the  Pliocene  San 
Diego  Formation  and  to  be  either  late  Pliocene  or  early  Pleistocene  in  age.  Milow  and 
Ennis  (1961:  28)  called  the  "Lindavista  Formation"  upper  Pleistocene,  but  they  were  re- 
ferring instead  to  deposits  of  topographically  lower  and  younger  terraces  than  the  Linda 
Vista  Terrace.  Their  combined  Sweitzer  Formation  and  an  overlying  unnamed  Sandstone 
comprise  the  Lindavista  Formation  of  current  usage.  Most  recently  Peterson  (1970:  122) 
has  assigned  the  Lindavista  Formation  to  the  middle  Pleistocene  because  of  its  medial  po- 
sition between  the  "Early  Pleistocene?"  higher  greatly  dissected  Poway  Terrace  and  the 
late  Pleistocene  lower  terrace  associated  with  the  Bay  Point  Formation.  Fossils  collected 
from  the  Linda  Vista  Terrace  (see  below;  also  Fig.  3)  indicate  either  a  late  Pliocene  or 
early  Pleistocene  age  for  the  formation.  Because  of  the  greater  number  of  tectonic-related 
events  experienced  by  the  late  Pliocene  San  Diego  Formation  (see  above),  the  Lindavista 
Formation  may  actually  be  early  Pleistocene  in  age.  although  further  evidence  is  needed 
before  any  age  determination  can  be  substantiated. 

FAUNA  OF  THE  LINDAVISTA  FORMATION 

The  fauna  of  the  Lindavista  Formation  is  essentially  a  modern  one,  with  a  few  ex- 
ceptions. Two  of  these.  Area  sisquocensis  Reinhart  and  Pecten  belhis  (Conrad),  are  known 
only  from  Pliocene  and  lower  Pleistocene  strata  in  California.  Turritella  gonostoma  hemp- 
hilli  Merriam,  only  questionably  found  in  the  Lindavista  Formation,  also  occurs  in  upper 
Pliocene  rocks  in  California.  Tegula  hemphilli  Oldroyd  occurs  in  both  the  upper  Pliocene 
San  Diego  Formation,  and  the  upper  Pleistocene  of  Pacific  Beach,  San  Diego.  The  re- 
maining molluscan  species  are  all  extant,  but  range  back  into  the  Pliocene.  The  barnacle 
Balanus  pacipcus  Pilsbry.  also  only  doubtfully  identified,  is  not  positively  known  to  occur 
in  Pliocene  or  older  rocks  (ZuUo.  1969:  10).  These  fossils  indicate  either  a  late  Pliocene  or 
early  Pleistocene  age  for  the  fauna. 

The  possibility  that  the  fossils  have  been  reworked  from  the  Pliocene  San  Diego  For- 
mation is  slight,  but  cannot  be  discounted  entirely.  The  Lindavista  Formation  in  the  vi- 
cinity of  the  fossil  exposures  unconformably  overlies  the  Eocene  Friars  Formation  and 
Stadium  Conglomerate  (Kennedy  and  Moore,  1971).  Field  investigations  have  revealed 
no  outcrops  of  the  San  Diego  Formation  anywhere  in  the  area  (G.  W.  Moore,  pers.  com- 
mun.;  Hertlein  and  Grant,  1944:  50).  The  closest  exposures  of  Pliocene  strata  are  all  sev- 
eral kilometers  distant,  to  the  south  on  the  south  side  of  Mission  Valley,  and  to  the  west  in 
the  vicinity  of  Mission  Bay  and  on  Mt.  Soledad. 

The  following  species  were  found  in  exposures  of  the  Lindavista  Formation  on  the 
east  side  of  Murphy  Canyon  in  San  Diego.  Nearly  all  the  species  are  from  one  locality 
(SDSNH  loc.  0325);  numbers  following  the  species  name  are  the  number  of  specimens 
(fragments  in  parentheses)  collected  from  this  locality,  unless  otherwise  noted.  For  local- 


122 


Figure  2.     SDSNH  locality  0329:  Fossiliferous  exposure  on  Santo  Road,  San  Diego,  showing  heavy-mineral 
sand,  cobble  conglomerate,  and  fragmented  valves  of  Tivela  siullorum.  Meterstick  for  scale. 

ity  data  see  Register  of  Localities. 

Mollusca.  Gastropoda:  Diodora  arnoldi  McLean,  1966—3;  Calliosloma  spp.—{\\);  Te^uhi  hemphilli  Old- 
royd,  1921—3(4);  Tegula  funebralis  (Adams,  1855)— 2(6);  Turritella  sp.  cf.  T.  gonostoma  hemphilli  Mer- 
riam,  1941— (6);  Turriiella  sp.—  l(\)\  Crepidula  spp.— (2);  Crucibulutn  spinosum?  (Sov^erhy.  1824)— (1); 
Polinices  recluzianus  (Deshayes,  1839)— 1(?12);  Acanthina  spirata  (Blainville,  1832)— (21);  unidentified 
fragments— (3).  Mollusca,  Bivalvia:  Yoldia  cooperi  Gabb,  1865— (1);  Area  sisquocensis  Reinhart,  1937— 
(1);  Oslrea  sp.— (1);  Pecten  bellus  (Conrad,  1857)— 1;  "Pecten'  spp.— (16);  Anomia'}  sp.— 1;  Pododesmus 
sp.-  (2);  Cardita  sp.  aff.  C  affinis  Sowerby,  1833-1  at  loc.  0329;  Lucinisca  nuttalli  (Conrad,  1837)-2(15); 
Tivela  siullorum  (Mawe,  1823)- 12,  1(15)  at  loc.  0321.  (6)  at  loc.  0322,  fragments  not  collected  at  Iocs.  0323 
and  0324,  (1)  at  loc.  0326,  3(10)  at  loc.  0329;  Proloihacal  sp.-(l)  at  loc.  0329;  Pelricola  carditoides  (Con- 
rad. 1837)-(1):  Spisula  hemphilli  (Dall,  1894)-(2);  Tellind!  sp.-(2);  Maeoma  nasuia'^.  (Conrad,  1837)- 
(1);  Zirfaea  pilshryi  Lowe,  1931-(6,  ?2);  Peniiella  sp.-(?l),  (4)  at  loc.  0326;  unidentified  fragments-(9). 
Annelida,  Polychaeta:  spionid  worm  burrows— 15  [in  single  Tegula  funebralis].  Echinodermata,  Ech- 
inoidea:  echinoid  spines— 8.  Arthropoda,  Crustacea  (Cirripedia):  Balanus  sp.  cf  B.  pacificus  Pilsbry, 
1916—1;  Megabalanus  sp.— 2;  unidentified  barnacle  wall  plates— 75  + . 


PALEOECOLOGY 

The  fauna  collected  does  not  represent  the  remains  of  any  single  biotic  community, 
but  rather  is  a  detrital  death  assemblage  from  several  near  shore  marine  habitats.  Speci- 
mens have  been  derived  mainly  from  two  habitats:  sandy  beach  and  cobble  or  rocky-bot- 
tom. 

An  exposed  open  coast  sandy  beach  habitat  at  littoral  or  adlittoral  depths  is  strongly 
suggested  by  the  great  abundance  of  the  Pismo  clam  Tivela  stultorum,  as  well  as  by  the 
presence  of  Spisula  hemphilli.  Donax  gouldi,  a  common  member  of  this  habitat  group  was 
unexpectedly  absent.  Many  specimens  are  quite  fragmented  (most  post  depositionally), 
but  their  as.sociation  with  the  cobble  conglomerate  indicates  either  local  transport  before 
deposition,  or  mixing  with  an  offlap  regressive  facies. 

A  cobble  or  rocky-bottom  habitat  is  suggested  by  many  of  the  species  in  the  fauna, 
including  those  in  the  genera  Diodora,  Calliosloma,  Tegula,  Acanthina,  Area,  Cardita, 
Protothaca,  Pelricola,  Peniiella,  Balanus,  and  Megabalanus.  Most  of  the  specimens  are 
fragmentary  and  while  the  conglomeratic  nature  of  the  outcrop  may  have  been  similar  to 
the  paleosubstrate  (see  above),  mixing  and  local  transport  are  indicated  here. 


123 


In  addition  to  the  above  habitats,  there  are  representatives  of  soft-bottom  (sand  and 
mud)  habitats  which  could  have  occurred  in  shallow  protected  bays  or  offshore  below  the 
level  of  effective  wave  action.  These  include  species  of  Turhtella,  Polinices,  Yoldia,  Pecten, 
Lucinisca,  Tellina,  Macoma,  and  Zirfaea. 

Most  species  are  largely  represented  by  only  a  relatively  few  fragments,  and  indicate 
at  least  local  transport  and  mixing.  The  high  degree  of  breakage  may  well  be  a  con- 
sequence of  the  conglomeratic  substrate  and  proximity  to  surf  action. 

Of  the  extant  species  represented  in  the  fauna,  only  Cardita  sp.  aff.  C.  affinis  does  not 
occur  today  in  the  vicinity  of  San  Diego.  Cardita  affinis  occurs  from  Bahi'a  de  Pequefia 
(26°  12'  N)  on  the  outer  coast  of  southern  Baja  California,  throughout  the  Gulf  of  Cali- 
fornia, and  south  to  northern  Peru.  Tiirritella  sp.  cf.  T.  gonostoma  hemphilli,  a  relative  of 
the  living  T.  gonostoma  s.s.,  which  occurs  today  from  the  Gulf  of  California  southward  to 
Ecuador,  also  suggests  warmer  water.  The  remainder  of  the  fauna  suggests  a  water  tem- 
perature comparable  with  that  of  the  present  Californian  Province.  There  are  no  cold-wa- 
ter or  strictly  northern  species  in  the  fauna.  The  occurrence  in  Pleistocene  sediments  of 
both  cooler  water  (Californian)  and  warmer  water  (Panamic)  species  cannot  yet  be  satis- 
factorily explained. 

Reconnaissance  geology  of  the  area  of  outcrop  by  George  W.  Moore  (pers.  comm.) 
indicates  that  the  Lindavista  Formation  lies  on  the  Eocene  Friars  Formation  (soft  sand- 
stone) directly  north  of  the  eroded  edge  of  the  overlying  Eocene  Stadium  Conglomerate, 
the  contact  between  the  two  Eocene  formations  trending  northeasterly.  A  ridge  of  the 
more  resistant  southward  dipping  conglomerate  seems  to  have  stood  slightly  in  relief  dur- 
ing erosion  of  the  wave-cut  platform.  The  fossiliferous  deposits  of  the  Lindavista  Forma- 
tion lie  in  an  embayment  etched  into  the  poorly  cemented  Friars  Formation  directly 
north  of  a  Pleistocene  rocky  headland  of  Stadium  Conglomerate.  This  physiographic  con- 
figuration may  have  been  responsible  for  the  accumulation  of  different  habitat  forms  in 
the  fauna.  Subsequent  deposition  of  an  offlap  facies  (conglomerate  and  tight  deltaic 
sandy  claystone,  never  deposited  to  the  west),  protected  the  deposits  from  weathering  and 
erosion  which  probably  accounts  for  the  general  lack  of  fossils  in  the  formation  as  a 
whole. 

SYSTEMATIC  NOTES 

Mollusca:  Gastropoda 

Diodora  arnoldi  McLc'dn,  1966 

Fig.  3c 

Range— Crescent  City,  Del  Norte  County,  California,  to  Isla  San  Martin,  Baja  Cali- 
fornia ( McLean,  1966:6). 

Remarks.— This  small  keyhole  limpet  occurs  exclusively  in  the  sublittoral  zone  and  is 
not  uncommon  on  the  undersides  of  rocks  below  a  depth  of  9  m  (McLean,  1969:  13). 
Diodora  arnoldi  is  also  known  from  the  upper  Pliocene  San  Diego  Formation  from  south- 
westernmost  San  Diego  County  (LACMIP  loc.  305A). 

Tegula  hemphilli  Oldroyd,  1921 

Fig.  3  a,b 

Remarks.— This  extinct  low  spired  Tegula  was  described  from  upper  Pleistocene  de- 
posits on  the  La  JoUa  Terrace  at  Pacific  Beach,  San  Diego  (Oldroyd,  1921:  115).  It  is  also 
known  to  occur  in  the  upper  Pliocene  San  Diego  Formation,  where  it  is  exposed  on  Tele- 
graph Canyon  Road,  east  of  the  city  of  Chula  Vista. 

Tegula funebralis  (Adams,  1855) 

Fig.  3d 

/?a/jge.— Vancouver  Island,  British  Columbia,  to  central  Baja  California  (McLean, 
1969:  22). 

Remarks.— This  species  occurs  strictly  intertidally  and  is  abundant  in  rocky  areas  at 


124 


■•ifA 


'^ 


%• 


^oiP' 


Figure  3.  Fossils  from  the  Lindavista  Formation,  a,  b,  Tegula  heniphilli.  SDSNH  15581,  width  16.2  mm;  c, 
Diodora  amoldi.  SDSNH  14886.  length  9.1  mm;  d.  Tei^iiUi  fiwehralls,  SDSNH  14889,  width  27.2  mm;  e,  Pecten 
hellus,  SDSNH  13117,  height  22  mm;  f",  Cardila  sp.  aff.  C.  affinis,  SDSNH  16678,  length  59  mm;  g,  Tivela  stulto- 
rum,  SDSNH  14887.  length  78.7  mm;  h,  Megabalanus  sp.,  SDSNH  14888,  height  60  mm. 

the  midtide  level  (McLean,  1969:  22).  Dead  shells  are  retained  in  the  intertidal  zone  by 
hermit  crabs,  which  use  them  for  their  own  protection.  The  columella  of  one  specimen  has 
been  extensively  bored  by  spionid  worms. 

MoUusca:  Bivalvia 

Area  sisquocensis  Reinhart,  1937 

Remarks.— One  fragment  of  this  distinctive  Pliocene  and  lower  Pleistocene  Area  was 
found.  This  species,  described  from  the  Pliocene  Careaga  Formation,  also  occurs  in  the 
lower  Pleistocene  Santa  Barbara  Formation  (Reinhart,  1943:  25),  as  well  as  in  the  upper 


125 


Pliocene  San  Diego  Formation  exposed  in  southwesternmost  San  Diego  County  (LAC- 
MIP  Iocs.  305  and  305A). 

Pecten  bellus  (Conrsid,  1857) 

Fig.  3e 

Remarks.— One  small  flat  left  valve  referable  to  Pecten  hellus  has  the  apical  angle, 
number  of  ribs,  and  muscle  scar  typical  of  the  species.  This  characteristic  middle  to  late 
Pliocene  species  also  occurs  in  the  lower  Pleistocene  Santa  Barbara  Formation  (J.  W.  Val- 
entine, pers.  commun.).  Sixteen  additional  pectinid  fragments  remain  unidentified. 

Cardita  sp.  aff'.  C.  affinis  Sowerby,  1833 
Fig.  3f 

Range— [oi  C.  affinis]  Bahi'a  de  Pequena,  and  the  Gulf  of  California  south  to  north- 
ern Peru  (Keen,  1958:  85;  Olsson,  1961:  190). 

Remarks.— One  complete  right  valve  differs  from  Recent  specimens  examined  by  its 
more  central  umbo,  rounded  anterior  and  posterior  margins,  and  greater  thickness,  al- 
though these  diff'erences  may  only  be  phenotypic.  Cardita  affinis  occurs  under  rocks  or  in 
crevices  intertidally  and  off'shore  to  a  depth  of  27  meters  (Keen,  1971:  107).  This  is  the 
only  living  species  in  the  fauna  of  the  Lindavista  Formation  that  does  not  presently  occur 
along  the  San  Diego  coastline. 

Fossil  occurrences  of  C.  affinis  are  known  only  from  the  upper  Pliocene  and  Pleisto- 
cene of  the  southern  part  of  the  Gulf  of  CaUfornia  (Durham,  1950:  72;  Hertlein,  1957:  62; 
Emerson  and  Hertlein,  1964:  341). 

Tivela  siultorum(Ma'^e,  1823) 

Fig-  3g 

Range.— Hsilf moon  Bay,  San  Mateo  County,  California,  to  Bahia  Magdalena,  Baja 
California  (Fitch,  1953:60). 

Remarks.— This  is  the  most  abundant  of  any  species  found,  and  fragments  and  occa- 
sional complete  valves  were  present  at  every  locality  in  the  area  which  produced  fossils. 
No  paired  valves  were  found,  and  all  appear  to  be  detrital.  A  few  specimens  still  exhibit 
faint  coloration  patterns.  Many  of  the  valves  have  been  post-depositionally  fragmented. 
Tivela  stultorum  usually  occurs  in  the  intertidal  zone  on  flat  sandy  beaches  on  the  open 
coast  (exposed  to  the  full  force  of  the  surO.  or  in  channels  leading  into  bays  and  estuaries 
(Fitch,  1953:  60). 

Penitella  sp.  indet. 

Remarks. —Se\er2i\  small  specimens  of  a  poorly  preserved  Penitella  were  removed 
from  sandstone  cobbles  at  two  localities.  The  umbonal  regions  of  all  the  specimens  are 
too  damaged  for  specific  identification.  Species  of  Penitella  are  commonly  found  in  cob- 
bles in  molluscan  death  assemblages,  and  usually  represent  intertidal  or  high  inner  sub- 
littoral  zones  on  the  open  coast  where  wave  action  is  strong  and  fine  sedimentation  does 
not  occlude  the  siphonal  openings. 

Arthropoda:  Crustacea 
Cirripedia 

Remarks.— 0\er  75  fragments  of  barnacle  wall  plates  were  found  at  one  locality. 
These,  although  mostly  unidentified,  represent  several  species  belonging  to  both  Balanus 
s.s.  and  Megahalanus  (Fig.  3h).  Numerous  fragments  were  recovered  from  the  con- 
glomeratic sandstone,  but  none  were  found  attached  to  cobbles,  nor  were  any  bases  found 
on  any  of  the  cobbles.  A  single  opercular  plate  has  been  tentatively  identified  as  Balanus 
sp.  cf  B.  pacificus  Pilsbry,  a  species  not  positively  known  from  Pliocene  or  older  rocks 
(Zullo,  1969:  10).  Balanus  pacificus  occurs  today  from  San  Francisco,  California,  to  north- 


126 


em  Peru,  and  is  common  in  Pleistocene  deposits  of  California  and  northern  Baja  Califor- 
nia (Zullo,  1969:  10). 


REGISTER  OF  LOCALITIES 


► 


All  of  the  following  localities  are  from  the  lower  Pleistocene  (?)  Lindavista  Formation  from  exposures  on 
the  Linda  Vista  Terrace  east  of  Murphy  Canyon  in  the  city  of  San  Diego,  San  Diego  County.  California.  Most  of 
the  specimens  were  collected  by  me  in  September  (Iocs.  0321-0326)  and  late  November  (loc.  0329),  1971.  These 
localities  are  now  mostly  on  private  residential  property.  Specimens  have  been  deposited  in  the  Department  of 
Invertebrate  Paleontology  in  the  San  Diego  Natural  History  Museum  and  bear  its  locality  numbers. 

Loc.  0321.  Northwest  trending  bank  facing  southwest  on  east  (back)  side  of  residence  at  5349  Jazmin 
Court.  San  Diego.  Unconsolidated  sand.  Altitude  142  m.  Approximate  coordinates:  32°  49.7'  N.,  117°  5.6'  W. 
Locality  found  by  R.  C.  Schwenkmeyer. 

Loc.  0322.  West-facing  bank  at  the  northeast  corner  of  the  intersection  of  Sandia  Place  and  Gabacho 
Drive  (10810  Gabacho  Drive),  San  Diego.  Fossil  fragments  in  gray  laminated  and  cross-bedded  non-indurated 
beach  sand  with  occasional  scattered  pebbles.  Approximate  coordinates:  32°  49.8'  N.,  1 17°  5.8'  W. 

Loc.  0323.  North-facing  bank  on  south  side  of  lot  at  10805  Gabacho  Drive  (in  cul-de-sac  opposite  Sandia 
Place).  San  Diego.  Laminated  gray  unconsolidated  beach  sand  overlain  bv  well-indurated  fossiliferous  con- 
glomeratic sandstone,  which  in  turn  is  overlain  by  a  terrestrial  conglomeratic  facies.  Approximate  coordinates: 
32°  49.8' N.,  117°  5.8' W. 

Loc.  0324.  East-west  trending  utilities  ditch  in  south  side  of  street  along  north  side  of  10825  Gabacho 
Drive,  San  Diego.  Approximate  coordinates:  32°  49.8'  N.,  117°  5.8'  W. 

Loc.  0325.  North-facing  bank  on  south  side  of  residence  at  10735  Montego  Drive  (in  southeast  corner  of 
first  cul-de-sac  on  Montego  Drive  west  of  El  Noche  Way),  San  Diego.  Unconsolidated  fossiliferous  con- 
glomeratic sandstone.  Altitude  131  m.  Approximate  coordinates:  32°  49.7'  N.,  117°  5.8'  W.  Locality  found  by  G. 
W.  Moore. 

Loc.  0326.  West-facing  bank  on  east  side  of  residence  at  10735  Montego  Drive  (in  southeast  corner  of 
first  cul-de-sac  on  Montego  Drive  west  of  El  Noche  Way),  San  Diego.  Approximate  coordinates:  32°  49.7'  N., 
1I7°5.8' W. 

Loc.  0329.  Ninety-meter  stretch  along  west-facing  roadcut  on  east  side  of  Santo  Road,  beginning  approx- 
imately 35-40  meters  north  of  intersection  of  Santo  Road  and  Monte  Negro  Drive,  San  Diego.  Fossiliferous  len- 
ses mixed  with  black  heavy-mineral  sand  and  cobble  conglomerate.  Locality  found  by  G.  W.  Moore. 

ACKNOWLEDGEMENTS 

I  am  grateful  to  Richard  C.  Schwenkmeyer  (San  Diego  Mesa  College)  for  bringing  his  specimens  and  local- 
ity to  my  attention.  George  W.  Moore  (USGS)  extended  many  courtesies,  including  extensive  data  from  his  own 
field  investigations,  and  constructive  criticism  of  the  manuscript.  The  molluscan  identifications  have  been 
checked  by  J.  G.  Vedder  (USGS).  Arnold  Ross  (SDSNH)  kindly  identified  the  cirripeds,  and  read  the  manu- 
script. Figure  1  was  prepared  by  Lanci  Valentine  (University  of  California  at  Davis). 

LITERATURE  CITED 

Berry,  S.  S. 

1 922.  Fossil  chitons  of  western  North  America.  Proc.  Calif  Acad.  Sci.,  ser.  4,  1 1 :  399-525. 

Bishop,  M.  J.,  and  S.  J.  Bishop 

1972.  New  records  of  Pleistocene  marine  Mollusca  from  Pacific  Beach,  San  Diego,  California.  Veliger. 
15:6. 

Durham,  J.  W. 

1950.  Megascopic  paleontology  and  marine  stratigraphy,  216  p.  In  1940  E.  W.  Scripps  cruise  to  the  Gulf  of 
California.  Geol.  Soc.  Amer.,  Mem.  43. 

Ellis,  A.  J.,  and  C.  H.  Lee 

1919.  Geology  and  ground  waters  of  the  western  part  of  San  Diego  County,  California.  U.  S.  Geol.  Surv., 
Water-Supply  Paper  446. 

Emerson,  W.  K.,  and  L.  G.  Hertlein 

1964.  Invertebrate  megafossils  of  the  Belvedere  Expedition  to  the  Gulf  of  California.  Trans.  San  Diego 
Soc.  Nat.  Hist.,  13:  333-368. 

Emery,  K.  O. 

1950.   Ironstone  concretions  and  beach  ridges  of  San  Diego  County,  California.  Calif  Jour.  Mines  Geol., 
46:213-221. 

Fitch.  J.  E. 

1953.  Common  marine  bivalves  of  California.  Calif  Dept.  Fish  Game.  Mar.  Fish.  Br.,  Fish  Bull.  90:  1-102. 

Hanna,  M.  A. 

1926.  Geology  of  the  La  Jolia  quadrangle.  California.  Univ.  Calit:  Publ.  Geol.  Sci..  16:  188-247. 

Hertlein,  L.  G. 

1957.   Pliocene  and  Pleistocene  fossils  from  the  southern  portion  of  the  Gulf  of  California.  Bull.  So.  Calif. 
Acad.  Sci..  56:  57-75. 


127 


Hertlein,  L.  G..  and  U.  S.  Grunt  IV 

1939.  Geology  and  oil  possibilities  of  southwestern  San  Diego  County.  Calif.  Jour.  Mines  Geol..  35:  57-78. 
1944.  The  geology  and  paleontology,'  of  the  marine  Pliocene  of  San  Diego.  California.  Part  1,  geology. 
Mem.  San  Diego  Soe.  Nat.  Hist.,  2:  1-72. 

Keen,  A.  M. 

1958.  Sea  shells  of  tropical  west  America.  Stanford  Univ.  Press,  Stanford.  624  p. 

1971.  Sea  shells  of  tropical  west  America  [second  edition],  Stanford  Univ.  Press,  Stanford.  1064  p. 

Kennedy,  M.  P.,  and  G.  W.  Moore 

1971.  Stratigraphic  relations  of  Upper  Cretaceous  and  Eocene  formations,  San  Diego  coastal  area,  Califor- 
nia. Amer.  Assoc.  Petrol.  Geol.  Bull.,  55:  709-722. 

Kern.  J.  P. 

1971.  Paleoenvironmental  analysis  of  a  late  Pleistocene  estuary  in  southern  California.  Jour.  Paleont  45- 
810-823. 

Kern.  J.  P..  T.  E.  Stump,  and  R.  J.  Dowlen 

1971.  An  upper  Pleistocene  marine  fauna  from  Mission  Bay,  San  Diego.  California.  Trans.  San  Diego  Soc. 
Nat.  Hist.,  16:  329-338. 

McLean,  J.  H. 

1966.  A  new  genus  of  Fissurellidae  and  a  new  name  for  a  misunderstood  species  of  west  American  Diod- 
ora.  Los  Angeles  Co.  Mus.  Contrib.  Sci.,  100:  1-8. 

1969.  Marine  shells  of  southern  California.  Los  Angeles  Co.  Mus.  Nat.  Hist,  Sci.  ser.  24,  Zool.  11:  1-104. 
Milow,  E.  D.,  and  D.  B.  Ennis 

1961.  Guide  to  geologic  field  trip  of  southwestern  San  Diego  County,  p.  23-43.  In  B.  E.  Thomas,  ed.. 
Guidebook  for  field  trips  [57th  Ann.  Meet.  Cordilleran  Sec,  Geol.  Soc.  Amer.]. 

Minch.  J.  A. 

1967.  Stratigraphy  and  structure  of  the  Tijuana-Rosarito  Beach  area,  northwestern  Baja  California,  Mex- 
ico. Geol.  Soc.  Amer.,  Bull.  78:  1155-1177. 

Moore.  E.  J. 

1968.  Fossil  moUusks  of  San  Diego  County.  San  Diego  Soc.  Nat.  Hist.,  Occas.  Paper  15. 

Moore.  G.  W. 

1972.  Ofi"shore  extension  of  the  Rose  Canyon  Fault.  San  Diego,  California,  U.  S.  Geol.  Surv.,  Prof  Paper 
800-C:  113-116. 

Oldroyd,  T.  S. 

1921.  New  Pleistocene  mollusks  from  California.  Nautilus,  34:  1 14-1 16. 

Olsson.  A.  A. 

1961.  Mollusks  of  the  tropica!  eastern  Pacific  particularly  from  the  southern  half  of  the  Panamic-Pacific 
faunal  province  (Panama  to  Peru).  Panamic-Pacific  Pelecypoda.  Paleont.  Res.  Inst..  Ithaca  (New 
York).  574  p. 

Peterson.  G.  L. 

1970.  Quaternary  deformation  of  the  San  Diego  area,  southwestern  California,  p.  120-126.  In  E.  C.  Allison, 
et  al,  eds..  Pacific  slope  geology  of  northern  Baja  California  and  adjacent  Aha  California.  [Geological 
guidebook  for  1970  fall  field  trip.  Pacific  sees.  AAPG.  SEPM.  and  SEG]. 

Reinhart.  P.  W. 

1943.  Mesozoic  and  Cenozoic  Arcidae  from  the  Pacific  slope  of  North  America.  Geol.  Soc.  Amer..  Spec. 
Paper  no.  47. 

Valentine.  J.  W..  and  R.  F.  Meade 

1961.  Californian  Pleistocene  paleotemperatures.  Univ.  Calif  Publ.  Geol.  Sci.,  40:  1-45. 
Zullo,  V.  A. 

1969.  Thoracic  Cirripedia  of  the  San  Diego  Formation.  San  Diego  County.  California.  Los  Angeles  Co. 
Mus.  Contrib.  Sci..  159:  1-25. 


Department  of  Geology,  University  of  California,  Davis,  California  95616 


S-/\)P^-S 


MUS.  COMP.  ZOOL 
LIBRARY 

HARVARD 
UNIVERSITY 


POST-BATHOLITHIC  GEOLOGY  OF  THE  JACUMBA  AREA, 
SOUTHEASTERN  SAN  DIEGO  COUNTY,  CALIFORNIA 

JOHN  A.  MINCH  AND  PATRICK  L.  ABBOTT 


TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  11  10  APRIL  1973 


POST-BATHOLITHIC  GEOLOGY  OF  THE  JACUMBA  AREA, 
SOUTHEASTERN  SAN  DIEGO  COUNTY,  CALIFORNIA 

JOHN  A.  MINCH  AND  PATRICK  L.  ABBOTT 


ABSTRACT.— The  post-batholithic  history  of  the  Peninsular  Range  is  documented  by  sparse  exposures  of 
fluvial  and  volcanic  rocks  in  widely  separated  areas.  The  Jacumba  Valley  outcrops  present  one  of  the  most 
complete  stratigraphic  records  in  the  range. 

The  intial  erosion  surface  upon  the  Southern  California  Batholith  probably  formed  in  the  Late  Cre- 
taceous. This  surface  was  fairly  deeply  weathered  as  was  the  earliest  granitic  gravel  deposited  upon  it. 
Clasts  resembling  those  of  the  Table  Mountain  Gravels  were  transported  across  the  Jacumba  area  to  the 
Pacific  Coastal  region,  where  they  appear  in  the  Cabrillo  Formation  of  Late  Cretaceous  age.  Eocene 
"Poway-type"  gravel  was  transported  acro.ss  an  essentially  parallel  surface  just  south  of  Jacumba  Valley 
near  La  Rumorosa.  Erosion  partially  removed  and  reworked  the  gravel  until  Early  Miocene  outpourings  of 
basalt  and  pyroclastic  debris  filled  in  much  of  the  Jacumba  Valley  area.  The  first  basalt  flows  from  eruptive 
centers  within  the  valley  were  followed  by  faulting  and  the  deposition  of  andesitic  pyroclastic  and  lahar  depos- 
its. The  source  of  this  andesite  may  have  been  the  plug-like  masses  at  Round  Mountain  and  Jade  Bench- 
mark. Continued  faulting  ofl'set  the  volcanic  rocks  before  the  eruption  of  a  second  series  of  basalt  flows  which 
covered  the  andesite  in  the  northeast  and  east  portions  of  the  valley.  Intermittent  erosion  within  the  volcanic 
sequence  is  evidenced  by  fluvial  and  eolian  volcaniclastic  deposits.  Post-volcanic  faulting  elevated  the  Penin- 
sular Range  and  accelerated  erosion  to  produce  the  present  topography. 

The  Jacumba  Valley  area  is  located  at  the  crest  of  the  Peninsular  Range  and  straddles 
the  Mexico-United  States  boundary.  Interstate  8  and  the  San  Diego  and  Arizona  Eastern 
Railroad  pass  through  the  valley  between  San  Diego  and  the  Imperial  Valley.  Rainfall  is 
hght  resulting  in  sparse  vegetation  and  excellent  outcrops.  Field  work  for  this  report  was 
accomplished  in  the  fall  of  1971  and  spring  of  1972  in  conjunction  with  a  field  geology  class 
at  California  State  University,  San  Diego. 

The  first  account  of  the  post-batholithic  geology  of  Jacumba  Valley  was  by  Fairbanks 
(1893),  who  indicated  the  presence  of  the  gravels  and  volcanics  at  the  crest  of  the  Peninsular 
Range.  Miller  (1935a)  followed  with  a  brief  description  of  the  rocks  and  included  specula- 
tions on  their  former  widespread  extent,  exotic  origin,  and  Miocene  age. 

In  the  late  1940s  and  early  1950s  field  classes  from  San  Diego  State  College  used  the 
area  for  reconnaissance  mapping  exercises.  These  were  compiled  as  map  sheet  23  (Brooks 
and  Roberts,  1954)  in  Bulletin  170  of  the  California  Division  of  Mines  and  Geology.  Ja- 
cumba Valley  remains  an  excellent  area  for  students  to  map  in  a  terrane  exposing  a  variety 
of  eruptive  features  resting  on  sedimentary,  plutonic,  and  metamorphic  rocks. 

Gastil  and  Bushee  ( 1 96 1 )  and  Weber  ( 1 963 )  briefly  mention  the  valley.  Hawkins  ( 1 970) 
provided  a  detailed  analysis  of  the  chemistry  of  the  volcanic  rocks  in  the  valley  and  tied 
them  into  the  over-all  picture  of  sea-floor  spreading  in  Southern  Cahfornia.  Minch  (1971) 
described  the  Table  Mountain  Formation  and  indicated  its  nonlocal  origin. 

BASEMENT  ROCKS 

The  crystalline  rocks  flooring  the  valley  have  been  mapped  as  two  separate  units  (We- 
ber, 1963).  The  older  mass  is  comprised  of  metamorphic  rocks  mixed  with  granodiorite, 
diorite,  and  pegmatites.  These  metasedimentary  rocks  are  dominated  by  the  quartz-  and 
mica-rich  Julian  Schist,  with  minor  amounts  of  gneiss  and  quartzite  and  occasional  pods  of 
marble.  These  are  cut  by  abundant  pegmatites  and  plutonic  bodies. 

The  younger  plutonics  of  the  Southern  California  Batholith  are  here  composed  mostly 
of  quartz  diorite  along  with  granodiorite  and  minor  pods  of  gabbro. 

TABLE  MOUNTAIN  GRAVELS 
The  Table  Mountain  Gravels  are  light  yellow-brown,  moderately  bedded,  fairly  well- 

SANDIEGOSOC.  NAT.  HIST.  TRANS.  1 7(  1 1):  129-136,  10  APRIL  1973 


130 


sorted,  very  friable,  medium  to  coarse-grained  sandstones  and  conglomeratic  sandstones 
which  crop  out  in  and  near  Jacumba  Valley.  In  addition  to  local  granitic  clasts  they  contain 
clasts  of  low-grade  green  metavolcanic  and  metasedimentary  rocks  and  quartzites  that  are 
not  found  locally. 

Miller  (1935a:  138)  defined  the  Table  Mountain  Formation  from  exposures  at  Table 
Mountain  7  km  northeast  of  Jacumba  as:  "Moderately  consolidated  deposits  of  yellowish  to 
reddish-brown  gravels  and  sands.  Various  kinds  of  pre-Cretaceous  crystalline  rock  frag- 
ments occur  in  the  formation  . . .  These  sediments  are  rather  variable  in  character,  crudely 
stratified,  and  gently  dipping." 

Several  authors  have  indicated  the  exotic  nature  of  these  gravels  found  high  in  the  Pen- 
insular Range  (Fairbanks,  1893;  Brooks  and  Roberts,  1954;  Weber,  1963).  Brooks  and  Rob- 
erts (1954)  compare  the  clasts  to  the  Santiago  Peak  Volcanics  of  western  San  Diego  County: 
"They  contain  fragments  of  dacites  and  other  aphanitic  rocks  that  show  strong  similarities 
to  the  Jurassic?Santiago  Peak  Volcanics  of  western  San  Diego  County.  These  gravels  are 
partially  interbedded  with  and  principally  overlain  by  other  volcanic  rocks." 

DISTRIBUTION. -The  Table  Mountain  Gravels  crop  out  on  the  erosion  surface  in  a  belt 
about  10  km  wide  and  25  km  long  that  lies  roughly  parallel  to  the  axis  of  the  Peninsular 
Range.  They  are  the  remnants  of  an  extensive  fluvial  deposit.  In  the  Jacumba  area  they 
stretch  another  5  km  down  the  frontal  scarp  of  the  range.  The  principal  outcrops  are  in 
and  around  Jacumba  Valley  and  in  the  area  just  west  of  La  Rumorosa  in  Baja  California. 
The  small  isolated  patches  of  the  gravels  which  occur  at  lower  elevations  on  the  frontal 
scarp  of  the  range  are  the  easternmost  exposures. 

The  best  exposures  of  the  Table  Mountain  Gravels  are  in  the  area  of  Jacumba  Valley 
where  the  Jacumba  Volcanics  form  a  resistant  cap  above  the  gravels  (Fig.  1).  A  typical 
section  measured  on  a  flat-topped  hill  just  north  of  Jacumba  in  the  northwest  corner  of 
Sec.  5,  T  18  S,  R  8  E  consists  of  75  m  of  interbedded  light  yellow-brown,  moderately  to 
thickly  bedded,  very  friable,  medium-  to  coarse-grained  sandstone  and  Vi  to  1  m  thick 
beds  of  conglomeratic  sandstone.  The  sandstones  within  the  Table  Mountain  Formation 
are  sheet-wash  to  fluvially  deposited,  plutonic  lithic  arkose.  The  framework  grains  are 
very  angular,  poorly  to  very  poorly  sorted,  mineralogically  immature,  and  are  cemented 
by  poikilotopic,  very  coarsely  crystalline  calcite  where  unleached.  Common  grains  include 
both  fresh  and  heavily  sericitized  plagioclase  and  orthoclase,  polycrystalline  quartz,  schist 
and  plutonic  rock  fragments  along  with  hornblende,  biotite,  muscovite,  and  other  acces- 
sory minerals. 

The  gravel-sized  clasts  are  subangular  to  subrounded  and  average  2.5  to  5  cm  in  di- 
ameter with  clasts  commonly  to  10  cm  and  rarely  to  30  cm.  Fifty  percent  of  the  clasts  are 
extraregional,  low-grade,  green  metavolcanic  and  metasedimentary  rocks.  Other  signifi- 
cant components  of  these  gravels  are  quartzites  and  resistant  sandstones  (25%),  and  local 
granitic  and  gneissic  basement  rocks  (12%).  The  thickness  of  the  gravels  is  quite  variable, 
ranging  from  a  thin  mantle  on  the  surface  to  greater  than  75  m.  Most  exposures  are  less 
than  30  m  in  thickness. 

Other  good  exposures  of  these  gravels  occur  on  the  northeast  side  of  Jacumba  Peak 
(N.  Center  Sec.  7,  T  18  S,  R  8  E),  on  Table  Mountain  (T  17  S,  R  8  E),  and  in  Myer  Valley 
(NE  '/4,  Sec.  26,  T  17  S,  R  19  E)  about  halfway  down  the  frontal  scarp. 

In  the  Jacumba  area  the  Table  Mountain  Gravels  seem  to  have  been  highly  eroded 
before  the  deposition  of  the  Jacumba  Volcanics.  This  is  evident  because  the  basalts  and 
breccias  have  an  irregular  basal  contact  which,  in  one  case,  rests  on  75  m  of  gravel  at  one 
end  of  a  hill  but  sits  on  the  granitic  erosion  surface  120  m  lower  in  altitude  at  the  other 
end  of  the  hill. 

AGE.  The  Table  Mountain  Gravels  are  certainly  older  than  the  18.5  m.y.  Jacumba 
Volcanics  which  overlie  them,  and  they  are  younger  than  the  rocks  of  the  Peninsular 
Range  Batholith  upon  which  they  rest  (90-105  m.y.,  Bushee  et  al.,  1963;  cooling  age  of  65- 
80  m.y.,  R.  G.  Gastil,  pers.  comm.).  They  were  deeply  eroded  before  the  deposition  of  the 
Jacumba  Volcanics, suggesting  that  they  may  be  significantly  older  than  the  Miocene  vol- 
canics. 

In  the  San  Diego  coastal  area  the  conglomerate  of  the  Upper  Cretaceous  Cabrillo 


131 


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Hypersthene  andes i te  plugs;  contain  xeno- 
liths  of  quartz  diorite;  exhibit 
quaquaversal  dips 
Upper  basalt  flows;  olivine  bearing;  platy 

to  massive  to  scoriaceous 
Basa It  d  i  kes 
Cinder  agglomerate;  baked  reddish  cinders 

with  abundant  bombs 
Andesite  breccia  and  conglomerate,  lahar 
deposits  and  vol  can i cl ast i c  sandstone; 
i  ntergradat  iona I 
Tas  0-60m   Vol  can i clas t i c  sandstone;  fluvial 

and  eol i  an 
Taf  0-6m    Andesite  flows 
Tat  0-12m   Tuff 
0-40m    Lower  basalt  flows;  olivine  bearing;  platy 

to  massive  to  scoriaceous 
0-90m    Cinder  cone  and  associated  cinder  carpet 
Tcy  0-15m   Yellow  cinder  bed 


Ktm  0-90m    Sandstones  and  conglomeratic  sandstones; 

fluvial  with  minor  basal  arkose;  plutonic 
lithic  arkose;  majority  of  gravel  clasts 
are  extra  regional 


Kqd 
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Quartz  diorite  and  granodiorite  of  Southern 

California  batholith 
Julian  schist  with  gneiss,  quartzite  and 

marble  mixed  with  quartz  diorite, 

granodiorite  and  pegmatite 


Figure  1.  GENERALIZED  COLUMNAR  SECTION 

Formation  (Kennedy  and  Moore,  1971)  contains  a  small  percentage  of  black  to  blue-gray 
quartzite,  light  colored  quartzite,  and  chert  pebbles,  and  several  types  of  metavolcanic 
clasts  not  associated  with  the  local  basement.  These  clasts  are  not  found  in  the  locally  de- 
rived gravel  of  the  underlying  Lusardi  Formation,  indicating  that  they  are  extra-region- 
ally  derived  clasts.  These  exotic  clast  types  are  also  found  in  the  Table  Mountain  Forma- 
tion. 

The  presence  of  the  extra-regional  clasts  in  the  Cabrillo  Formation,  which  has  been 
dated  as  Maestrichtian  (Kennedy  and  Moore,  1971),  suggests  a  relation  to  the  Table 
Mountain  Gravels,  which  in  turn  suggests  that  the  two  formations  might  be  the  same  age. 
If  so,  this  would  place  the  initial  deposition  of  the  gravels  in  the  Late  Cretaceous. 


I 


JACUMBA  VOLCANICS 

The  Jacumba  Volcanics  were  defined  by  Miller  (1935a:  138-139)  for  "...the  exten- 
sive rocks  which  are  excellently  exposed  in  the  several  areas  north  to  east  of  Jacumba  .  .  . 
Some  volcanic  breccias  or  pyroclastics  here  occur  toward  the  bottom  of  the  lava  beds."  In 
the  present  report  the  Jacumba  Volcanics  are  subdivided  into  three  basic  units.  They  are: 
(1)  basalt  flows  and  cinder-cone  deposits,  (2)  basaltic-andesite  plugs,  and  (3)  andesite 
breccia,  lahar  deposits,  conglomerate,  and  volcaniclastic  sandstone. 

The  same  authors  who  discussed  the  Table  Mountain  Gravels  also  generally  dis- 
cussed the  Jacumba  Volcanics  (Fairbanks,  1893;  Miller,  1935a,  b;  Brooks  and  Roberts, 


132 


1954;  Gastil  and  Bushee,  1961;  and  Weber,  1963).  In  addition,  Hawkins  (1970)  discussed 
the  petrochemistry  of  the  volcanics  and  cited  a  K/Ar  whole  rock  date  of  18.7  ±1.3  m.y. 
for  the  lower  part  of  the  basalt  sequence. 

The  Jacumba  Volcanics  form  a  20  by  55  km  belt  of  outcrops  parallel  to  the  axis  of  the 
Peninsular  Range.  However,  the  principal  areas  of  outcrop  are  in  a  narrow  belt  5  to  10 
km  wide  by  55  km  long  on  and  along  the  frontal  scarp  of  the  range,  with  the  majority  of 
the  outcrops  near  the  base  of  the  scarp.  The  Jacumba  Volcanics  also  crop  out  in  and 
around  Jacumba  Valley. 

The  basalt  flows  are  the  most  extensive  part  of  this  unit,  with  outcrops  over  most  of 
the  valley.  The  andesite  breccia,  lahar  deposits,  and  volcaniclastic  sandstone  are  as  exten- 
sive but  more  limited  in  outcrop.  Remnants  of  at  least  five  cinder  cones  and  two  promi- 
nent hypersthene  andesite  plugs  are  exposed  in  the  valley. 

FLOWS.-The  lavas  consist  of  up  to  40  meters  of  gray  to  dark  gray,  platy  to  massive, 
vesicular  olivine  basalt  and  basaltic  andesite.  The  basalt  flows  crop  out  at  the  base  and  at 
the  top  of  the  volcanic  sequence.  As  many  as  five  or  six  flow  units  may  be  represented  in 
any  given  outcrop.  These  flow  units  tend  to  be  massive  in  their  upper  portions  but  exhibit 
platy  jointing  due  to  flow  shearing  at  their  bases  or  where  they  abut  other  rocks  (see 
Hawkins,  1970,  for  detailed  chemical  description). 

DIKES  AND  CINDER  CONES.-Squaw  Tit  is  an  excellent  example  of  one  of  several 
prominent  dikes  exposed  on  Table  Mountain  (Map-Fig.2).  This  dike  system  appears  to 
have  partially  followed  a  northwest-trending  fracture  system.  The  basalt  in  these  dikes 
ranges  from  a  dense  gray-green  to  dark  gray,  well-jointed  olivine  basalt  to  a  scoriaceous 
ohvine  basalt  which  appears  to  have  been  close  to  the  surface  of  a  vent  represented  by  the 
spine  of  Squaw  Tit.  These  dikes  cut  the  lower  basalt  unit  and  may  be  the  source  for  the 
upper  basalt  unit. 

Associated  with  these  dikes  are  at  least  two  cinder  cones.  The  oldest  cinder  cone  is  at 
the  base  of  the  section  east  of  a  fault  on  the  southern  side  of  Table  Mountain.  It  is  75  to 
90  m  high  and  is  well  exposed  as  a  result  of  quarrying  operations.  The  cinders  comprising 
the  cone  are  red-  to  purple-brown,  well  sorted,  thinly  bedded,  and  tend  to  be  lapilh  to 
dust  size  with  a  small  percentage  of  blocks  and  very  few  bombs.  The  upper  15  m  of  the 
cone  has  been  altered  by  gasses  to  a  yellow-brown  color.  Also  deposited  during  this  pyro- 
clastic  episode  is  a  thinly  bedded  cinder  carpet  up  to  4.5  m  thick,  which  appears  to  be 
thicker  to  the  east  and  southeast  of  the  cone,  suggesting  a  paleowind  direction  similar  to 
the  present  prevailing  wind  pattern.  This  cone  may  have  been  the  source  for  the  earlier 
basalt  flows.  A  fault  truncates  its  west  side  and  moves  the  western  portion  right-laterally 
about  90-150  m. 

Transitional  lithologies  occur  where  the  first  pyroclastic  eruptives  are  mingled  with 
the  granitic  wash  mantling  the  Table  Mountain  Formation.  Commonly  found  here  are 
volcanic  glass-cemented,  spherical  concretions  of  slightly  granular,  bimodal,  very  coarse- 
and  very  fine-grained,  volcanic  lithic  arkose.  The  coarser  mode  contains  numerous  pluton- 
ic  rock  fragments  along  with  microcline,  orthoclase,  and  perthite  from  the  surrounding 
highlands.  The  finer  mode  comprises  idiomorphic  volcanic  plagioclase,  relict  shards,  1am- 
probolite,  zircon,  biotite,  and  apatite.  The  non-concretionary  arenite  in  places  contains 
montmorillonite  formed  from  altered  pyroclastics. 

West  of  the  major  fault  and  interlayered  with  but  partially  overlying  the  earlier  flows 
is  a  series  of  agglomerate  and  cinder  deposits.  These  pyroclastics  are  distinguished  from 
the  earlier  cone  materials  because  they  overlie  the  lower  basalt,  are  coarser,  lack  sorting, 
and  contain  a  large  percentage  of  bombs  and  agglutinate  material.  Some  of  the  bombs 
and  blocks  in  this  younger  cone  are  up  to  1.3  m  in  diameter,  although  most  average  3  to  5 
cm.  Several  basalt  flow  units  appear  to  have  issued  from  the  base  of  the  younger  cone 
along  the  west  and  northwest  side  of  Table  Mountain.  The  cone  morphology  is  not  ob- 
vious as  it  has  largely  been  destroyed  or  obscured  by  later  deposits.  Cinder  and  scoria  de- 
posits suggestive  of  smaller  eruptive  centers  are  found  on  the  east  side  of  the  border  hill 
(VABM  3572),  in  a  pit  along  Carrizo  Gorge  road  just  north  of  old  Highway  80,  in  a  pit 
east  of  Round  Mountain  and  south  of  Interstate  8,  and  in  the  low  hills  north  of  the  free- 
way and  west  of  the  Jacumba  turnoff'.  All  have  an  abundance  of  calcite  cement,  and  in  the 


133 


eruptive  center  just  south  of  the  freeway  the  calcite  percentage  is  so  high  that  the  cinders 
initially  appear  to  lack  a  supporting  framework. 

ANDESITE  PLUGS.-Round  Mountain  and  the  hill  at  Jade  Benchmark  (Map-Fig.  2) 
are  plugs  whose  chemistry  differs  significantly  from  the  other  volcanic  rocks  in  the  valley 
(Hawkins,  1970).  They  contain  significant  percentages  of  hypersthene  in  place  of  the 
hornblende  typical  of  the  basalt  flows.  These  plugs  are  largely  intact  and  have  typical  on- 
ion-skin jointing  and  a  bulbous  dome-like  structure.  Both  plugs  contain  xenoliths  of  gran- 
odiorite,  quartz  diorite,  and  schist  from  the  basement  rock. 

The  plugs  are  younger  than  at  least  that  part  of  the  andesite  breccia  sequence  which 
they  rest  upon.  No  clear-cut  evidence  can  be  presented  for  their  minimum  age  and  they 
could  be  much  younger  than  the  lavas  and  andesites.  Similar  appearing  plugs  of  Pliocene 
age  (R.  G.  Gastil,  pers.  comm.)  occur  in  other  parts  of  the  Peninsular  Range  near  the  in- 
ternational boundary. 

ANDESITE  BRECCIA,  LAHAR  DEPOSITS,  CONGLOMERATE,  AND  SANDSTONE.-A  hetero- 
geneous sequence  of  andesitic  lahar  deposits,  breccia,  conglomerate  and  sandstone  form  a 
significant  portion  of  the  Jacumba  Volcanics.  The  andesites  are  up  to  150  m  thick  in  some 
localities  and  generally  average  90  m  thick  over  much  of  the  area.  The  breccia  appears  to 
be  the  dominant  form  of  the  andesite,  but  other  forms  are  more  common  locally.  The  in- 
tergradation  of  the  component  rock  types  and  the  presence  of  a  heavy  lag  gravel  over  the 
andesite  prevented  the  mapping  of  the  individual  andesitic  rock  units. 

The  best  exposure  of  the  andesite  breccia  is  just  east  of  the  Table  Mountain  quarry  in 
NE  '/4,  Sec.  35,  T  17  S,  R  8  E  (Map-Fig.  2).  There  the  breccia  consists  of  red-brown  to 
brown  to  gray  massive  hornblende  andesite  which  has  been  intensely  sheared,  broken, 
and  comminuted  to  form  a  flow  breccia.  No  bedding  or  stratification  can  be  discerned  at 
the  outcrop. 

The  andesite  breccias  grade  into  lahars  (mudflows  of  volcanic  detritus).  In  the  vicin- 
ity of  the  Table  Mountain  quarry  an  approximately  9  m  thick  lahar  deposit  consists  of  a 
red-brown  to  brown,  massive  andesitic  breccia  with  andesite  and  cinder  particles  ranging 
from  dust  to  small  boulder  size.  The  complete  lack  of  bedding  and  sorting  coupled  with 
monohthologic  andesite  fragments  are  characteristic  of  a  lahar  deposit.  Intermittent  flu- 
vial action  has  reworked  the  breccia  and  lahar  deposits  producing  volcaniclastic  con- 
glomerate and  sandstone  which  occur  throughout  the  section.  These  fluvial  units  resemble 
the  breccia  in  clast  composition,  and  they  are  distinguished  by  the  presence  of  bedding 
and  of  increased  sorting  of  the  clasts.  A  typical  sandstone  exposed  above  the  quarry  on 
Hill  4089  (SW  '/4,  Sec.  26,  T  17  S,  R  8  E)  is  a  very  poorly  sorted,  very  angular  to  subangu- 
lar,  pebbly  medium  sandstone.  This  and  similar  sandstone  beds  are  lithic  arkoses  and  are 
loaded  with  volcanic  rock  fragments  and  extrusive  euhedral  minerals,  while  plutonic-de- 
rived  sediment  is  usually  absent. 

Mixed  plutonic-volcanic  lithic  arkoses  are  found  between  tuff's  (below)  and  volcanic 
mudflows  (above)  on  the  south  side  of  Round  Mountain.  This  mixed-provenance  lith- 
ology  illustrates  the  intermittent  nature  of  volcanic  sedimentation  that  allowed  inworking 
of  granitic-derived  debris. 

Intercalated  within  the  volcanic  sequence  west  of  Gray  Mountain  is  a  volcanic  lithic 
arkose  mass  exhibiting  0.6  to  1.3  m  thick  planar-wedge  sets  of  cross-laminae.  These  fine 
sandstone  grains  are  mostly  plagioclase,  volcanic-rock  fragments,  hypersthene,  horn- 
blende, and  biotite  that  are  moderately  sorted,  skewed  toward  the  fines,  and  subrounded 
to  subangular.  The  sedimentary  structures,  presence  of  abrasion,  fine  grain  size,  and  best 
sorting  in  the  valley  all  indicate  an  episode  of  reworking  of  volcanic  sediment  into  eolian 
dunes. 

The  various  units  of  the  Jacumba  Volcanics  are  considered  to  be  of  Early  Miocene 
age.  The  K/Ar  whole  rock  age  date  of  18.7  ±1.3  m.y.  (Hawkins,  1970)  for  a  basalt  on  Ja- 
cumba Peak  corresponds  with  concordant  hornblende  and  plagioclase  dates  of  18.5  ±0.9 
m.y.  and  18.6  ±0.8  m.y.,  respectively,  obtained  from  a  clast  in  the  andesite  breccia  on 
Table  Mountain  (K/Ar  laboratory  CSUSD).  Thus,  the  bulk  of  the  Jacumba  Volcanics 
were  erupted  in  the  Early  Miocene. 


134 


STRUCTURE 

The  structure  of  the  Jacumba  area  is  dominated  by  a  series  of  northwest-trending 
normal  and  reverse  faults.  Two  east-trending  faults  bound  the  area  on  the  north  and 
south.  Only  a  few  northwest-trending  faults  could  be  traced  beyond  the  east-trending 
faults. 

The  northwest-trending  set  of  faults  produces  a  horst  and  graben  effect  within  the 
valley.  Throws  of  up  to  300  m  are  necessary  to  produce  some  of  the  observed  features. 
The  fault  which  bisects  Table  Mountain  has  approximately  90  to  150  m  of  right-lateral 
separation  and  30  to  60  m  of  vertical  separation.  In  many  cases  a  fault  which  can  be 
traced  for  some  distance  with  certainty  in  the  volcanic  rocks  is  lost  within  a  few  feet  in  the 
granitic  rocks.  In  other  cases  a  single  fault  in  the  volcanic  rocks  splits  into  several  seg- 
ments as  it  enters  the  granitic  rocks. 

An  interesting  feature  noted  along  several  large  faults  is  the  inclusion  of  a  thin  sliver 
of  Table  Mountain  Formation  in  the  fault  zone.  In  a  number  of  cases  the  presence  of  a 
fault  was  recognized  by  the  thin  strip  of  conglomerate  between  the  granitic  and  the  vol- 
canic rocks. 

The  two  east-trending  faults  are  primarily  in  granitic  rocks  and  have  helped  erosion 
produce  long  linear  valleys.  The  northern  fault  exhibits  a  0.6  to  1.5  m  shear  zone  along 
the  north  side  of  Table  Mountain. 

The  structure  in  the  Jacumba  Valley  closely  parallels  the  regional  structure  of  the 
northwest  side  of  the  Peninsular  Range.  The  valley  itself  is  directly  along  the  projected 
trend  of  the  Elsinore  fault  zone  as  it  passes  out  of  the  main  mountain  mass  near  Banner 
Grade.  The  right-lateral  separation  on  at  least  one  fault  in  the  Jacumba  area  may  suggest 
that  some  of  the  displacement  on  the  Elsinore  is  taken  up  by  movement  along  its  pro- 
jected trend  in  the  Jacumba  Valley  area. 

ACKNOWLEDGEMENT 

We  would  like  to  thank  George  W.  Moore  for  his  helpful  review  of  this  manuscript. 


LITERATURE  CITED 

Brooks,  B.,  and  E.  Roberts 

1954.  Geology  of  the  Jacumba  area,  San  Diego  and  Imperial  Counties.  In.  Jahns.  R.  H..  ed..  Geology  of 
Southern  California.  Calif  Div.  Mines  and  Geology  Bull.  170,  map  sheet  23. 

Bushee,  J.,  J.  Holden.  B.  Geyer,  and  G.  Gastil 

1963.  Lead-alpha  dates  for  some  basement  rocks  of  southwestern  California.  Geol.  Soc.  Amer.  Bull.  74: 
803-806. 

Fairbanks,  H.  W. 

1893.  Geology  of  San  Diego  County.  Calif.  State  Mining  Bur.  Repl.  11:  76-120. 

Gastil,  G..  and  J.  Bushee 

1961.  Geology  and  geomorphology  of  eastern  San  Diego  County,  p.  8-22.  In,  Field  trip  guidebook,  San 
Diego  County.  Geol.  Soc.  Amer.  (Cordilleran  Section)  57th  Ann.  Mtg. 

Hawkins,  J.  W. 

1970.  Petrology  and  possible  tectonic  significance  of  late  Cenozoic  volcanic  rocks,  southern  California  and 
Baja  California.  Geol.  Soc.  Amer.  Bull.  81:  3323-3338. 

Kennedy,  M.  P..  and  G.  W.  Moore 

1971.  Stratigraphic  relations  of  Upper  Cretacious  and  Eocene  formations,  San  Diego  coastal  area,  Califor- 
nia. Amer.  Assoc.  Petrol.  Geol.  Bull.  55:  709-722. 

Miller,  W.  J. 

1935a.   Geologic  section  across  southern  Peninsular  Range  of  California.  Calif  Jour.  Mines  and  Geology  31: 
115-142. 

1935b.  Geomorphology  of  the  southern  Peninsular  Range  of  California.  Geol.  Soc.  Amer.  Bull.  46:   1535- 
1561. 

Minch,  J.  A. 

1970.  Early  Tertiary  paleogeography  of  a  portion  of  the  northern  Peninsular  Range,  p.  83-87.  //;,  Pacific 
slope  geology  of  northern  Baja  California  and  adjacent  Alta  California.  Amer.  Assoc.  Petrol.  Geol. 
(Pacific  Section)  Fall  Field  Trip  Guidebook. 


i 


135 


Weber.  F.  H..  Jr. 

1963.  Mines  and  mineral  resources  of  San  Diego  County,  California.  Calif  Div    Mines  and  Geoloey 
Count)  Rcpl.  3:  1-309. 


Department  of  Geology,  Saddleback  College,  Mission  Viejo,  California  92675,  and  De- 
partment of  Geology,  California  State  University,  San  Diego,  California  92115. 


s-m-s 


MUS.  COMP.  ZOOL. 
LIBRARY 

JUN221973 

HARVARD 
UNWERS^TYi 

REVISION  OF  THE  CORAL-INHABITING  BARNACLES 
(CIRRIPEDIA:  BALANIDAE) 


ARNOLD  ROSS  AND  WILLIAM  A.  NEWMAN 


TRANSACTIONS 

OF  THE  SAN  DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  12  20  APRIL  1973 


Figure  I.     Boscia  anglicum  on  Carvophvllia  smiihii  Stokes  and  Broderip:  Eddystone,  South  Devon,  England: 
British  Museum  (Nat.  Hist.)  1904.6.27.13.  British  Museum  photograph. 


REVISION  OF  THE  CORAL-INHABITING  BARNACLES 
(CIRRIPEDIA:  BALANIDAE) 

ARNOLD  ROSS  AND  WILLIAM  A.  NEWMAN 


ABSTRACT.— The  biogeography,  growth,  morphology  and  host  specificity  of  all  known  taxa  of  coral-in- 
habiting barnacles  in  the  Pyrgomatinae  are  reviewed.  In  addition  to  Pvri^oma  and  Cretisia.  among  which  all 
of  the  species  were  previously  divided,  we  resurrect  five  genera  and  propose  Hoekia,  Hiroa  and  Cantellius. 
These  10  genera  fall  into  three  groups:  Boscia  (cosmopolitan),  Ceratoconcha  (Pliocene  in  the  eastern  Pacific, 
Mio-Pliocene  in  the  Mediterranean  Basin,  and  as  Miocene  relicts  in  the  western  Atlantic),  and  Cantellius 
and  its  derivatives  (Miocene  to  Recent  in  the  Indo-west  Pacific).  The  Pyrgomatinae  are  apparently  poly- 
phyletic;  Cantellius  and  possibly  Boscia  arose  tYom  different  armatobalanid  stocks,  while  Ceratoconcha 
arose  from  an  indeterminate  balanoid  stock.  Cantellius  and  Ceratoconcha  first  appeared  in  the  Miocene 
during  the  break  up  of  the  Tethyan  Sea  and  the  initiation  of  faunal  provincialism. 

The  reef  coral  community  has  been  characterized  by  Cloud  (1959:  387)  as  essentially 
a  steady-state  oasis  of  high  population  density,  intense  calcium  metabolism,  and  complex 
nutrient  cycling,  generally  surrounded  by  waters  of  relatively  low  nutrient  and  plankton 
content.  Aside  from  interesting  parallels  with  tropical  rain  forests  and  man-made  mega- 
lopolises, Newell  (1971:  2)  argued  that  the  organisms  comprising  the  coral  reef  commu- 
nity are  "superlatively  coadapted."  One  of  the  remarkably  coadapted  animal  groups  is 
the  coralliophilic  pyrgomatines. 

Barnacles  comprising  the  Pyrgomatinae  are  obligatory  symbionts  or  parasites  prima- 
rily of  scleractinian  corals.  They  occur  in  all  regions  of  the  world  that  support  major 
growths  of  hermatypic  corals,  and  they  have  been  found  in  sediments  dating  from  the 
early  Miocene.  Modern  pyrgomatines  were  probably  recognized  by  naturalists  in  the  18th 
century,  but  the  group  did  not  receive  serious  attention  until  the  middle  of  the  19th  cen- 
tury. The  present  study  is  the  first  general  revision  on  a  world-wide  basis. 

HISTORICAL  ACCOUNT 

Studies  on  pyrgomatine  cirripeds  may  be  grouped  into  three  periods.  The  work  of 
Leach  (1817,  1818,  1825),  Sowerby  (1823),  Gray  (1825,  1831),  and  many  other  con- 
chologists  characterize  the  earliest  period  as  one  of  describing  new  taxa  and  grouping 
these  into  a  hierarchy. 

The  second  period,  covering  about  70  years,  began  with  the  publication  of  Darwin's 
(1854)  monograph  of  the  Balanidae  and  Veruccidae.  Darwin  attempted  to  embody  the 
best  features  of  earlier  studies;  however,  he  chose  not  to  follow  the  generic  divisions  pro- 
posed by  Leach  and  Gray,  and  retained  only  Pyrgoma  and  Creusia,  and  the  latter  he  as- 
signed subgeneric  status.  Leach  and  Gray  had  attempted  to  group  a  seemingly  meager 
number  of  species  into  a  maximum  number  of  poorly  defined  genera.  Consequently,  Dar- 
win's conservative  approach  was  generally  accepted;  and  such  caution  has  proved  a  deter- 
rent to  unraveling  the  systematics  of  this  group. 

The  third  period  began  with  the  work  of  Annandale  (1924),  followed  by  Withers 
(1926,  1929),  Hiro  (1935,  1938),  and  Nilsson-Cantell  (1938),  and  more  recently,  by  Brooks 
and  Ross  (1960),  Utinomi  (1962,  1967),  Baluk  and  Radwahski  ( 1967a,  1967b,  1967c),  and 
Ross  and  Newman  (1969).  Annandale,  Hiro,  and  Nilsson-Cantell  provided  names  for  the 
majority  of  Darwinian  numerical  varieties,  and  they  added  considerably  to  our  knowl- 
edge of  the  Indo-Pacific  members  of  the  subfamily. 

Of  the  many  classifications  proposed  for  this  group,  the  earliest  were  taxonomic 
rather  than  phylogenetic,  except  that  of  Gray  (1825:  102),  which  was  based  on  an  ecologi- 
cal concept.  The  most  promising  classification  was  proposed  recently  by  Baluk  and  Rad- 
wahski (1967c)  who  resurrected  the  generic  groupings  initiated  by  Leach,  Gray  and 

SAN  DIEGO  SOC.  NAT.  HIST..  TRANS.  17(12):  137-174.  20  APRIL  1973 


138 


Sowerby,  and  proposed  several  new  names.  The  present  study  somewhat  revises  and 
greatly  extends  their  classification. 

BIOGEOGRAPHY  *" 

Modern  pyrgomatines  occur  in  all  regions  of  the  world  that  support  major  growths  of 
hermatypic  corals.  Fossils  occur  predominantly  in  the  Tertiary  and  Pleistocene  of  the 
western  Atlantic  and  the  Mediterranean  Basin  (Withers,  1929:  2;  Brooks  and  Ross,  1961: 
326;  Baluk  and  Radwahski,  1967c;  Newman  and  Ladd,  in  press).  The  western  Atlantic 
contains  but  a  few  morphologically  primitive  pyrgomatines,  while  the  Indo-west  Pacific 
has  the  greatest  variety  and  abundance  and  the  morphologically  most  advanced  species. 
The  disparity  between  these  faunal  realms  may  relate  to  the  greater  number  of  reef  corals 
available  as  hosts  in  the  Indo-Pacific,  of  which  there  are  80  genera  and  500  species  as 
compared  to  20  genera  and  65  species  in  the  western  Atlantic  (Newell,  1971:  26),  but  the 
latter  has  also  witnessed  a  general  decline  in  the  biota  dating  from  the  Miocene  (Newell, 
1971:  23). 

Three  major  morphological  groups  of  Pyrgomatinae  are  recognized  in  this  paper 
{Ceratoconcha,  Boscia  and  CantelUus  and  its  derivatives),  and  these  have  interesting  im- 
plications. The  first  and  most  generalized  is  the  creusioid  Ceratoconcha,  which  first  ap- 
pears in  sediments  of  lower  Miocene  age.  Based  on  studies  by  Brooks  and  Ross  (1961: 
362),  Baluk  and  Radwahski  (1967c),  and  Newman  and  Ladd  (in  press),  it  is  evident  that 
during  the  Miocene  Ceratoconcha  was  not  only  more  diverse  in  terms  of  species  than  it  is 
today,  but  also  ranged  throughout  the  tropical  Atlantic  and  its  eastern  Pacific  outpost, 
while  it  survives  as  a  Miocene  relict  in  the  western  Atlantic  (Fig.  2).  Ceratoconcha  appar- 
ently never  ranged  into  the  Indo-Pacific,  probably  because  communications  between  the 
Indian  Ocean  and  the  Mediterranean  had  ceased  in  early  Miocene  times  (Ruggieri,  1967: 
284)  with  the  northward  movement  of  the  African  land  mass. 

The  Pliocene  fauna  in  the  Mediterranean  Basin  includes  only  C.  costata  (see  Baluk 
and  Radwahski,  1967c:  483;  Moroni,  1967:  17);  apparently  no  Pleistocene  ceratocon- 
choids  are  found  there.  The  short  stratigraphic  range  of  Ceratoconcha  in  the  Mediterra- 
nean Basin  is  not  surprising,  because  climatic  cooling  which  had  already  begun  in  the 
Oligocene  (Wells,  1956;  Ekman,  1953),  coupled  with  isolation  (Ruggieri,  1967:  284),  re- 
sulted in  a  decline  in,  if  not  total  destruction  of,  the  hermatypic  corals  and  other  tropical 
elements  of  the  fauna. 

The  Mediterranean  ceratoconchoids  are  probably  western  Atlantic  derivatives  de- 
spite the  great  distance  separating  these  two  regions.  Numerous  other  invertebrates  pres- 
ently have  trans-Atlantic  distribution  patterns  (Briggs,  1970),  and  apparently  many  of 
these  animals  have  larvae  that  were  transported  eastward  from  the  western  Atlantic 
(Robertson,  1964:  21;  ScheUema,  1971:  284). 

Ceratoconcha  was  represented  by  at  least  five  species  in  the  western  Atlantic  in  the 
early  Miocene  (Newman  and  Ladd,  in  press),  and  some  time  thereafter  by  a  few  species 
in  the  eastern  Pacific.  At  least  two  species  are  found  in  Pliocene  corals  of  the  Imperial 
Formation  of  the  Carrizo  Creek  and  Coyote  Mt.  areas  of  southern  California  (Ross,  un- 
publ.).  There  are  no  Pleistocene  or  living  ceratoconchoids  in  the  eastern  Pacific.  The  west- 
ern Atlantic  Pliocene  fauna  contains  only  preflorldanum:  the  Pleistocene  fauna  contains 
barbadensis,  possibly  prefJoridanum,  and  several  undescribed  species  (Brooks  and  Ross, 
1960:  362).  The  Recent  western  Atlantic  contains  two  or  possibly  three  species. 

The  second  group,  containing  only  the  primitive  pyrgomoid  Boscia  (Fig.  2),  has  been 
found  in  sediments  of  Pliocene  age  in  the  Mediterranean  Basin  (Baluk  and  Radwahski, 
1967c:  483)  and  England  (Darwin,  1854b;  Withers,  1926).  Pleistocene  occurrences  include 
Italy  (Alessandri,  1906)  and  Japan  (Sakakura,  1938).  Although  Sakakura  reported  the  in- 
dividuals he  found  on  an  ahermatypic  coral  as  anglicum,  restudy  of  these  may  reveal  that 
they  represent  either  a  new  species  or  oulastreae  (see  Utinomi,  1967:  232),  since  anglicum 
appears  to  be  restricted  to  the  western  Mediterranean  and  eastern  Atlantic  (Moyse,  1961: 
384;  Utinomi,  1967:  231;  cf  Rees,  1962:  412).  Boscia  occurs  on  hermatypic  corals  in  the 
western  Atlantic  (madreporarum)  and  western  Pacific  (oulastreae),  whereas  in  the  eastern 
Atlantic  and  Mediterranean  (anglicum)  it  settles  only  on  ahermatypic  corals.  There  are  no 


139 


records  of  Boscia  in  the  eastern  Pacific. 

The  third  group  is  wholly  Indo-west  Pacific  with  species  ranging  from  the  Red  Sea  to 
the  Great  Barrier  Reef  and  to  the  Line  Islands  (Fig.  2  and  3).  There  are  but  two  Miocene 
records  for  the  eight  genera  in  this  group,  but  the  specimens  have  not  yet  been  identified 
(Newman  and  Ladd,  in  prep.).  Nubia,  Cantellius,  and  Savignium  are  found  throughout 
the  Indo-Pacific,  but  only  Savignium  ranges  as  far  south  as  the  Great  Barrier  Reef  and  as 
far  east  as  the  Line  Islands.  Pyrgoma,  Creusia,  and  Hoekia  range  from  eastern  India  to  Ja- 
pan, although  Hoekia  has  been  reported  from  Mauritius  (Ross  and  Newman,  1969).  There 
is  only  one  record  for  Hiroa,  in  the  Caroline  Islands. 

From  the  foregoing,  two  provincial  coral-barnacle  faunas  can  be  recognized,  one 
centering  in  the  Caribbean  portion  of  the  western  Atlantic  and  the  other  in  the  Austra- 
lasian portion  of  the  Indo-Pacific.  Comparable  biogeographic  patterns  have  long  been 
recognized  in  other  invertebrates  and  in  fishes  (see  Briggs,  1970).  The  coral  barnacles 
were  evolving  when  the  continuity  of  the  Tethyan  Sea  was  being  destroyed,  ultimately 
leading  to  faunal  provincialism.  In  light  of  the  geological  history  of  these  regions  and  con- 
sidering the  morphological  features  of  these  two  groups,  it  is  apparent  that  they  devel- 
oped independently  in  the  two  regions  from  different  balanoid  ancestors.  Boscia,  which 
appears  to  be  a  third  independent  group,  may  owe  its  widespread  distribution  to  its  abil- 
ity to  settle  on  deep-water  ahermatypic  corals. 

GROWTH  AND  FORM 

The  early  growth  stages  of  pyrgomatines  look  much  like  those  of  ordinary  balanids. 
It  is  in  the  later  stages  that  their  adaptations  to  an  intracoralline  hfe  become  evident. 
Knowledge  of  the  larval  stages  is  limited:  Kolosvary  (1950:  293)  described  typically  bala- 
noid nauplii  of  Savignium  milleporae  and  Moyse  (1961:  371)  described  all  larval  stages  of 
Boscia  anglicum.  Duerden  (1904:  39)  suggested  that  the  cyprid  bores  through  the  living 
tissue  of  the  polyp  and  that  in  the  process  of  growth  the  skeletons  of  the  two  become 
fused. 

Utinomi  (1943:  16)  followed  the  ontogeny  of  the  earliest  juveniles  of  Creusia  indicum 
Annandale,  and  found  that  the  juvenile  does  not  initially  attach  to  the  coral  skeleton  but 
remains  imbedded  in  the  coral  tissue.  While  the  four  plates  making  up  the  wall  and  the 
opercular  valves  are  calcified,  the  cup-shaped  basis  of  the  juvenile  is  wholly  membranous. 
Even  after  the  basis  calcifies  there  is  a  period  when  the  juvenile  remains  free  in  the  coral 
tissue  before  the  basis  and  corallites  come  into  contact  and  fuse.  Moyse  (1971:  127)  noted 
similar  relationships  in  Boscia  anglicum. 

Subsequent  growth  is  rapid,  especially  laterally,  so  that  the  shell  reaches  essentially 
maximum  diameter  early  in  life.  This  is  well  illustrated  by  Hiro  (1938,  fig.  1 1  and  12).  In 
general,  the  wall  becomes  proportionately  less  conical  as  its  diameter  and  basal  height  in- 
crease, the  aperture  enlarging  by  diametric  growth  in  four-plated  forms,  or  by  corrosion 
and  cirral  rasping  in  single-plated  forms.  In  the  scatter  diagram  plotted  by  Hiro  (1938,  fig. 
12)  for  Creusia  indicum,  after  the  period  when  the  basal  height  and  shell  width  increase 
uniformly,  width  stabilizes  while  basal  height  continues  to  increase,  as  it  must  throughout 
the  life  of  the  barnacle.  During  the  early  period  of  rapid  increase  in  width,  the  barnacle 
may  rotate  its  position  by  as  much  as  90°  (Baluk  and  Radwahski,  1967b,  fig.  2,  1;  New- 
man and  Ladd,  in  press,  pi.  2h). 

Creusioids  with  well  developed  radii  are  commonly  overgrown  to  some  extent  by  the 
coral,  and  enlargment  in  both  basal  height  and  shell  diameter  requires  breaking  the  over- 
growth along  the  sutures.  Creusioids  with  radii  indicated  by  simple  sutures,  and  pyrgo- 
moids  in  general  (except  Boscia,  see  Moyse,  1971),  have  the  ability  to  suppress  coral 
skeleton  deposition  around  the  margin  of  the  shell  so  that  vertical  growth  can  proceed 
without  mechanical  breakage.  In  some  cases,  the  coral  may  lay  down  skeletal  elements  on 
the  wall  of  the  barnacle  suggestive  of  normal  septa,  and  the  barnacle  then  takes  on  the 
appearance  of  a  corallite  (Duerden,  1904:  39);  this  is  an  unusual  form  of  mimicry  to  say 
the  least.  In  other  cases,  only  coral  tissue  grows  over  the  wall  of  the  barnacle,  and  in 
Hoekia  this  tissue  proliferates  over  the  aperture  where  it  is  fed  upon  by  the  barnacle 
(Ross  and  Newman,  1969). 


140 


140°  120'  100°  60'  60*  40'  20'  »   0°  E  ;0'  40°  60° 


1 r 


100°  120°  140°  160°        E   160°  W        160°  140° 


-^ 


Caniellius 


140°  120°  100°  80°  60°  40°  20°  W  0°  E  20°  40°  60° 


100°  120°  140°  160°        E   180°  W        160°  140° 


140°  120°  100° 


60°  40°  20°  W  0°  E  20°  40°  60' 


100°  120°  140°  160°        E   180°  *        160°  140° 


60°        E   160°  W        160° 


Figure  2.     Distributional  records  for  Nobia,  Fyrgoma,  Hiroa,  Savignium,  and  Creusia.  Data  from  same  sources 
as  Figure  3. 

In  specimens  of  Savignium  crenatum  that  we  have  observed  growing  between  low 
branches  of  the  surface  of  Merulina  ampliala,  the  rate  of  growth  of  the  barnacle  exceeds 
that  of  the  coral  so  that  the  barnacle  extends  well  above  the  general  surface  of  the  coral- 
lum.  In  most  cases  a  thin  layer  of  coral  skeleton  grows  up  onto  the  surface  of  the  basis  of 
the  rapidly  advancing  barnacle,  aiding  in  its  support,  but  in  some  a  fair  proportion  of  the 
basis  stands  free  of  the  coral.  While  it  might  appear  that  the  barnacle's  growth  rate  is  sim- 
ply out  of  phase  with  that  of  the  coral,  there  is  adaptive  value  in  growing  in  this  manner. 
The  barnacles  are  growing  up  between  branches  of  the  coral  which  will  eventually  fuse 
laterally  at  higher  levels.  If  the  barnacles  simply  kept  pace  with  the  growth  of  the  surface, 
they  would  more  than  likely  be  buried. 

Boscia  anglicum  grows  in  a  similar  manner,  but  on  solitary  ahermatypic  corals,  along 
the  margin  of  the  corallite  (Fig.  1).  In  this  position  there  is  relatively  little  interference 
with  the  normal  feeding  mechanism  of  the  coral.  Established  individuals  frequently  serve 
as  sites  for  subsequent  generations.  Cloud  (1959:  392)  suggested  that  the  barnacles  re- 
place the  coral  polyps,  and  although  this  is  certainly  not  true  here,  it  may  more  frequently 


141 


IJO"  120'  100' 


100°  80*  60'  40'  20°  W  0°  t  20°  '0'  60 


\2<y  IW  160"        E   180"  *        160° 


Figure  3.  Distributional  records  for  Caniellius.  Hoekia,  Ceratoconcha,  Pyrgopsella,  and  Boscia.  Circles  and 
squares  represent  Recent  records,  triangles  fossil  records.  Data  based  on  specimens  in  the  American  Museum, 
San  Diego  Natural  History  Museum,  Scripps  Institution  of  Oceanography.  Florida  State  Museum,  British  Mu- 
seum (Natural  History),  Museum  of  Comparative  Zoology,  Harvard  University,  and  available  literature. 


142 


be  true  in  those  corals  having  smaller  caHces.  Duerden  (1904:  39)  found  that  in  Side- 
rastrea  radians.  Ceratoconcha  fixes  itself  in  the  calicinal  cavity,  never  on  the  ridges  con- 
necting two  calices  and  that  the  presence  of  the  barnacle  results  in  imperfections  in  the 
surrounding  polyps. 

The  only  pyrgromatine  growing  on  the  stinging  hydrocoral  Millepora  is  Savignium 
milleporae.  It  is  not  uncommon  to  find  a  thick-walled  chimney  of  the  host  skeleton  ele- 
vated about  5  mm  above  the  general  surface  of  the  coral  supporting  the  barnacle.  The 
basis  of  the  barnacle  occupies  the  whole  chimney,  with  the  initial  point  of  attachment  es- 
sentially at  the  level  of  the  surrounding  colony.  The  top  of  the  chimney  is  flush  with  the 
flat  top  of  the  barnacle.  Evidently  the  general  surface  of  the  coral  does  not  grow  fast 
enough  to  accommodate  the  rapidly  growing  body  chamber  of  the  barnacle.  Whether  the 
barnacle  is  able  to  regulate  the  growth  rate  of  the  coral,  so  that  the  supporting  chimney  is 
formed,  or  the  coral  is  simply  reacting  to  the  presence  of  a  foreign  object  and  attempting 
to  bury  it,  has  not  been  determined.  Interestingly,  Balamis  stultus,  the  only  other  barnacle 
occurring  on  Millepora,  likewise  extends  well  above  the  general  surface  of  the  coral.  It  is 
also  covered  by  a  layer  of  coral  skeleton,  but  it  appears  to  be  simply  encrusted,  rather 
than  contained  within  a  thick-walled  chimney  as  is  S.  milleporae.  Balanus  stultus  contin- 
ues to  grow  diametrically  by  fracturing  the  coral  skeleton  along  the  sutures  in  the  wall. 
Thus  in  both  cases  the  coral  is  reacting  in  ways  that  favor  the  diff'erent  growth  habits  of 
the  barnacles,  and  this  suggests  that  these  barnacles  are  exercising  some  control  over  the 
growth  habits  and  defense  mechanisms  of  the  coral. 

In  general,  shell  color  in  the  pyrgomatines  is  white.  Boscia  juveniles  have  a  white 
shell,  but  with  later  growth  the  shell  takes  on  a  pinkish  or  pinkish-purple  hue.  Hoekia  has 
a  pinkish-purple  shell,  while  that  of  Nobia  is  white  and  splotched  with  pink  or  purple.  In 
Cantellius  some  species  are  all  white,  whereas  in  others  the  apical  portion  of  the  opercular 
plates  is  tinted  purple.  The  shell  in  Savignium  is  commonly  pinkish  red,  and  in  Pyrgoma  it 
is  a  pale  pink.  Ceratoconcha  is  invariably  white.  While  the  basis  is  never  pigmented,  the 
exposed  shell  of  most  genera  is.  Consequently,  there  must  be  some  adaptive  or  functional 
significance  to  these  colors.  Since  the  colors  do  not  match  those  of  their  hosts,  they  appar- 
ently do  not  serve  as  protective  coloration. 

Generalized  creusioids  have  well  developed  radii,  and  undergo  diametric  growth 
during  the  better  part  of  their  lives.  The  radii  range  in  form  from  triangular  to  rectangu- 
lar, or  they  may  be  indicated  externally  simply  by  sutures.  When  the  radii  are  triangular, 
the  base  of  the  isosceles  triangle  forms  part  of  the  apertural  margin  and  indicates  that  the 
aperture  has  enlarged  disproportionately  to  the  total  diameter  of  the  shell.  Rectangular 
radii  indicate  proportionate  increments.  Where  radii  are  evidenced  simply  by  sutures,  di- 
ametric growth  has  all  but  terminated,  and  the  total  diameter  of  the  wall  can  increase 
only  by  marginal  increments.  The  aperture  either  remains  the  same  or  is  enlarged  by  cor- 
rosion and  (or)  by  the  rasping  eff'ect  of  cirral  movement;  such  forms  have  eff'ectively 
reached  the  pyrgomoid  level  of  organization. 

The  surface  of  the  basis  is  commonly  marked  by  longitudinal  ribs,  corresponding  to 
the  internal  radiating  ribs  of  the  wall,  and  by  transverse  growth  lines.  The  growth  lines 
are  generally  very  fine,  ranging  between  4  and  24  per  mm  (Newman  and  Ladd,  in  press), 
and  are  interrupted  by  discontinuities  at  more  or  less  regular  intervals  (Baluk  and  Rad- 
wahski,  1967a,  fig.  2;  Newman  and  Ladd,  in  press,  pi.  l,b).  The  interruptions  are  fre- 
quently at  intervals  of  5  mm  or  so  and  probably  correspond  to  the  annual  density  bands 
in  coral  described  by  Knutson  et  al  ( 1972:  270).  This  suggests  that  coral  barnacles  live  for 
several  years,  which  agrees  with  the  age  estimate  given  by  Hiro  (1938:  410).  Unfortu- 
nately, the  barnacles  in  which  these  bands  have  been  observed  are  fossil  forms  that  have 
been  leached  out  of  the  coral  so  that  the  host  species  is  unknown.  With  intact  specimens, 
agreement  between  the  bands  in  the  coral  and  the  barnacle  probably  could  be  deter- 
mined by  the  x-ray  techniques  employed  by  Knutson  et  al  (1972),  but  such  work  remains 
to  be  done. 

Coral  barnacles  do  not  live  as  long  as  their  hosts,  and  eventually  they  become  en- 
tombed. In  some  cases  the  opercular  parts  of  the  entombed  barnacles  are  cemented  in  the 
position  they  occupied  in  life,  while  in  others  they  have  fallen  into  the  body  chamber.  In 


143 


the  first  case  the  coral  undoubtedly  overwhelmed  the  barnacle  while  alive.  This  could  be 
true  in  the  second  case,  although  it  may  be  that  the  barnacle  died  before  the  coral  over- 
grew it.  In  any  event,  the  coral  usually  forms  a  "stopper,"  growing  into  the  aperture  a 
short  distance,  before  attaining  a  normal  growth  pattern  over  the  barnacle  (Baluk  and 
Radwahski,  1967c:  490). 

To  the  best  of  our  knowledge  all  pyrgomatines  have  solid  walls,  at  least  fundamen- 
tally. Some  species  develop  parietal  tubes  where  the  longitudinal  ribs  on  the  interior  of 
the  wall  become  fused  with  the  sheath,  while  others  form  tubes  between  external  longitu- 
dinal ribs.  In  still  others,  where  the  sheath  becomes  fully  fused  to  a  much  thickened  wall, 
several  rows  of  more  or  less  regularly  spaced  tubes  develop.  In  none  of  these  cases  are  the 
tubes  formed  in  the  same  way  as  in  the  tubiferous  balanids  (subgenera  Balanus  and  Mega- 
halanus)  where  interlaminate  figures  can  be  observed  in  the  longitudinal  septa  separating 
the  inner  and  outer  laminae  of  the  wall. 

The  ontogenetic  and  phylogenetic  development  of  tubiferous  walls  has  been  ana- 
lyzed in  a  number  of  cases  (Costlow,  1956;  Newman  et  al,  1967;  Ross  and  Newman, 
1967),  but  their  function  has  only  been  a  point  of  speculation.  A  few  systematists  have 
suggested  applying  the  general  engineering  principle  that,  for  a  given  amount  of  material, 
a  properly  designed  tubiferous  structure  would  be  mechanically  stronger  than  a  solid  one. 
If  it  were  necessary  for  a  barnacle  to  be  economical  in  its  use  of  calcium  carbonate,  then  a 
tubiferous  wall  should  be  advantageous  in  high  energy  environments.  Barnes  et  al  (1972) 
tested  the  resistence  of  certain  species  to  impaction  and  found  that  breakage  occurred  not 
in  the  plates  themselves  but  at  the  sutures  between  them.  They  concluded  that  the 
strength  of  the  plates  generally  exceeded  the  strength  of  the  articulating  joints.  The  nature 
of  the  articulation  between  the  wall  and  calcareous  basis  is  also  of  great  importance  (New- 
man et  al,  1967:  170).  These  structural  features  are  well  developed  in  the  pyrgomatines. 
However,  wall  strength  in  coral  barnacles  can  hardly  be  related  to  withstanding  impac- 
tion as  in  many  free-living  forms,  but  rather  is  related  to  the  pressures  required  to  sustain 
growth  in  an  intracoralline  habitat. 

Considering  the  array  and  independent  occurrences  of  tubiferous  walls,  and  the  sec- 
ondary modifications  found  in  them,  e.g.  sealing  off  into  chambers,  secondarily  filling 
with  calcareous  material,  or  filling  with  chitin  during  construction,  one  might  look  for 
some  adaptive  value  other  than  simply  strength.  Ross  (1970:  9)  and  Newman  and  Ross 
(1971)  suggested  that  such  adaptations  might  include  defensive  mechanisms  against  bo- 
rers, specifically  against  the  drilling  of  gastropods.  In  this  regard,  Orton  (1927:  653)  noted 
that  "oysters  are  frequently  attacked  and  abandoned  (by  gastropods) . .  .  if  either  a  cham- 
ber or  loose  horny  layer  is  encountered  . .  ."  It  would  be  expected  that  free-living  barn- 
acles, which  are  frequently  attacked  by  gastropods,  would  also  have  developed  defense 
mechanisms  against  them.  However,  in  the  pyrgomatines  predation  by  borers  has  not 
been  reported.  Their  tubiferous  walls,  then,  developing  in  different  ways  in  different 
members  of  the  group,  undoubtedly  have  some  other  function.  Strength  is  probably  the 
important  one,  but  it  is  also  likely  that  these  tubes  allow  for  physiological  interactions  be- 
tween the  barnacle  and  its  host.  In  many  species  the  tubes  are  arranged  so  as  to  leave  gaps 
around  the  margin  of  the  shell,  which  appear  to  allow  the  uncalcified  integument  of  the 
barnacle  to  come  into  intimate  contact  with  the  tissue  of  the  coral.  Moyse  (1971)  sugges- 
ted that  the  barnacle  may  receive  metabolic  substances  from  the  host  by  this  route.  How- 
ever, we  believe  it  more  likely  or  important  that  these  are  the  sites  where  physiological 
control  of  coral  growth  are  initiated. 

The  opercular  valves  function  to  guard  the  aperture  and  range  in  form  from  wholly 
balanoid  to  highly  modified.  In  Cantellius  and  Ceratoconcha.  the  two  most  generalized 
genera,  the  four-plated  wall  varies  from  high  conic  to  virtually  flat.  Yet  the  valves  are  al- 
ways tall  and  typically  balanoid.  The  same  can  be  said  of  Boscia,  except  that  it  has  a  con- 
crescent  shell.  In  these  three  genera  the  terga  as  well  as  the  scuta  occlude  the  aperture. 

In  the  Savignium  line  (Fig.  5),  the  opercular  valves  are  generally  thin  and  fragile,  and 
the  wall  is  totally  concrescent.  The  scuta  are  relatively  elongate  and  the  reduced  terga  be- 
come completely  fused  to  them.  Likewise,  the  aperture  is  elongate,  and  it  is  guarded 
primarily  by  the  scuta.  The  epitome  of  modified  valves  is  seen  in  Hoekia.  However,  it  has 


144 


a  minute  orifice  and  tiiis  is  related  to  its  wholly  parasitic  way  of  life  (Ross  and  Newman, 
1969). 

In  the  Hiroa  lineage  the  opercular  valves  tend  to  remain  balanoid,  although  the  scuta 
alone  occlude  the  aperture.  Modifications  within  the  lineage  include  elongation  and  nar- 
rowing of  the  terga  and  reduction  of  articular  margins  on  one  hand  {Hiroa,  Pvrgoma), 
and  broadening  and  concrescence  on  the  other  (Nobia,  Creusia).  In  Hiroa- Pyrgoma  the 
opercular  plates  are  relatively  thin  and  fragile  while  in  Nobia-Creusia  they  are  thick  and 
massive.  This  disparity  correlates  to  some  extent  with  size,  but  the  difference  probably 
also  relates  to  the  amount  of  protection  each  requires  from  predators.  Baluk  and  Rad- 
wahski  ( 1967c:  463),  with  reference  to  Darwin's  plate  13,  fig.  ld,(Fig.l2,  c  herein),  misun- 
derstood the  anatomical  relationships  between  the  opercular  valves  and  the  wall  in  Nobia, 
and  concluded  that  the  valves  no  longer  guard  the  aperture.  Apparently,  they  were  not 
distinguishing  between  the  scutal  and  tergal  portions  of  the  concrescent  valves  and 
thought  that  the  occludent  margins  of  the  scutal  portions  were  fused  together  and  no 
longer  functional. 

In  summary,  cyprids  of  coral  barnacles  apparently  first  settle  on  coral  tissue  where 
they  metamorphose  into  juveniles.  A  juvenile  doesn't  attach  to  the  coral  skeleton  until  af- 
ter the  cup-shaped  basis  has  become  calcified.  During  this  period,  and  to  some  extent  af- 
ter attachment,  the  juvenile  may  undergo  reorientation  in  relation  to  the  host  of  as  much 
as  90°.  Unlike  ordinary  barnacles,  subsequent  growth  is  primarily  through  elongation  of 
the  basis  rather  than  the  wall.  Species  with  radii  generally  undergo  diametric  growth  and, 
in  the  process,  frequently  fracture  the  coral  skeleton  overgrowing  them.  Advanced  species 
apparently  gain  a  degree  of  control  over  coral  tissue,  and  its  ability  to  lay  down  new  skele- 
ton. While  barnacles  live  for  several  years,  they  eventually  become  entombed. 

HOST  SPECIFICITY 

Gray  (1825:  102)  proposed  the  Pyrgomatidae  to  accommodate  several  balanoid  gen- 
era peculiar  to  certain  zoophytes;  Pvrgoma  and  Creusia  imbedded  in  scleractinian  corals, 
Conopea  in  gorgonians  and  Acasta  in  sponges.  The  unification  of  these  genera  under  one 
family  was  based  primarily  on  comparable  habitats. 

Figure  4  summarizes  available  data  on  distribution  of  the  various  genera  of  Pyr- 
gomatinae  among  the  scleractinian  suborders.  Of  the  ten  genera  all,  except  Pyrgopsella  in 
sponges  (not  included  in  the  figure)  and  Savignium  milleporae  on  nine  species  of  Mille- 
pora,  occur  exclusively  on  hermatypic  and  ahermatypic  corals.  Of  these,  seven  genera  oc- 
cur on  Faviina,  five  on  Fungiina,  five  on  Astrocoeniina,  four  on  Dendrophylliina,  and  two 
on  Caryophylliina.  Faviina  then,  with  the  greatest  diversity  of  genera,  supports  the  great- 
est diversity  of  coral  barnacles.  Caryophylliina,  while  nearly  equal  to  Faviina  in  numbers 
of  genera,  supports  the  least.  This  is  no  doubt  because  Faviina,  Fungiina,  and  Astrocoe- 
niina are  hermatypic,  while  Dendrophylliina  and  Caryophylliina  are  ahermatypic  with 
representatives  ranging  into  deep  water.  Balanoids  in  general  are  shallow  water  organ- 
isms. 

Only  the  cosmopolitan  genus  Boscia  is  known  to  inhabit  all  five  scleractinian  subor- 
ders and  it  is,  as  far  as  opercular  valves  are  concerned,  among  the  most  generalized  of  the 
Pyrgomatinae.  Hoekia,  Pyrgoma,  and  Hiroa  are  each  limited  to  but  one  scleractinian  sub- 
order, and  each  is  monotypic.  Of  these,  the  first  two  are  among  the  more  specialized 
members  of  the  subfamily,  Hoekia  being  the  most  specialized  balanoid  known.  Hiroa  on 
the  other  hand  resides  at  the  stem  of  the  other  higher  forms  {Creusia,  Nobia,  and  Pyr- 
goma) which,  between  themselves  share  all  five  scleractinian  suborders,  with  Nobia,  a  rel- 
atively highly  modified  form,  occurring  on  four  of  them. 

Ceratoconcha  is  one  of  the  most  generalized  forms,  yet  it  inhabits  but  two  of  the  scle- 
ractinian suborders.  In  being  an  Atlantic  genus,  it  has  survived  in  a  situation  where  coral 
diversity  has  declined  since  the  Oligocene  or  Miocene  (see  Biogeography).  The  remaining 
generalized  genus,  Cantellius,  stands  at  the  stem  of  the  Indo-Pacific  members  of  the  sub- 
family and  is  well  represented  on  the  three  principal  shallow  water  suborders,  Faviina, 
Fungiina,  and  Astrocoeniina. 

From  the  foregoing,  what  can  be  said  of  host  specificity  among  Pyrgomatinae  at  the 


145 


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MILLEPORA 

Figure  4.  Distribution  of  genera  of  Pvrgomatinae  among  scleractinian  coral  suborders  derived  from  the  liter- 
ature. Space  for  each  coral  suborder  is  proportional  to  the  total  number  of  coral  genera  within  each  suborder; 
bar  width  indicates  number  of  occurrences  of  barnacles  within  each  suborder;  bar  height  represents  the  number 
of  coral  genera  on  which  the  barnacles  are  known  to  occur. 

scleractinian  subordinal  level?  One  might  expect  highly  modified  forms  to  be  highly  host 
specific,  and  for  such  extremes  as  Hoekia  this  is  indeed  the  case.  However,  Nobia  is 
among  the  most  modified  forms,  yet  it  occurs  on  all  scleractinian  suborders  except  As- 
trocoeniina,  while  Hiroa,  an  intermediate  in  the  transition  between  Cantellius  and 
Creusia,  has  been  found  only  on  Astrocoeniina.  Savignium  and  Creusia  exploit  mainly  Fa- 
viina  while  Cantellius  exploits  mainly  Fungiina  and  Astrocoeniina,  but  the  division  is  not 
precise  as  there  is  considerable  overlap  among  the  three. 

What  then  seems  to  be  the  situation  as  regards  host  specificity  at  the  specific  level? 
Hiro  (1935:  23)  found  that  in  Tanabe  Bay,  Japan,  ''A  given  species  of  Pyrgoma  is  prac- 
tically confined  to  a  single  species  of  coral,  whereas  the  same  variety  of  Creusia  may  be 
found  on  various  kinds  of  coral."  In  1938  he  reported  on  his  findings  in  the  more  tropical 
Palau  Islands.  While  expanding  the  number  of  corals  playing  host  to  species  of  Pyrgoma 
(1938:  404),  he  again  came  to  the  same  general  conclusion  (1938:  392).  Presently,  his 
statement  holds  best  for  the  monotypic  genera. 

The  suborder  Faviina  plays  host  to  most  of  the  genera  of  Pyrgomatinae.  Of  the  Fa- 
viina,  Pontes  apparently  has  been  selected  most  frequently  as  a  host.  Other  balanids  also 
have  invaded  Pontes,  the  Recent  armatobalanid  Balanus  allium,  the  Pliocene  to  Recent 
armatobalanid  Balanus  durhami  and  the  Miocene  balanid  Balanus  duvergieri  are  notable. 
Growth  on  Pontes  is  independent  of  the  growth  of  individual  coral  polyps  and  con- 


146 


sequently  requires  little  specialization.  This  appears  to  us  to  be  the  reason  why  Porites 
plays  host  to  a  variety  of  barnacles.  Millepora,  on  the  other  hand,  does  not,  and  one  might 
suspect  that  this  is  due  to  its  stinging  ability. 

In  summary  then— 1)  the  greatest  diversity  of  Pyrgomatinae  in  terms  of  numbers  of 
genera  is  found  among  hermatypic  suborders,  particularly  Faviina  and  this  is  probably 
because  balanids  in  general  are  shallow  water  organisms;  2)  there  are  no  marked 
differences  between  the  occurrences  of  pyrgomoids  and  creusioids  as  a  whole  on  the  vari- 
ous suborders  of  scleractinian  corals— in  both  groups  Faviina  is  preferred,  with  scattered 
occurrences  between  the  other  scleractinian  suborders;  and  3)  the  rule  (Hiro  1938:  408), 
that  the  more  peculiar  the  morphological  characteristics  of  species,  the  more  rigid  their 
host  specificity,  holds  in  a  general  way.  The  same  rule  holds  only  weakly  when  applied  to 
barnacle  genera  and  scleractinian  suborders,  for  some  relatively  specialized  genera,  such 
as  Nobia  and  Boscia,  occur  on  a  wide  variety  of  corals  and  are  notable  exceptions. 

ORIGIN  AND  EVOLUTION 

The  Pyrgomatinae  are  a  well  defined  group  (Baluk  and  Radwahski,  1967b:  465),  but 
to  what  lineage  of  the  Balaninae  the  subfamily  owes  its  origin  has  not  been  resolved.  Al- 
though the  consensus  is  that  the  Pyrgomatinae  are  polyphyletic,  only  the  broader  outlines 
of  their  evolution  have  been  elucidated  (Withers,  1929:  564;  1935:  38;  Hiro,  1938:  402, 
412;  Zullo,  1961:  72;  1967:  127;  Baluk  and  Radwahski,  1967c:  500).  Existing  problems 
stem  from  a  lack  of  critical  data  on  fossil  and  Recent  forms,  as  well  as  from  Darwin's 
(1854)  conservative  handling  of  genera  and  species.  His  treatment  of  Creusia  as  a  sub- 
genus of  Pyrgoma  has  not  been  accepted  by  later  workers.  Also,  his  reluctance  to  recog- 
nize geographic  populations  of  Creusia  as  species,  even  though  a  sample  from  a  given 
locality  showed  markedly  uniform  characteristics  (Darwin,  1854:  376),  and  the  failure  of 
subsequent  workers  to  rectify  this,  resulted  in  a  plethora  of  subspecific  and  in- 
frasubspecific  taxa  that  make  little  sense  biologically.  Therefore,  before  looking  into  the 
origins  of  these  barnacles  relationships  within  the  subfamily  are  discussed. 

Pyrgoma.  in  the  broad  sense,  contains  the  most  highly  evolved  members  of  the  Pyr- 
gomatmae  (Darwm,  1854:  355,  375;  Hiro,  1938:  402).  Baluk  and  Radwahski  (1967b:  691; 
1967c;  486)  revised  Pyrgoma,  dividing  it  into  Pyrgomina  (  =  Megatrema  of  Utinomi,  1967: 
232)  and  Pyrgoma  with  its  subgenera  Nobia  and  Daracia.  We  recognize  somewhat  similar 
groupings,  with  minor  differences  in  the  arrangement  of  species,  but  all  at  the  generic 
level.  The  relationships  of  the  genera  are  indicated  in  Figure  5. 

Pyrgoma  s.  s.,  Nobia,  Savignium,  Hoekia,  and  Pyrgopsella  are  Indo-Pacific  shallow- 
water  pyrgomoids.  Boscia  is  a  cosmopolitan  pyrgomoid,  having  both  shallow  and  deep- 
water  representatives.  The  Indo-Pacific  pyrgomoids  differ  morphologically  from  Boscia  in 
having  highly  modified  opercular  valves  and  in  lacking  paired  fissures  ('sutures')  in  the 
sheath;  they  can  be  derived  readily  from  Indo-Pacific  creusioids  {Cantelliiis,  Creusia, 
Hiroa),  as  will  be  discussed,  but  they  cannot  be  derived  readily  from  Boscia.  We  infer  that 
Boscia  has  had  a  separate  origin— that  is,  that  the  Pyrgomatinae  are  at  the  least  diphyletic. 
In  contrast,  the  Indo-Pacific  pyrgomoids  apparently  are  related  through  two  major  lines 
derived  from  different  creusioid  lineages.  Hence,  we  infer  that  the  pyrgomoid  level  of  or- 
ganization has  been  achieved  at  least  four  times  (Fig.  5). 

Creusia,  in  the  broad  sense,  contains  the  most  generalized  members  of  the  Pyrgoma- 
tinae. Baluk  and  Radwahski  (1967c:  484)  attempted  a  modest  revision  oi  Creusia,  which 
they  divided  into  the  nominate  subgenus  and  a  new  subgenus,  Withersia.  Their  revision 
was  based  mainly  on  fossil  forms,  thereby  considering  only  the  Atlantic  fauna  and  thus 
failed  to  come  to  grips  with  the  Indo-Pacific  Creusia  spimdosa  complex;  and  the  natural 
groupings  that  exist  within  Creusia  were  overlooked.  All  of  the  Atlantic  species,  both  liv- 
ing and  fossil,  form  a  natural  unit  for  which  the  name  Ceratoconcha  is  available.  Cerato- 
concha  has  relatively  unmodified  balanoid  opercular  valves  of  a  characteristic  type  that 
differ  markedly  in  form  from  what  would  be  considered  generalized  balanoid  valves  of 
the  Indo-Pacific  forms  contained  within  our  newly  proposed  genus  Cantellius.  This  in- 
dicates that  the  generalized  or  primitive  creusioids  are  not  closely  related  and  if  the  creu- 
sioids descended  from  a  common   balanoid  stock,  they  did  so  independently  in  the 


147 


CREUSIA 


NOBIA 


PYRGOMA 


BOSCIA 


CANTELLIUS 


CERATOCONCHA 


®Z)V       ^M 


Figure  5.  Diagram  depicting  inferred  phylogenetic  relationships  within  the  Pyrgomatinae.  The  group  or  groups 
from  which  Boscni.  Ceratoconcha.  and  Caniellius  evolved  remain  unknown.  The  solid  lines  radiating  from  the 
orifice  of  the  shell,  shown  in  plan  view,  indicate  relative  position  of  the  sutures  separating  the  compartments;  the 
dotted  lines  in  Bosci'u  indicate  the  position  of  the  pseudoalae.  Dotted  lines  on  the  opercular  plates  indicate  a 
structures  present  in  only  a  few  species  of  that  group. 

Atlantic  and  the  Indo-Pacific.  Thus  we  set  Ceratoconcha  from  the  Atlantic  apart  and  inde- 
pendent from  the  Indo-Pacific  creusioid  genus  Caniellius,  and  consider  the  subfamily  to 
be  triphyletic  (see  Fig.  5). 

Although  Ceratoconcha  has  remained  much  the  same  throughout  its  history,  the 
Indo-Pacific  creusioids  have  undergone  marked  diversification.  There  is  no  fossil  record  to 
document  the  lineages  leading  to  contemporary  forms,  but  among  Recent  representatives 
there  are  sufficient  forms  upon  which  to  draw  inferences.  First,  there  presently  appears  to 
be  no  reason  to  suggest  that  the  Indo-Pacific  creusioids  are  other  than  a  natural  group 
since  they  can  be  derived  readily  from  one  another.  Caniellius  is  the  most  generalized  and 
is  envisaged  as  the  stem  from  which  the  remaining  genera  evolved.  Secondly,  there  are 
apparently  two  major  lineages,  one  stemming  from  Hiroa,  the  other  from  Savignium;— 
that  is,  two  parallel  lines,  each  leading  independently  from  Caniellius  to  pyrgomoid  forms 
(Fig.  5).  Pvrgopsella,  occurring  in  sponges,  appears  to  be  an  ofi'shoot  of  the  Savignium- 
Hoekia  line.  The  modifications  that  ensue  in  each  line  concern  alterations  in  the  form  of 
the  opercular  valves  and  concrescence  of  the  wall  plates,  presumably  better  adapting  the 
barnacles  to  different  host  corals.  Interestingly,  the  most  modified  form,  Hoekia,  has  the 
most  reduced  wall  plate,  aperture,  and  opercular  valves  of  any  pyrgomoid.  It  has  also 
modified  its  nutritional  source,  shifting  from  setose  feeding  to  feeding  directly  on  the  tis- 
sues of  the  host  coral  (Ross  and  Newman,  1969:  255). 

Baluk  and  Radwanski  (1967c:  465)  believed  that  Pyrgopsis  annandalei  Gruvel 
(  =  P\rogopsella  nom.  nov.,  Zullo,  1967),  dredged  from  90m  off  the  Andaman  Islands, 
should  not  be  assigned  to  the  Pyrgomatinae  because  the  basis  is  membranous.  Gruvel 
(1907:  8)  had  three  specimens,  but  the  habitat  and  (or)  actual  relationship  of  the  barnacle 
to  the  substratum  were  unknown.  He  inferred  that  the  membranous  elongate  basis  func- 
tioned as  a  peduncle  or  stalk,  analogous  to  the  fleshy  stalk  of  Xenohalanus,  by  which  the 
animal  attached  to  the  substratum.  Indeed,  Zullo  (1967:  123)  referred  to  Pvrgopsella  as  an 
"unusual  pedunculate  balanid."  Recently,  however.  Resell  (pers.  comm.,  1971)  reported 
finding  a  new  species  of  Pvrgopsella  imbedded  in  a  sponge  from  the  Philippines,  and  we 
believe  that  this  explains  the  peculiar  anatomical  structure  of  the  genus.  The  mem- 
branous stalk  is  not  a  "peduncle"  in  the  sense  used  by  Gruvel,  but  rather  it  is  an  elongate 
basis  comparable  to  and  serving  the  same  function  as  the  elongate  basis  of  other  Pyr- 
gomatinae. In  inhabiting  sponges,  rather  than  a  coral,  the  basis  is  membranous  rather 
than  calcareous,  analogous  to  the  situation  seen  in  Memhranohalanus  also  inhabiting 
sponges.  The  single  plate  making  up  the  wall  and  the  pyrgomoid  valves  suggest  that  Pvr- 
gopsella is  an  off-shoot  of  the  coral-inhabiting  pyrgomatines.  Indeed  the  valves  are  similar 
to  those  of  Savigniunh  and  it  is  from  this  genus  that  we  infer  it  has  evolved. 

In  summary  then,  the  Pvrgomatinae  are  a  diverse  group  of  coral-inhabiting  balanids. 
dominated  by  a  central  group  of  eight  wholly  Indo-Pacific  genera  stemming  from  Can- 


148 


lellius,  and  flanked  by  the  cosmopolitan  genus  Boscia  and  the  Atlantic  genus  Cerato- 
concha.  Cantellius,  Boscia,  and  Ceratoconcha  have  rather  generalized  balanoid  opercular 
valves,  but  there  is  no  indication  that  one  gave  rise  to  the  other.  Rather  it  is  inferred  that 
they  descended  independently  from  balanoid  ancestors,  and  therefore  the  subfamily  is 
considered  triphyletic.  We  can  now  ask  from  which  balanines  these  three  lines  may  have 
evolved. 

There  is  ample  evidence  that  the  Pyrogomatinae  have  been  derived  from  balanines; 
the  rostrum  overlaps  the  laterals,  the  opercular  plates  are  balanoid,  the  labrum  is 
notched,  and  the  intromittant  organ  bears  a  basidorsal  point.  While  the  most  primitive 
living  balanid  {Chelonibia)  has  eight  plates  making  up  the  wall,  it  is  apparently  a  special- 
ized survivor  of  an  ancient  stock  that  presumably  gave  rise  to  the  more  typical  balanines. 
The  vast  majority  of  typical  balanines  have  six  plates  making  up  the  wall,  and  it  has  gen- 
erally been  assumed  that  the  Pyrgomatinae  descended  from  some  six-plated  ancestor 
(Withers,  1929:  564;  1935:  38;  Hiro,  1938:  402;  Zullo,  1967:  127;  Baluk  and  Radwahski, 
1967c:  504).  Withers  (1935:  38)  suggested  that  Balamis  {Balanus)  duvergieri  (Alessandri) 
might  be  such  a  form,  and  Zullo  (1961:  72)  proposed  the  subgeneric  name  Hexacreusia 
for  Balamis  durhami,  a  species  he  thought  also  likely  to  be  such  a  form. 

Balanus  duvergieri,  with  its  tubiferous  wall  and  basis,  appears  to  belong  to  the  sub- 
genus Balanus,  where  Withers  placed  it.  The  wall  of  all  known  pyrgomatines  is  solid; 
while  tubes  may  be  found  in  some  species,  they  are  formed  between  the  sheath  and  the 
internal  ribs  or  between  external  ribs  of  the  wall  and  therefore  are  not  homologous  with 
the  tubes  of  Balanus.  All  generalized  pyrgomatines,  except  Boscia  anglicum,  have  a  solid 
basis.  The  opercular  valves  of  5.  duvergieri  are  generalized,  resembling  those  of  Cantellius 
more  than  those  of  Boscia  and  Ceratoconcha,  but  this  is  no  doubt  simply  because  Cantel- 
lius has  the  most  generalized  valves  of  any  of  the  pyrgomatines.  In  light  of  the  differences 
in  the  wall  between  B.  duvergieri  and  the  Pyrgomatinae,  and  in  light  of  the  evidence  indi- 
cating that  the  Pyrgomatinae  had  a  solid-walled  ancestry,  we  must  agree  with  Baluk  and 
Radwahski  ( 1967c:  504)  that  B.  duvergieri  is  not  an  ancestor  of  the  Pyrgomatinae  as  Withers 
suggested,  nor  is  it  closely  related  to  the  stock  from  which  the  Pyrgomatinae  must  have 
been  derived. 

Balanus  durhami  appears  closer  than  B.  duvergieri  to  the  stem  hne  of  the  Pyrgoma- 
tinae since  it  has  a  solid  wall  and  basis,  and  since  the  opercular  valves  are  superficially 
comparable.  If  5.  durhami  had  but  four  wall  plates  rather  than  six,  would  it  then  belong 
to  the  Pyrgomatinae,  and  if  so,  to  which  of  the  three  major  groups  would  it  be  assigned? 
It  would  belong  to  the  Pyrgomatinae  as  presently  defined,  but  it  is  not  readily  assignable 
to  any  one  of  the  three  existing  divisions.  The  tergum,  with  broad  spur  and  strongly 
developed  depressor  muscle  crests,  and  the  scutum,  with  a  broadly  developed  adductor 
ridge  descending  from  the  occludent  margin,  differ  markedly  from  the  generalized  types 
seen  in  the  subfamily,  so  that  B.  durhami  would  have  to  be  placed  as  a  fourth  and  inde- 
pendent line.  Although  for  different  reasons,  we  agree  with  Baluk  and  Radwahski  ( 1967c: 
504)  that  B.  durhami  is  not  a  surviving  ancestor  of  the  Pyrgomatinae.  Yet  it  is  much  closer 
to  what  must  have  been  the  balanine  stock  from  which  one  or  more  of  the  Pyrgomatinae 
lines  were  derived,  and  it  is  therefore  necessary  to  look  closely  at  the  affinities  of  B.  durhami. 

Zullo  (1961:  75)  stated  that  B.  durhami  resembles  species  of  the  subgenus  Armatoha- 
lanus  but  differs  from  them  in  having  the  anterior  margin  of  cirrus  III  toothed  rather  than 
only  cirrus  IV,  and  he  placed  it  in  a  new  subgenus,  Hexacreusia.^  However,  when  Zullo 
(1963:  590)  described  B.  [Armatobalanus)  nefrens  from  California,  he  noted  that  this  spe- 
cies lacks  hooks  or  spines  on  cirrus  IV,  as  does  B.  {A.)  oryza  Broch  from  the  southwest 
Pacific.  Zullo  (1967:  127)  later  noted  that  Darwin  (1854)  confused  specimens  of  B.  dur- 
hami with  B.  (A.)  allium  from  the  southwest  Pacific.  He  stated  that  such  species  of  Arma- 
tobalanus, as  terebratus  Darwin  are  so  similar  to  B.  durhami  that  it  appears  reasonable  to 
assume  that  the  armatobalanids  were  the  ancestral  stock  from  which  the  coral  barnacles 


'We  wholly  concur  that  Hexacreusia  and  Arniatohalaniis  are  similar,  and  in  fact,  except  for  the  development 
of  the  scutal  adductor  ridge  in  the  former,  there  are  no  diagnostic  difi'erences  between  them.  Rather  than  elevate 
Hexacreusia  to  generic  rank  as  did  Zullo,  et  al  (1972:  72),  we  consider  it  synonymous  with  Annatobaluiuis.  If 
Armatohalanus  were  raised  to  generic  rank,  it  would  be  reasonable  to  consider  //tn^/cref/.v/asubgenerically  distinct. 


149 


Figure  6. 
enhagen. 


Cantellius  pallidus  on   Fungia  fungites  (Linnaeus).   Indo-west   Pacific.  Zoologisk  Museum,  Cop- 


were  derived. 

Darwin  (1854:  282),  Hiro  (1938:  402),  and  Zullo  (1967:  127)  looked  to  Armatoba- 
lanus  as  the  stem  line  from  which  the  Pyrgomatinae  evolved.  Members  of  the  subgenus 
are  found  in  both  the  Atlantic  and  the  Indo-Pacific,  some  occur  exclusively  on  corals,  and 
one  is  known  from  the  late  Miocene  of  the  United  States  (Ross,  1965:  337).  There  is  a  fair 
diversity  of  opercular  valves,  and  in  general  these  bear  a  closer  resemblance  to  those  of 
Cantellius  than  to  those  of  Boscia  or  Ceratoconcha..  If  the  lines  within  the  Pyrgomatinae 
have  in  fact  evolved  three  times,  it  seems  likely  that  at  least  the  Indo-Pacific  Cantellius 
and  its  derivatives  and  perhaps  Boscia  have  an  Armatobalanus  ancestry.  The  affinities  of 
Ceratoconcha  are  still  too  obscure  to  conjecture  (see  Newman  and  Ladd,  in  press). 

SYSTEMATICS 

Descriptions  of  the  54  or  more  species  in  this  subfamily  are  not  included  here,  be- 
cause many  require  redescription  and  adequately  preserved  material  is  unavailable.  To 
obviate  the  problem  of  deciding  on  the  author's  intent  in  relegating  subspecific  or  in- 
frasubspecific  rank  to  a  taxon  (ICZN,  Art.  45),  we  have  blanketly  endorsed  all  known 
nominal  taxa,  and  accordingly  assigned  them  appropriate  rank.  Our  reasons  for  placing  a 
nominal  species  or  genus  in  synonymy  are  given  in  the  remarks  section  under  the  respec- 
tive taxon.  For  each  species,  following  citation  of  the  author  and  date  of  publication,  we 
cite  type  locality  and  host-coral.  A  list  of  species  incertae  sedis  follows  the  systematics  sec- 
tion. 


Family  Balanidae,  Leach,  1817 

Subfamily  Pyrgomatinae  Gray,  1825 

Balanidae  Leach,  1817:  68,  in  part;  Darwin,  1854a:  33,  in  part. 

Pyrgomatidae  Gray,  1825:  102,  in  part;  Reichenbach,  1828:  89,  in  part. 

Bifora  Latreille,  1825:  234,  in  part. 

Pyrgomacea  Menke,  1830:  92,  in  part;  Philippi,  1853:  424,  in  part. 

Sessilia:  Philippi,  1836:  247,  in  part. 

Tetrameridae  Gruvel,  1903:  159,  in  part;  Alessandri.  1922:  226,  in  part. 

Creusiinae  Baluk  and  Radwahski,  1967c:  468. 


150 


Defjfiition.—Shell  of  four  parietal  plates  with  radii  and  alae,  or  totally  concrescent, 
with  or  without  carinal  "pseudoalae"  discernible  in  sheath;  walls  solid  or  tubiferous,  the 
tubes  occurring  in  one  or  more  rows  either  between  the  sheath  and  internal  ribs  or  be- 
tween external  ribs  of  wall;  scutum  and  tergum  either  separate,  cemented,  or  calcified  to- 
gether; basis  membranous  or  calcareous,  when  calcareous  cup-shaped  and  shallow  or 
cylindrical  and  deep;  labrum  with  deeply  incised  notch;  intromittant  organ  with  basi-dor- 
sal  point.  Obligatory  symbionts  or  parasites  primarily  of  scleractinian  corals.  (One  species 
occurs  on  a  hydrocoral,  another  on  a  sponge.)  Type  genus:  Pyrgoma  Leach,  1817. 

Key  to  Genera  of  Pyrgomatinae 

1.  Basis  membranous  (1  sp.) Pyrgopsella 

1 .  Basis  calcareous 2 

2.  Shell  concrescent 3 

2.  Shell  separable  into  4  plates 7 

3.  Scutum  at  least  twice  as  long  as  high 4 

3.  Scutum  as  long  as  high 5 

4.  Tergal  spur  well  developed  (1  sp.) Pyrgoma 

4.  Tergal  spur  rudimentary 6 

5.  Opercular  plates  balanoid,  separable;  tergum 

triangular  (4  spp.) Boscia 

5.  Opercular  plates  modified,  fused  together;  tergum 

quadrate  (6  spp.) Nobia 

6.  Shell  irregular  in  outline;  aperture  minute  (1  sp.) Hoekia  n.  gen. 

6.  Shell  regular  in  outline;  aperture  large  (4  spp.) Savignium 

1 .  Opercular  valves  fused  (3  spp.) Creusia 

7.  Opercular  valves  not  fused 8 

8.  Opercular  valves  highly  modified  (1  sp.) Hiroa  n.  gen. 

8.  Opercular  valves  balanoid 9 

9.  Scutum  with  basal  margin  entire,  depressor  muscle  pit  present, 
but  no  rostral  tooth;  tergum  with  broad  pad  in  area  normally 

occupied  by  depressor  muscle  crests  (16  spp.) Ceratoconcha 

9.  Scutum  with  basal  margin  notched  near  basi-tergal  angle, 
commonly  with  depressor  muscle  pit,  and  a  rostral  tooth; 
tergum  without  broad  pad  (17  spp.) Cantellius  n.  gen. 

Cantellius  n.  gen. 

Definition— V^aW  of  four  plates,  conical  to  flat;  compartments  separated  by  well  de- 
fined radii;  scutum  varies  from  high  triangular  to  transversely  elongated,  and  bearing 
prominent  adductor  ridge  and  lateral  depressor  muscle  depression;  scutum  commonly 
with  rostral  tooth  and  notch  in  basal  margin  near  basitergal  angle;  spur  of  tergum  essen- 
tially confluent  with  scutal  margin,  and  about  Vi  width  of  basal  margin;  crests  for  tergal 
depressor  muscles  feebly  developed  or  wanting. 

Type  species.— Cantellius  transversalis  (Nilsson-Cantell),  1938;  Recent,  Andaman  Isl- 
ands. 

Etymology.— ^Simtd  in  honor  of  Carl  August  Nilsson-Cantell. 

Species  assigned  to  genus: 

Cantellius  acutum  (Hiro),  1938;  398  (syn.:  Creusia  spinulosa  var.  6  subvar.  2  Darwin,  1854:  379);  Palao  Is- 
lands, Caroline  Islands;  on  Acropora  formosa. 

Cantellius  arcuatum  (Hiro),  1938:  395;  Palao  Islands,  Caroline  Islands;  on  Porites  capricornis. 

Cantellius  hreviter^um  (Hiro),  1938:  397;  Palao  Islands,  Caroline  Islands;  on  Acropora  sp. 

Cantellius  et4spinulosum  {Broch).  1931:  118  (syn.:  Creusia  spinulosa  vslt.  1  Darwin,  1854:  377);  Amboina, 
Molucca  Islands;  on  Herpetolitha  sp. 

Cantellius  ^regarea  (Sowerby),  1823  [no  pagination]  (syn.:  Creuisa  spinulosa  var.  3  Darwin,  1854:  378; 
Creusia  spinulosa  pseudoseptima  Kolosvary,  1948:  362;  Creuisa  spinulosa paeudoseptima  [sic]:  Kolosvary,  1951  lb: 
292);  near  Kei  Islands  (5°3rS.,  132°47'E.);  on  Acropora  cvtherea.  type  host  here  designated. 

Cantellius  iwavama  (Hiro),  1938,  p.  393;  Palao  Islands,  Caroline  Islands;  on  Porites  iwayamaensis. 

Cantellius  madreporae  {Bormdaile).  1903:  443  (syn.:  Pvrgoma  madrepoarae  [sic]:  Nilsson-Cantell,  1938:  65); 
Hulule,  Male  Atoll,  Maldive  Islands;  on  Madrepora  sp. 


151 


Figure  7.  Opercular  plates  of  Cantellius.  a-c,  C.  sextus.  after  Hiro.  1938;  d,  e,  C.  arciiatus.  after  Hiro.  1938;  f. 
C.  madreporarae,  after  Borradaile,  1903;  g-i,  C.  euspinulosum.  after  Hiro,  1938;  j-m,  C.  iransversalis.  after  Nil- 
sson-Cantell,  1938;  n-i,  C.  iwavama,  after  Hiro,  1938;  q,  r,  C.  breviiergum.  after  Hiro.  1938;  s,  t.  C.  iredecimus. 
after  Kolosvary,  1947;  u-x,  C.  acutum.  after  Hiro,  1938. 


152 


CanlelUus  octavus  Ross  and  Newman,  n.  sp.  (syn.:  Creusia  spinulosa  var.  8  Darwin,  1854a:  380);  type  local- 
ity, distribution  and  host  coral  not  known. 

Cantellius pallidus  (Broch),  1931:  1 18;  Banda  Sea  (5°32'S.,  132°37'E.);  on  Pocillopora  damkornis.  type  host 
here  designated.  r 


Figure  8.  Opercular  plates  of  CanlelUus.  a-d,  C.  secundus.  after  Hiro,  1938;  e,  C.  septiimis.  after  Darwin,  1854; 
f-h,  C  sumhawae,  after  Hoek,  1913;  i,  j,  C.  gregarea,  after  Nilsson-Cantell,  1938;  k-m,  C.  sepiimus,  after  Hiro, 
1938;  n,  C.  quinius,  after  Darwin,  1854;  o-r,  C  pallidus,  after  Hiro,  1935. 


153 


Caiuelliiis  quinius  Ross  and  Newman,  n.  sp.  (syn.:  Creusia  spinulosa  var.  5  Darwin,  1854a:  379);  type  local- 
ity, distribution  and  host  coral  not  known. 

Canicllius pseudopullidum  (Kolosvarv),  1948:  362:  Pacific  area:  on  Pavona  varians. 

Canicllius  secuncjus  (Broch),  1931:  118  (syn.:  Creusia  spinulosa  var.  2  Darwin,  1854:  378);  offNaira,  Banda 
Islands;  on  Pavonia  sp. 

Canicllius  sepii'mus  (Hiro),  1938:  395  (syn.:  Creusia  spinulosa  var.  7  Darwin,  1854:  380;  Creusia  spinulosa 
duodecima  Koiosvary  and  Wagner,  1941:  9);  Palao  Islands,  Caroline  Islands;  Monlipora  sp. 
cf.  M.  cacutus. 

Canicllius  sextus  (Hiro).  1938  (syn.:  Creusia  spinulosa  var.  6  suhvar.  3  Darwin.  1854:  379);  Palao  Islands, 
Caroline  Islands;  on  Pachvseris  rugosa. 

Cantellius  sumbawae  (Hoek),  1913:  265;  east  of  Dangar  Besar,  Saleh  Bay;  on  Heieropsammia  sp. 

Caniellius  transversalis  (Nilsson-Canteli),  1938:  61  (syn.:  Creusia  spinulosa  var.  6  subvar.  1  Darwin.  1854: 
379);  North  Bay,  Port  Blair,  Andaman  Islands;  on  Madrepora  sp. 

Caniellius  iredecimus  (Koiosvary).  1947:  426;  Island  of  Singapore;  on  Tridacophyllia  lactiua. 

Remarks —Cantellius  is  proposed  for  those  Indo-Pacific  creusoids  with  unfused  oper- 
cular valves  of  which  the  scutum  commonly  possesses  a  notch  in  the  basal  margin  near 
the  basi-tergal  angle,  a  rostral  tooth,  an  adductor  ridge,  and  a  lateral  depressor  muscle  pit. 
The  tergum  has  either  feebly  developed  crests  for  the  depressor  muscles,  or  no  crests. 

In  critically  comparing  the  illustrations  and  brief  description  of  Creusia  spinulosa 
duodecima  Koiosvary  (1941:  9)  with  that  of  Cantellius  septima.  the  authors  find  no 
differences  that  warrant  continued  recognition  of  duodecima.  We  also  find,  for  the  same 
reasons,  that  C.  spinulosa  pseudoseptima  is  synonymous  with  C  gregarea. 

Hiroa  n.  gen. 

Definition— VsldW  of  four  plates,  small,  flat  or  low  conical;  parietal  tubes  present; 
sheath  occupying  whole  inner  wall;  basis  cylindrical  and  deep;  triangular  scutum  high 
and  elongated  transversely;  adductor  ridge  projecting  below  basal  margin  of  valve;  ter- 
gum narrow,  with  spur  about  Vi  or  less  height  of  valve,  lacking  crests  for  depressor  mus- 
cles; overall  height  of  tergum  greater  than  that  of  scutum  and  about  equal  in  bulk  to 
scutum. 

Type  species.— Hiroa  stubbingsi,  new  species. 

Etvmology.—Na.med  in  honor  of  Dr.  Fijio  Hiro  (  =  Huzio  Utinomi),  in  appreciation  of 
his  numerous  studies  on  the  Pyrgomatinae. 

Remarks.— Hiroa  bridges  the  gap  between  Cantellius  and  the  morphologically  ad- 
vanced Indo-Pacific  creusoids  and  pyrgomids.  In  having  a  shell  with  four  distinct  plates,  it 
is  readily  separable  from  Nobia  and  Pvrgoma.  The  bizarre  development  of  the  opercular 
plates,  which  are  separate,  distinguishes  it  from  Cantellius  on  one  hand,  and  from  Creusia 
on  the  other. 

Hiroa  stubblngsi  n.  sp. 

Diagnosis.— ^QCSi\i?,Q  there  is  but  a  single  known  species,  the  diagnosis  is  the  same  as 
that  for  the  genus. 

Material. -^umtrous  specimens  in  Stvlophora  sp.,  type  host;  OUan  Island,  Truk  Islands,  7°14'N,  151°38'E, 
type  locality;  CARMARSEL  Exped.  sample  CRS  811:  25  February  1967;  coral  blasted  from  ba.se  of  seaward 
reef  front  at  8  m;  c"  dating  indicates  age  of  less  than  500  BP. 

/)e.9fr//>//o«.— Specimens  were  entombed  in  coral  so  that  the  external  surfaces  of  the 
wall  could  not  be  observed;  wall  of  four  plates,  flat  or  low  conic;  outline  ovate;  rostrocari- 
nal  diameter  less  than  5  mm,  lateral  diameter  less  than  3  mm;  parietes  non-tubiferous 
and  thickened  marginally,  thinning  toward  aperture;  sutural  surfaces  of  radii  strongly 
denticulate;  sheath  extending  to  basal  margin  of  wall  with  basal  edge  depending  freely. 

Basis  deep  (greater  than  26  mm);  cylindrical;  strongly  ribbed  internally;  non-tubi- 
ferous; gradually  expanding  from  point  of  initial  growth. 

Scutum  high  and  transversely  elongated  (1.7  mm  high  x  1.6  mm  wide);  exterior  sur- 
face sculptured  with  irregular,  high,  growth  ridges;  tergal  margin  about  Vi  length  of  basal; 
occludent  margin  coarsely  toothed;  internal  surface  smooth;  slight  indication  of  adductor 
muscle  depression;  adductor  plate  extends  well  below^  basal  margin  of  valve  proper;  ros- 
tral angle  of  adductor  plate  slightly  produced. 


154 


Figure    9.     Opercular    plates    of    Hiroa 
stubbingsi  n.  gen.,  n.  sp. 


Tergum  T-shaped  or  narrowly  triangular  (2.3  mm  high  x  1.6  mm  wide);  external  sur- 
face ornamented  with  irregular  low  growth  ridges;  external  longitudinal  furrow  deep, 
steep-walled  and  open  throughout  its  length;  internal  surface  smooth,  lacking  crests  for 
depressor  muscles,  deep  depression  present  in  area  bordering  basi-carinal  angle. 

Disposition  of  types— Tho.  holotype  and  two  paratypes  are  deposited  in  the  collections 
of  the  National  Museum  of  Natural  History.  The  remaining  paratypes  are  housed  in  the 
collections  of  Scripps  Institution  of  Oceanography. 

Etymology— ^dimtd  in  honor  of  H.  G.  Stubbings,  long-time  student  of  the  Cirri- 
pedia,  on  the  occasion  of  his  retirement. 

Genus  Creusia  Leach 

Creusia  Leach,  1817:  68.  Genus  without  originally  included  nominal  species;  first  species  assigned  to  genus: 
Creusia  spinulosa  Leach,  1818,  Recent,  type  locality  unknown,  ipso  facto  type  species  by  subsequent  monotypy 
(Leach,  1818:  171). 

Cerusia  (eTTOT  for  Creusia  Leach,  1817):  Ranzani,  1818:  92;  Ranzani.  1820:  56. 

Creusa  (error  for  Creusia  Leach,  1817):  Catlow,  1843:  39. 

Definit ion. —SheW  flat,  ribbed,  compartments  separated  by  narrow  radii;  parietal 
tubes  absent  in  small  species,  rarely  present  in  larger  ones;  scutum  and  tergum  calcified 
together  without  visible  indication  of  line  of  juncture;  adductor  "plate"  commonly  ex- 
tending below  basal  margin  of  valve;  where  plate  extends  below  margin  it  is  produced  as 
basi-rostral  tooth;  no  distinct  lateral  depressor  muscle  depression  on  scutum;  tergal  por- 
tion of  valve  somewhat  quadrate,  occupying  Vi  or  more  of  total  area;  basis  oval,  or  nearly 
circular  in  outline  and  commonly  deep. 

Species  assigned  to  genus: 

Creusia  decima  Ross  and  Newman,  n.  sp.  (syn.:  Creusia  spinulosa  var.  10  Darwin,  1854:  381);  type  locality, 
distribution,  and  host  coral  not  known. 

Creusia  inJicum  (Annanddle),  1924:  64  (syn.:  Creusia  spinulosa  var.  11  Darwin,  1854:  381:  Pyrgoma  indicum 
phase  merulinae  Annandale,  1924:  65;  Pyrgoma  indicum  phase  svmphvlliae  Annandale,  1924:  65;  Creusia  spin- 
ulosa angustiradiaia  Broch,  1931:  118;  Creusia  spinulosa  angustiterga  [sic]:  Nilsson-Cantell,  1938:  63);  Padaw 
Bay,  King-Fsland,  Mergui  Archipelago;  on  Favia  valenciennesii. 

Creusia  spinulosa  Leach,  1818:  171  (syn.:  Creusia  spinulosa  var.  9  Darwin,  1854:  380);  type  locality,  distri- 
bution, and  host  coral  not  known. 


155 


Figure  10.  Creusia.  a,  b,  shell  and  oper- 
cular plate  of  C.  indicum\  c,  opercular 
plate  of  C.  spinulosa:  opercular  plate  of 
C  decima:  all  figures  after  Darwin,  1854. 


Remarks— J\vQ  original  definition  of  Creusia  follows:  "Testa  quadripartita;  oper- 
culum valvis  unipartitis"  (Leach,  1817:  68).  This  was  later  given  by  Leach  (1818:  171)  as: 
"Shell  quadripartite;  parts  equal.  Valves  of  the  operculum  unipartite.  Base  in- 
fundibuliformis."  In  reference  to  Leach's  statement  that  the  opercular  valves  are  fused. 
Gray  (1825:  103)  stated,  in  his  discussion  of  C.  spinulosa,  "Dr.  Leach  describes  the  valves 
of  the  operculum  as  soldered  two  and  two,  but  they  are  not  so  in  the  Museum  specimens." 
Probably  the  opercular  valves  of  the  specimens  in  question,  which  are  not  necessarily  a 
species  of  Creusia  {sensu  stricto),  are  only  cemented  together,  rather  than  calcified  to- 
gether, and  this  would  account  for  the  discrepancy  between  the  two  descriptions. 

Of  the  13  numbered  varieties  and  sub-varieties  of  C.  spinulosa  described  by  Darwin 
(1854a),  three  have  not  been  redescribed  nor  assigned  formal  names.  For  "variety  5"  we 
propose  the  name  Cantellius  quintus:  for  "C  spinulosa  var.  8,"  the  name  Cantellius  oc- 
tavus;  for  "C.  spinulosa  var.  10,"  the  name  Creusia  decima. 

Nilsson-Cantell  (1938:  63)  considered  Annandale's  phase  merulinae  and  phase  sym- 
phylliae  to  be  synonymous  with  Creusia  spinulosa  angustiradiata.  This  taxon  is  a  junior 
subjective  synonym  of  C.  indica,  as  noted  by  Utinomi  (1962:  227),  who  also  followed  Nil- 
sson-Cantell's  suggestion  in  synonymizing  Annandale's  several  "phases." 

Genus  Nobia  Sowerby 

Nobia  Sowerby  (ex  Leach),  1839:  71.  Type  species:  N.  [obia]  grandis  Sowerby,  Recent,  Island  of  Singapore 
(type  locality  here  designated),  by  monotypy. 

Definition. —ShQ\\  flat  or  conical,  ribbed  or  smooth,  composed  of  one  piece  lacking  all 
evidence  of  radii  and  alae;  shell  perched  on  basis;  sheath  applied  directly  to  wall,  extend- 
ing to,  or  nearly  to  basis;  opercular  valves  nearly  of  equal  size  and  fused,  with  line  of  fu- 
sion invisible,  or  visible  either  externally,  internally,  or  both;  scutal  portion  of  valve 
quadrate  to  subquadrate  in  outline;  basis  deep,  cylindrical,  and  either  exserted  or  flush 
with  corallum. 

Species  assigned  to  genus: 

Nobia  conjugcilum  (Darwin).  1854:  364;  Red  Sea;  on  Cvphastraea  chalcidkum. 

Nobia  grandis  Sowerby,  1839:  71;  Singapore;  on  Galaxea  musicalis. 

Nobia  halomitrae  (Kolosvary),  1948:  363;  type  locality  and  distribution  unknown;  on  Halomitra  sp. 

Nobia  kuri(\\oQk).  1913;  259;  near  Kei  Islands  (5°28.4'S.,  132°0.2'E.,);  on  Caryophvllia  sp. 

Nobia  orbicellae  (Hiro),  1934:  367;  Tanabe  Bay,  Japan;  on  Goniopora  sp. 

Nobia  projectum  (Nilsson-Cantell),  1938:  70;  Persian  Gulf;  on  Carvophvlh'a  sp. 

Remarks.—SoweThy^  (1839:  71)  original  definition  of  Nobia  is:  "This  genus  resem- 
bles Pyrgoma,  Auct.  consisting  of  a  conical  paries  supported  upon  a  funnel-shaped  cavity 
in  the  madrepore,  but  diff'ers  in  its  operculum,  which  consists  of  two  valves;  whereas  that 
of  Pyrgoma  has  four." 


156 


Figure  11.     Nobia  grandis  on  Euphyllia  fimhriata  (Spengler);  Warrior  Reef,  Torres  Straits,  Australia;  Museum 
Comparative  Zoology  coral  5685. 


Genus  Pyrgoma  Leach 

Pyrgoma  Leach  (ex  Savigny  MS),  1817:  Genus  without  originally  included  nominal  species;  first  species  as- 
signed to  genus:  Pyrgoma  cancellata  Leach,  1818,  Recent,  Indo-Pacific,  ipso  facto  type  species  by  subsequent 
monotypy  (Leach,  1818,  171;  and  by  subsequent  designation  of  Brooks  and  Ross,  1960:  354). 

Pvrgone  (Qxxox  ior  Pyrgoma  hQ&dn,  1817):  Ferrusac,  1822:  144. 

Pyrgona  (quov  ior  Pyrgoma 'L&&c\\,  1817):  Catlow,  1843:  39. 

Pvrgomum  (error  for  Pyrgoma  Leach,  1817):  Darwin,  1854:  364  (footnote). 

Pyrogoma  (error  for  Pyrgoma  Leach,  1817):  Kolosvary  and  Wagner,  1941:  12;  Kolosvary,  1943:  95. 

Pytgoma  (error  for  Pyrgoma  Leach,  1817):  Johnson,  1963:  95. 

Definition. —Shell  large,  flat  to  sub-conical,  plates  totally  fused;  short  adpressed 
sheath  covers  about  1/5  height  of  inner  wall;  parietal  tubes  present;  triangular  scutum 
high  and  elongated  transversely;  adductor  ridge  projecting  below  basal  margin  of  valve; 
tergum  extremely  narrow,  with  spur  %  to  Va  height  of  valve;  lacking  crests  for  depressor 
muscles;  overall  height  of  tergum  greater  than  that  of  scutum,  but  about  '/>  bulk  of  scu- 
tum. 

Species  assigned  to  genus: 

Pyrgoma  cancellata  Leach,  1818:  171  (syn.:  Pyrgoma  lobata  Gray,  1825:  102;  Pyrgoma  cancellaliim  var.Japo- 
nica  Wehner,  1897:  255);  Sirahama,  Honshu  Island,  Japan,  type  locality  here  designated;  on  Turhinaria  contorta. 

Remarks.— Leach's  (1817:  68)  original  definition  of  Pyrgoma  is:  "Testa  unipartita; 
operculum  valvis  bipartitis."  In  subsequent  publications  Leach  (1818,  1825)  neither  en- 
larged nor  amplified  this  description. 

Pyrgoma  cancellata  is  the  only  species  assigned  to  this  genus.  The  unusual  devel- 
opment of  the  opercular  valves,  especially  the  tergum,  and  the  concrescent  shell,  serve  to 
distinguish  it  from  those  species  previously  referred  to  Pyrgoma. 


157 


Figure  12.  Opercular  plates  of  Nobia.  a-c,  N.  ^randis.  after  Darwin,  1854;  d,  e.  .V.  conjugaium.  after  Darwin. 
1854;  f,  g,  N.  kuri,  after  Hoek.  1913;  h.  ,V.  halomiirae.  after  Kolosvary.  1948;  i.  j.  ,V.  projecium.  after  Nilsson- 
Cantell,  1938;  k,  1,  N.  orbkellae,  after  Hiro,  1935. 


158 


Figure  13.     Pyrgoma  cancellata  Leach  on  Dendrophvllia  micranthus  grandis  Crossland;  Great  Barrier  Reef,  Aus- 
tralia; Zoologisk  Museum,  Copenhagen. 


Figure  14.     Opercular  plates  of  Pyrgoma  cancellata,  after  Hiro,  1935. 


Genus  Savignium  Leach 

Savignium  Leach,  1825  (not  Sowerby,  1823,  nomen  nudum):  210.  Genus  without  originally  included  nomi- 
nal species;  first  species  assigned  to  genus:  D.  [aracia]  linnaei  Gray,  1825  [  =  Savignium  crenaium  Sowerby.  1823], 
Recent,  Island  of  Singapore  (type  locality  here  designated),  ipso  facto  type  species  by  subsequent  monotypy 
(Gray,  1825:  102). 

Daracia  Gray,  1825:  102.  Type  species:  D.  [aracia]  linnaei  [  =  Savignium  linnaei  =  Savignium  crenatum  So- 
werby, 1823],  Recent,  Philippine  Archipelago,  by  monotypy. 

Z)orada  (error  for  Z)arac/a  Gray,  1825):  Weltner,  1897:  278. 

Definition —Shell  totally  fused,  flat,  oval  in  outline;  lower  margin  of  sheath  free,  ex- 
tending nearly  to  basal  edge  of  wall;  opercular  valves  separate,  cemented,  or  fused  to- 


159 


Figure  15. 
1868. 


Savignium  crenatum  on  Merulina  ampliata  Ellis  and  Solander;  Singapore;  American  Museum  coral 


gether;  scutum  transversely  elongated,  its  overall  length  exceeding  that  of  tergum;  tergum 
variable,  commonly  lacking  definitive  spur  and  lacking  crests  for  depressor  muscles;  scu- 
tum comprising  bulk  of  operculum;  basis  commonly  deep  and  cylindrical. 
Species  assigned  to  genus: 

Savignium  crenatum  Sowerby,  1823,  no  pagination  (syn.:  Pyrgoma  crenatum  phase  tridacophvlliae  Annan- 
dale.  1924:  66;  Pyrgoma  crenatiformis  Kolosvary,  1951:  287);  Singapore,  type  locality  here  designated;  on  Tri- 
dacophvllia  lactuca. 

Savignium  dcnialum  Darwin.  1854:  369;  Red  Sea;  on  Meandrina  spongiosa. 

Savignium  elongatum  Hiro.  1931;  154;  Sirahama.  Honshu  Island.  Japan;  on  Madrepora  sp. 

Savignium  milleporae  Darwin.  1854:  367  (syn.:  Pyrgoma  millepora  [sic]:  Nilsson-Cantell.  1938:  65;  Pyrgoma 
milleporae  forma  typica  Kolosvary,  1950:  292;  Pyrgoma  milleporae  CoTma  snelliusi  Kolosvary,  1950:  292);  Min- 
doro  Island.  Philippine  Archipelago;  on  Millepora  complanata. 

Remarks— Aiter  Leach  (1817,  1818)  published  his  first  two  studies  on  the  Cirripedia 
he  subdivided  Pyrgoma  and  proposed  the  genera  Savignium,  Megatrema,  and  Adna.  Al- 
though he  did  not  publish  these  names  until  1825,  he  did  leave  labeled  specimens  in  the 
British  Museum  (Natural  History)  collections  (see  Sowerby,  1823;  Gray,  1825:  107).  So- 
werby (1823)  found  ". . .  upon  examining  the  collection  of  Cirripedes,  in  the  British  Mu- 
seum, as  it  now  remains  arranged  by  Leach  himself,  that  since  the  publication  of  the 
'Supplement  to  the  Encyclopedia  Britannica,'  where  the  characters  of  the  genus  [Pyr- 
goma] first  appear  in  print,  he  [Leach]  had  divided  into  four;  upon  what  grounds  we  must 
acknowledge  ourselves  entirely  ignorant,  except  it  be  from  some  diff"erences  in  the  form  of 
the  shell,  and  the  valves  of  the  operculum  .  .  .  We  do  not  consider  .  .  .  these  four  genera  .  .  . 
sufficiently  distinct  to  constitute  several  genera  .  .  .  wherefore  we  still  include  all  above 
enumerated  [Megatrema,  Savignium,  and  Adna]  under  the  denomination  of  Pyrgoma^ 

Sowerby  is  not  considered  the  author  of  Megatrema,  Savignium,  or  Adna,  although 
his  publication  has  priority,  because,  "A  name  first  published  as  a  synonym  is  not  thereby 


160 


Figure  16.  Savignium  milleporae  on  Mil- 
lepora  sp.;  Heron  Island,  Great  Barrier 
Reef,  Australia. 


made  available  unless  prior  to  1961  it  has  been  treated  as  an  available  name  with  its  origi- 
nal date  and  authorship,  and  either  adopted  as  the  name  of  a  taxon  or  used  as  a  senior 
homonym"  (Article  1 1  (d),  ICZN).  Sowerby's  use  of  the  specific  names,  Savignium  crena- 
tum  and  Adna  anglica,  suggested  by  Leach,  on  the  other  hand,  entitles  him  to  the  author- 
ship of  these. 

Leach's  second  synopsis  of  the  Cirripedia,  published  in  July  of  1825,  contained  brief 
descriptions  of  Savignium,  Megatrema,  and  Adna.  At  the  same  time  Leach  was  working 
on  his  manuscript.  Gray  (1825)  was  also  preparing  a  synopsis  of  the  Cirripedia,  which  was 
pubhshed  in  the  August  issue  of  the  Annals  of  Philosophv.  In  his  synopsis.  Gray  (1825: 
102)  described  the  genus  Daracia  as  follows:  ""Daracia,  Gray,  Savignium,  Leach,  without 
character.  Valves  of  the  body  of  the  shell,  four,  soldered  together."  This  description  com- 
pares favorably  with  Leach's  abbreviated  description  of  Savignium:  "Testa  indivisa:  basis 
immersa,  valvae  indivisae"  (1825:  210).  The  only  species  mentioned  by  Gray  in  con- 
nection with  the  definition  of  Daracia  is  linnaei,  which  is  not  described.  In  the  discussion 
of  Pyrgoma,  Sowerby  (1823)  made  reference  to  Savignium  crenatum,  which  he  attributed 
to  the  authorship  of  Leach.  That  Gray  referred  to  the  same  specimens  as  did  Sowerby, 
who  figured  them,  seems  probable  at  this  time,  and  Gray  more  than  likely  based  his  con- 
cept of  D.  linnaei  on  these  specimens.  Therefore,  we  believe  that  Gray's  D.  linnaei  is  ac- 
tually a  junior  objective  synonym  of  Savignium  crenatum.  It  should  also  be  noted  that 
Gray,  proposed  Daracia  as  a  replacement  name  for  Savignium  (see  Gray  1825:  102,  foot- 
note). 

Based  on  the  general  aspects  of  barnacles  overgrown  by  a  milleporine,  Darwin  (1854: 
366)  suspected  Chenu's  (1843)  Creusia  madreporarum  to  be  synonymous  with  his  Pyr- 
goma  milleporae.  Chenu,  questionably,  ascribed  this  taxon  to  the  authorship  of  Leach, 
and  his  illustration  speaks  favorably  of  its  being  the  same  as  Darwin's  taxon.  So  far  as  we 
have  been  able  to  determine,  the  only  pyrgomatid  reported  from  a  milleporine  is  P.  mille- 
porae. In  the  interests  of  stability,  although  recognizing  that  C  madreporarum  has  prior- 
ity, Darwin's  name  is  used  here. 

Two  forms  of  Pyrgoma  milleporae  were  designated  by  Kolosvary  (1951:  292),  typica 
(  =  P.  milleporae  milleporae)  and  snelliusi.  These  are  not  recognized  here  because  the  mor- 
phological variations  recorded  fall  within  the  limits  of  variation  assumed  to  correlate  with 
different  infrageneric  milleporine  associations. 

In  the  coral  collections  of  the  American  Museum  there  are  two  large  specimens  of 
Merulina  ampliata  (cat.  no.  1868  and  3214)  from  the  Island  of  Singapore  infested  with 
pyrgomatids.  Our  study  of  these  indicated  that  they  are  conspecific  with  Pyrgoma  crena- 
tum. In  1951  Kolosvary  described  P.  crenatiformis  from  the  coral  Merulina  ampliata,  col- 


161 


ra 


Figure  17.  Opercular  plates  of  Savignium.  a-d,  S.  crenatum.  after  Hiro,  1935;  e-g,  5'.  milleporae.  after  Darwin, 
1854;  h-k,  S.  denialum.  after  Hiro.  1935  and  1938;  1-n.  S.  elonganinu  after  Hiro.  1938. 

lected  in  the  vicinity  of  the  Island  of  Singapore.  Comparison  of  our  specimens  with 
Kolosvary's  illustrations  and  brief  description  does  not  reveal  differences  that  enable  one 
to  separate  these  two  species. 

Hoekia  n.  gen. 

Definit ion. —SheW  totally  concrescent,  irregularly  lobate  in  outline,  and  exhibiting  no 
definitive  peripheral  shape;  region  surrounding  minute  ovate  orifice  elevated  above  exter- 
nally flat  or  undulatory  surface  of  shell;  sheath  short,  adpressed,  basal  margin  not  de- 
pending freely;  irregularly  scattered  wall  tubes  occur  at  varying  distances  from  shell 


162 


Figure  18.  Hoekia  monticulariae.  Left,  on  Hydnophora  exesa  (Pallas);  Mortensen  Java-South  Africa  Expedition 
1929-30;  Station  44,  Flat  Island,  Mauritius;  Zoologisk  Museum,  Copenhagen.  Right,  internal  view  of  shell  from 
Hydnophora  exesa;  Singapore;  American  Museum  coral  1883. 

margin;  operculum  minute;  scutum  and  tergum  fused  without  evidence  of  suture,  form- 
ing elongate  valve  with  broad  occludent  ledge;  tergal  end  of  valve  lacking  spur. 

Etymology —This  taxon  honors  the  late  Dutch  cirripedologist  Paulus  Peronius  Cato 
Hoek(1851-1914). 

Type  species.— Pyrgoma  monticulariae  Gray,  1831,  Recent,  Island  of  Singapore. 

Species  assigned  to  genus: 

Hoekia  monticulariae  (Gray),  1831:  6;  Singapore;  on  Hydnophora  exesa. 

Remarks.— ThQ  gross  differences  between  the  shell  and  opercular  valves  of  mon- 
ticulariae and  those  of  other  Pyrgomatinae  are  of  sufficient  magnitude  to  warrant  its  sepa- 
ration as  a  distinct  genus  (see  Ross  and  Newman,  1969). 

Although  Baluk  and  Radwahski  (1967b:  487)  resurrected  the  name  Daracia  to  in- 
clude Pyrgoma  monticulariae  and  P.  elongatum,  it  is  readily  apparent  that  not  only  was 
Daracia  proposed  as  a  replacement  name  for  Savignium  (see  Gray,  1825:  102,  footnote), 
but  also  the  type  species  D.  linnaei  appears  to  be  a  junior  objective  synonym  of  5".  crena- 
tum.  Consequently,  we  feel  justified  in  proposing  a  new  taxon. 

Hoekia  is  allied  morphologically  to  Savignium  in  that  the  fused  opercular  valves 
show  some  affinity  to  those  of  S.  crenatum,  S.  dentatum,  and  less  so  to  S.  milleporae,  as 
pointed  out  by  Darwin  (1854a:  374).  But,  in  these  species  the  valves  are  separate  or  only 
cemented  together. 


Figure  19.     Opercular  plate  of  Hoekia  monticulariae: 
top,  after  Darwin,  1854;  bottom,  after  Hiro,  1935. 


163 


Figure  20.  Shell  and  opercular  plates  of 
Pvr^opseUa  annandalei.  redrawn  after 
Gruvel,  1907. 

Aside  from  the  fact  that  H.  monticulariae  is  the  only  known  wholly  parasitic  balanid, 
the  trophi  of  this  species  depart  radically  from  those  of  other  pyrgomatines  (Ross  and 
Newman.  1969:  255).  Apparently,  critical  study  of  the  mouth  field  should  provide  addi- 
tional and  independent  criteria  for  recognition  of  pyrgomatine  generic  groups.  Our  pre- 
liminary studies  of  species  having  morphologically  primitive  shells  indicate  that  the 
trophi  of  these  depart  little  from  that  of  many  primitive  balanids  (see  Broch,  1924,  fig. 
10). 

Genus  Pyrgopsella  Zullo 

Pyrgopsis  Gruvel,  1907:  8.  Type  species:  Pyrgopsis  annandalei  Gruvel,  Recent,  Andaman  Islands,  by  mono- 
typy. 

Pyrgopsella  Zullo,  1967:  109  (substitute  name  for  Pyrgopsis  Gruvel,  1907,  not  Rochebrune,  1884). 

Definition.— V^dW  subconical,  rostro-carinally  elongate,  smooth,  composed  of  numer- 
ous calcareous  rods  contained  in  a  chitinous  envelope  continuous  with  basis;  basis  elon- 
gate and  membranous;  opercular  plates  separate,  well  calcified,  scutum  transversely  elong- 
ated; tergum  triangular  with  short  irregular  spur;  living  in  sponges. 

Species  assigned  to  genus: 
Pyrgopsella  annandalei  (Gr\i\Q\),  1907:  8:  Andaman  Islands;  host  unknown. 

Remarks— T\\Q  remarkable  feature  in  Pyrgopsella  is  the  membranous  basis  that  Gru- 
vel (1907:  9)  thought  served  as  a  peduncle.  Rosell  (pers.  comm.)  recently  discovered  a 
new  species  of  Pyrgopsella  living  in  a  sponge  in  the  Philippines,  and  from  this  it  is  clear 
that  the  function  of  the  elongate  membranous  basis  is  comparable  to  that  of  the  elongate 
calcareous  basis  of  the  other  pyrgomatines,  and  that  being  membranous  is  simply  a  sec- 
ondary adaptation  to  living  in  sponges  as  opposed  to  corals. 

Utinomi  (1943:  16)  studied  the  post  larval  settlement  stages  in  Creusia  indicum,  and 
found  the  basis  to  be  initially  cup-like  and  wholly  membranous.  It  is  evident  that  calcifi- 
cation of  the  basis  is  delayed,  at  least  in  C.  indicum,  and  hence  it  is  not  difficult  to  envis- 
age that  in  Pyrgopsella  ontogenetic  suppression  of  calcium  deposition  would  result  in  a 
membranous  basis. 

The  general  shape  of  the  shell  and  the  opercular  plates  of  Pyrgopsella  are  similar  to 
those  found  in  Savignium  dentatum.  From  the  morphology  of  the  hard  parts  it  is  apparent 
that  Pyrgopsella  was  derived  from  Savignium. 

Genus  Boscia  Ferussac 

Boscia  Ferussac.  1822:  145.  Type  species:  Balanus  madreporarum  Bosc,  1812  [  =  Boscia  madreporamm 
(Bosc)],  Recent,  Caribbean-western  Atlantic,  by  monotypy. 

Megatrema  Leach,  1825  (not  Sowerby,  1823,  nomen  nudum):  210.  Genus  without  originally  included  nomi- 
nal species;  first  species  assigned  to  genus:  M.  [egalrema]  slokesii  Gray,  1825  [  =  Boscia  madreporarum  (Bosc), 
1812],  Recent,  Caribbean-western  Atlantic:  ipso  facto  type  species  by  subsequent  monotypy  (Gray,  1825:  102), 
and  subsequent  designation  of  Philippi  ( 1853:  424). 

Adna  Leach,  1825  (not  Sowerby,  1823,  nomen  nudum):  210.  Genus  without  originally  included  nominal  spe- 


164 


'■*?-l*^,*  .lit 


r^^ 


■iMIiiiMi 


Figure  21.     Boscia  madreporarum  on  Agahcia  agahcites  (Linnaeus);  Dry  Rocks,  off  Key  West,  Florida. 

cies;  first  species  assigned  to  genus:  M.  [egatrema]  (A.  [dna])  anglica  Gray,  1825  \  =  Boscia  cmglicum  Sowerby, 
1823],  Recent,  coast  of  Devonshire,  England;  ipso  facto  type  species  by  subsequent  designation  of  Philippi  ( 1853: 
424). 

Pyrgominia  Baluk  and  Radwahski,  1967b:  691.  Type  species:  Pvrgominia  seguenzai  Baiuk  and  Radwanski, 
1967  [  =  Boscia  seguenzai  Baluk  and  Radwanski],  by  original  designation.  Pliocene,  Island  of  Crete,  Greece. 

Definition.— SheW  conical  in  juveniles  and  commonly  flat  or  low  conical  in  later 
stages;  shell  plates  totally  fused  externally;  pseudo-alae  may  be  present;  sheath  adpressed 
and  covering  %  to  entire  inner  wall;  opercular  valves  typically  balanoid;  terga  lacking 
depressor  muscle  crests;  basis  cup-shaped  or  sub-cylindrical,  exserted  or  flush  with  coral- 
lum. 

Species  assigned  to  genus: 

Boscia  angliciim  Sowerby,  1823  [no  pagination]  (syn.:  Pvrgoma  sulcatum  Philippi,  1836,  pi.  12,  fig.  24;  Pvr- 
goma  undaium  Michelotti,  1839:  140-141);  coast  of  Devonshire,  England;  on  Carvophyllia  smilhii. 

Boscia  madreporarum  {Bo^c),  1812:  66  (.syn.:  Creusiaboscii  DeBlainville,  1824:  378;  Pvrgoma  stokesii  Gray, 
1825:  103;  Creusia  decorata  Chenu,  1843  [no  pagination];  Pvrgoma  stockesi  [sic]:  Kruger,  1940:  382);  "Ame- 
rique"  [  =  Caribbean  western-Atlantic];  on  Agaricia  agaricites. 

Boscia  oulastreae  (Utinomi),  1962:  83;  Nomosaki,  Kyushu  Island,  Japan;  on  Oulastrea  crispata. 

Boscia  seguenzai  (Baluk  and  Radwahski),  1967b:  691;  Gournes,  Island  of  Crete,  Greece;  Pliocene. 

Remarks— FeTuss'dc''s  (1822:  14)  original  description  of  Boscia  follows:  "Test  univalve 
en  cones  tres-surbaisse,  a  parois  tubuleuses;  articule  avec  la  base.  Celle-ci,  plus  grande,  en- 
forme  de  godet  ou  de  cupule.  " 

in  the  year  following  the  publication  o{ Boscia,  Sowerby  (1823)  published  two  manu- 
script names  of  Leach:  Megatrema  and  Adna.  When  Sowerby  described  Megatrema  he 
failed  to  mention  any  nominal  species.  Subsequently,  stokesii  was  assigned  to  the  genus 
(Gray,  1825:  103).  However,  it  appears  that  Megatrema  stokesii  is  a  junior  subjective  syn- 


165 


Figure  22.  Boscia  madreporarum  ( =  Pvrgoma  stokesii).  Top,  slab  with  several  shells  and  opercular  valves 
mounted  and  identified  by  Darwin:  bottom  right,  external  view  of  shell  shown  at  top  center  of  slab;  bottom  left, 
internal  view  of  shell  shown  at  top  left  of  slab.  British  Museum  (Nat.  Hist.)  1962.  12.7.1. 


onym  of  Boscia  madreporarum,  because  there  are  no  differentiating  morphological  fea- 
tures, and  because  it  occurs  on  the  same  host  coral,  Agaricia  agaricites  (Linnaeus),  in  the 
same  geographical  region. 

Adna  was  described  by  Sowerby  (1823)  as  a  subjective  synonym  of  Pvrgoma.  How- 
ever, Sowerby  is  regarded  as  the  author  only  of  the  specific  name  anglica  (see  Article 
1 1(d),  ICZN).  Leach,  who  originally  proposed  Adna,  did  not  publish  the  name  until  1825. 
At  that  time  no  nominal  species  was  assigned  to  the  genus.  Gray  (1825:  103)  included 
only  one  species,  Adna  anglica. 

The  manner  is  which  Gray  (1825:  103)  cited  the  taxon  Adna  suggests  that  it  was  to  be 
recognized  as  a  subgenus  of  Megatrema.  He  did  not  state  why  this,  rather  than  a  generic 
assignment  was  made,  nor  did  Darwin  (1854:  360)  who  also  cited  Adna  as  a  subgenus. 

Leach's  original  definition  of  Megatrema  is  "Testa  indivisa:  basis  immersa,  valvae 
Balani,"  while  that  of  Adna  is:  "Testa  indivisa:  basis  exserta,  valvae  Balani"  (1825:  210). 
The  only  difference  between  the  two,  as  proposed  by  Leach,  is  in  the  basis,  which  in  Adna 
is  not  flush  with  the  surface  of  the  corallum. 

Of  the  pyrgomatids  known  to  Darwin  (1854a:  355),  only  Pyrgoma  stokesii  {  =  Boscia 
madreporarum)  and  P.  anglicum  ( =5.  anglicum),  ".  .  .  have  some  claims  to  be  generically 
separated  from  the  other  species  of  Pyrgoma  .  .  ."  This  opinion  was  based  on  the  sim- 
ilarity of  the  operculum,  and  the  conical  shells  which  internally  exhibit  carinal  pseu- 
doalae.  The  authors  have  adopted  Darwin's  suggestion  and  maintain  these  two  species,  in 
addition  to  Boscia  oulastreae  and  B.  seguenzai  in  a  distinct  genus. 

What  have  been  interpreted  as  carinal  sutures  are  a  pair  of  fines  where  the  arthrodial 


166 


Figure  23.     Opercular  plates  of  Boscia.  a,  b,  B.  anglicum,  after  Darwin,  1854;  c-f,  B.  oulastreae,  after  Utinomi, 
1962;  g,  h,  B.  madreporarum. 


membranes  of  the  scuta  and  terga  attach  to  the  sheath.  As  the  operculum  is  carried  ba- 
sally  with  growth  of  the  sheath,  the  Hnes  remain  marking  the  points  of  earher  attachment. 
These  Hnes  could  represent  vestiges  of  the  suture,  but  they  appear  only  in  the  sheath  and 
not  the  wall.  When  the  shell  is  ground  transversely,  the  area  beneath  the  lines  has  a 
folded  appearance;  the  overlapping  portion  being  termed  a  pseudoala. 

Genus  Ceratoconcha  Kramberger-Gorjanovic 

Ceratoconcha  Kramberger-Gorjanovic,  1889:  50.  Type  species:  Ceratoconcha  costata  Kramberger-Gorjano- 
vic [  =  Creusia  krambergeri  nom.  nov.  =  Ceratoconcha  krambergi  (Baluk  and  Radwanski),  1967:  145],  Miocene, 
Yugoslavia,  by  monotypy. 

Paracreusia  Abel,  1927:  101.  Type  species:  Paracreusia  trolli  Abel  [  =  Ceratoconcha  trolli  (Abel)],  Miocene, 
Italy,  by  monotypy. 

Andromacheia  Kolosvary,  1949:  4.  Type  species:  Andromacheia  noszkvi  Kolosvary  [  =  Ceratoconcha  noszkvi 
(Kolosvary)],  Upper  Miocene,  southern  Hungary,  by  monotypy. 

Withersia  Baluk  and  Radwanski,  1967c:  485.  Type  species:  Creusia  barbadensis  Withers,  Pleistocene,  Bar- 
bados by  original  designation. 

Definition. -~?i\\Q\\  of  four  compartments  separated  by  radii  and  alae,  the  latter  trend- 
ing toward  reduction  in  size;  shell  ribbed,  ranging  from  conical  to  nearly  flat;  sheath  ap- 
proximately Vi  height  of  wall,  with  basal  margin  depending  freely;  well  developed  ribs 
may  occur  on  inner  surface  of  shell  and  extend  from  base  to  sheath;  opercular  valves  typi- 
cally balanoid;  rostral  tooth  of  scutum  either  inconspicuous  or  wanting;  well  developed 
lateral  depressor  muscle  pit  present;  tergum  commonly  bears  a  prominent  ridge  or  plate 
on  carinal  segment,  rather  than  depressor  muscle  crests;  basis  commonly  deep  and  cy- 
lindrical. 

Species  assigned  to  genus: 

^Ceratoconcha  barbadensis  (Withers),  1926:  2  (syn.:  Creusia  barndensis  [sic]:  Nilsson-Cantell,  1938:  63); 
Barbados,  West  Indies;  Pleistocene. 

^Ceratoconcha  costata  (Sequenza),  1876,  p.  316  (syn:  Creusia  costata  elargata  (Sequenza),  1876:  317; 
Creusia  moravica  Prochazka,  1893:  20;  Creusia  spinulosa  forma,  praespinulosa  Kolosvary,  1949:  1,  fig.  5  only; 
Creusia  spinulosa  forma  kojumdgievae  Kolosvary,  1962:  86);  Messina,  Italy;  Pliocene  (Astian). 

^Ceratoconcha  darwiniana  {Proishdzkd).  1893:  23;  Leibnitz,  Australia;  Miocene. 

\ Ceratoconcha  diploconus  (Seguenzsi),  1876:  322;  Messina,  Italy;  Pliocene  (Astian). 

Ceratoconcha  domingensis  (Des  Moulins),  1867:  307;  Port-au-Prince,  Haiti;  on  Porites  astreoides. 

Ceratoconcha  floridanum  (Pilsbry),  1931:  81;  Gulf  of  Mexici>;  on  Maeandra  sp.  cf.  M.  areolata. 

1; Ceratoconcha  krambergeri  (Baluk  and  Radwaiiski),  1967a:  145  (see  Kramberger-Gorjanovic,  1889:  50); 
Podsused,  Yugoslavia;  Miocene. 


167 


Figure  24.  Ceratoconcha  fioridanum  on 
MvcetophvUia  lamarckana  Milne-Ed- 
wards and  Haime;  Recent,  Florida  Keys. 


^Ceratoconcha  miocaenica  (?xoc\\dLzk3i),  1893:  22;  Wollersdorf,  Austria:  Miocene. 

^Ceratoconcha  noszkvi  (V^o\o^\iL'[y),  1949:  4;  Magyarszek,  Hungary;  Miocene. 

^Ceratoconcha  prefioridana  (Brooks  and  Ross),  I960:  355  (syn.:  Creusia  neogenica  weisbord,  1972:  60); 
Florida,  U.S.A.;  Pliocene;  on  Manicina  mayori. 

Ceratoconcha  quaria  (Kolosvary),  1947:  426  (syn.:  Creusia  spinulosa  var.  4  Darwin,  1854:  378);  West  Indies; 
West  Indies;  on  Colpophvllia  nutans. 

^Ceratoconcha  rangi  rangi  (Des  Moulins),  1867:  302  (syn.:  Pyrgoma  miilticostatum  Seguenza.  1873:  319; 
Creusia  fuchsi  Prochazka,  1893:  18;  Creusia  spinulosa  forma,  caldangiae  Kolosvary.  1949:  1;  Creusia  spinulosa 
formd praespinulosa  Kolosvary,  1949:  I,  figs.  2-3  only);  Bazas,  France;  Miocene  (Aquitanian). 

j Ceratoconcha  rangi  latum  (Seguenza).  1876:  321;  Rometta,  Italy;  Miocene  (Tortonian). 

\ Ceratoconcha  sanctacrucensis  Baluk  and  Radwaiiski,  1967c:  468;  Korytnica,  Poland;  Miocene  (Torton- 
ian); on  Tarhellastraea  reussiana 

^Ceratoconcha  sturi  (?xoc\\azka),  1893:  15;  Sudic,  Czechoslovakia;  Miocene. 

^Ceratoconcha  trolli  (Abd).  1927:  101:  Voslau,  Austria;  Miocene:  on  Siderastraea  crenulata. 

Remarks— \n  view  of  the  allocation  of  the  coral-inhabiting  barnacles  to  different  gen- 
era, the  specific  name  Ceratoconcha  costata,  proposed  by  Kramberger-Gorjanovic  (1889: 
50),  becomes  a  junior  homonym  of^  Creusia  costatuni  (Sequenza,  1876:  316).  As  a  replace- 
ment for  this  preoccupied  name,  Baluk  and  Radwahski  (1967a:  145)  proposed  Creusia 
kramhergeri. 

The  validity  of  Paracreusia  has  long  been  questioned  (Withers,  1929:  565:  Hiro, 
1938:  414;  Kriiger,  1940:  452),  because  no  apparent  differences  allowing  separation  from 
Ceratoconcha  were  noted  by  Abel  (1927,  1928),  or  subsequent  workers.  Baluk  and  Rad- 
wahski (1967c:  482)  suggested  merging  Paracreusia,  a  proposal  we  have  adopted  here. 

Kolosvary  (1949:  4)  proposed  Andromacheia  on  the  basis  of  one  poorly  preserved 
specimen  with  visible  and  irregularly  developed  squamate  compartments.  The  shell  sur- 
face was  said  to  bear  three  rows  of  scales.  These  are  probably  the  result  of  weathering  in- 
asmuch as  the  same  feature  was  noted  in  Ceratoconcha  cladangiae  (Kolosvary,  1949:  2); 
therefore  they  are  not  considered  to  be  of  major  taxonomic  significance.  The  poor  demar- 
cation of  the  parietal  plates  and  poor  development  of  radii  and  alae,  as  noted  by  Kolos- 
vary, is  often  encountered  in  fossil  material.  Baluk  and  Radwahski  (1967c:  476) 
questioned  "^hQihtx  Andromacheia  was  even  a  barnacle. 

Many  fossil  Pyrgomatines  have  been  described  from  specimens  lacking  morphologi- 
cally important  details  (Baluk  and  Radwariski,  1967c:  482).  Many  of  these  are  based  on 
unique  specimens,  and  many  of  them  have  not  been  reported  or  described  since  their  in- 


168 


Figure  25.  Opercular  plates  of  Cerato- 
concha.  a,  b,  C.  quaria,  after  Darwin, 
1854;  c-e,  C.  preflondamim,  after  Brooks 
and  Ross.  1960. 


itial  publication.  Over  half  of  the  species  assigned  to  Ceratoconcha  were  originally  based 
on  incomplete  specimens  and  still  are  known  to  us  solely  on  the  basis  of  an  abbreviated, 
incomplete  description.  Invariably  these  descriptions  omit  the  morphology  of  the  oper- 
culum, which  is  perhaps  the  most  diagnostic  feature  of  this  genus.  Therefore,  the  assign- 
ment of  many  of  the  species  must  remain  tentative  until  well  preserved  and  more 
complete  material  becomes  available. 

Baluk  and  Radwahski  (1967c:  485)  proposed  the  subgenus  Withersia  for  two  species, 
one  of  which,  Creusia  barbadensis,  is  here  referred  to  Ceratoconcha,  and  the  other,  C  ou- 
lastreae,  to  Boscia.  The  reason  for  proposing  Withersia  was  that  the  radial  sutures  are  "in- 
distinct or  even  disappearing."  In  barbadensis  sutures  are  present,  but  poorly  discernible 
or  obscure  largely  due  to  secondary  calcification,  whereas  in  oulastreae  radial  sutures  are 
never  present  in  the  adult  stage. 

Incertae  Sedis 

The  following  taxa  cannot  be  assigned  to  any  of  the  genera  defined  herein:  Creusia 
childreni  Gray,  1825;  Balanus  duploconus  Lamarck,  1818;  Megatrema  semicostata  So- 
werby,  1839;  Pyrgoma  stellata  Chenu,  1843;  Pvrgoma  spongiarum  Chenu,  1843;  Pyrgoma 
corymbosa  "Valenciennes"  Chenu,  1843;  Creusia  radiata  Chenu,  1843;  Creusia  multi- 
striata  Chenu,  1843;  Creusia  madreporarum  "Leach?,"  Chenu,  1843;  Creusia  striata 
Chenu,  1843;  Pyrgoma  undata  Michelotti,  1839. 

Darwin  (1854:  365,  footnote)  noted  that  Balanus  duploconus  Lamarck  may  be  synon- 
ymous with  Nobia  grandis  Sowerby.  Lamarck's  (1818:  394)  description,  "R  testae  parte 
suprema  univalvi,  indivisa,  convexa;  inferiore  turbinata,  non  clausa;  apertura  elliptica," 
may  also  apply  to  other  species  here  included  in  the  genera  Nobia,  Pvrgoma,  or  Boscia. 
The  uncertainty  that  surrounds  the  nature  of  B.  duploconus,  which  Lamy  and  Andre 
(1932)  failed  to  clarify,  stems  from  the  lack  of  a  more  complete  description. 

Schluter  (1838:  38)  considered  Lamarck's  Balanus  duploconus  to  represent  a  distinct 


169 


genus,  for  which  he  proposed  Duplocona,  with  D.  laevigata  Schluter  (=Balanus  dupl- 
oconus  Lamarck,  1818)  as  the  sole  nominal  species.  Although  D.  laevigata  is  accompanied 
by  a  reference  to  Lamarck's  work,  no  description  or  illustrations  are  given,  the  section  on 
barnacles  being  for  the  most  part  a  list  of  names.  Pilsbry  (1916:  261),  without  any  comment, 
placed  Schluter's  taxon  in  the  synonymy  of  Pvrgoma  as  then  recognized.  Because  of  the 
dubious  nature  of  Balanus  duplocomis,  Duplocona  cannot  be  defined. 

Both  Creusia  childreni  and  Megatrema  semicostata  are  presented  without  description 
or  locality.  Sowerby's  illustration  of  the  external  surface  of  A/,  semicostata  is  too  small  and 
generalized  to  be  of  any  taxonomic  value.  Creusia  childreni  was  not  figured. 

In  his  "Illustrations  Conchyliologiques"  Chenu  (1843)  figured  seven  species  of 
Creusia  and  five  species  of  Pvrgoma.  Of  these,  only  Creusia  grandis,  C  decorata,  C.  mad- 
reporarum,  Pvrgoma  cancellatum,  and  P.  crenatum  can  be  identified  with  any  certainty. 
The  lack  of  text,  figure  explanations,  or  locality  data,  precludes  identification  of  the  re- 
maining seven  species. 

The  illustration  of  Creusia  striata  presented  by  Chenu  shows  only  the  internal  surface 
of  the  shell  in  situ.  Close  inspection  reveals  six  fines  marking  the  interior  surface  of  the 
sheath,  indicating  that  the  wall  is  composed  of  six  plates.  Therefore,  Chenu's  form  is  ei- 
ther a  species  of  Balanus  or  Hexacreusia. 

Darwin  (1854:  364  footnote)  stated  that  J.  E.  Gray  thought  Pvrgoma  stellata  Chenu 
was  a  synonym  of  P.  conjugatum  Darwin.  However,  Darwin  commented  that,  ".  .  .  it  may 
be  so;  but  the  figure  given  of  the  shell  will  do  equally  well  or  rather  better  for  the  Pvrgo- 
mum  [sic]  dentatum  of  this  work,  and  for  some  varieties  of  P.  crenatum."''  The  uncertainty 
regarding  the  identity  of  P.  stellata  stems  from  the  lack  of  illustrations  of  the  opercular 
plates. 

ACKNOWLEDGMENTS 

It  is  with  great  pleasure  that  we  acknowledge  the  assistance  of  numerous  friends  and  associates  during  the 
more  than  thirteen  years  we  have  devoted  to  preparing  this  revision.  We  are  especially  indebted  to  Patricia  L. 
Barker,  British  Museum  (Natural  History);  Roger  Batten,  American  Museum  of  Natural  History:  Harold  K. 
Brooks,  University  of  Florida;  William  K.  Emerson,  American  Museum  of  Natural  History;  Peter  Glynn. 
Smithsonian  Tropical  Institute  of  Research;  Ivan  Goodbody,  University  of  West  Indies;  the  late  Thomas 
Goreau.  University  of  the  West  Indies;  J.  P.  Harding,  British  Museum  (Natural  History);  Meredith  L.  Jones. 
Smithsonian  Institution;  Harry  Ladd,  U.  S.  Geological  Survey;  John  Moyse,  Iloilo  University,  Philippines;  An- 
drez  Radwahski,  University  of  Warsaw;  the  late  William  J.  Rees,  British  Museum  (Natural  History);  Jack  Rud- 
loe.  Gulf  Specimen  Corporation;  Donald  Squires,  Smithsonian  Institution;  Huzio  Utinomi,  Seto  Marine 
Biological  Laboratory;  Stephen  Wainwright,  Duke  University;  and  Victor  A.  Zullo,  California  Academy  of  Sci- 
ences. 


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Department  of  Paleontology,  Natural  History  Museum,  P.O.  Box  1390,  San  Diego, 
California  92112,  and  Scripps  Institution  of  Oceanography,  La  Jolla,  California  92037. 


O-  fvi-J 


MUS.  CO  MP.  ZOOL 
LIBRARY 

HARVARD 
UNlVeRSITYi 


BIOLOGY,  GEOGRAPHICAL  DISTRIBUTION,  AND  STATUS 
OF  ATTEVA  EXQUISITA  (LEPIDOPTERA:  YPONOMEUTIDAE) 


JERRY  A.  POWELL,  JOHN  ADAMS  COMSTOCK 
AND  CHARLES  F.  HARBISON 


TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  13  14  MAY  1973 


BIOLOGY,  GEOGRAPHICAL  DISTRIBUTION,  AND  STATUS 
OF  ATTEVA  EXQUISITA  (LEPIDOPTERA:  YPONOMEUTIDAE) 

JERRY  A.  POWELL,  JOHN  ADAMS  COMSTOCK  AND  CHARLES  F.  HARBISON 


ABSTRACT.— Moths  of  the  genus  Altera  are  colorful  insects  that  are  often  encountered  at  flowers  during 
daytime.  Taxonomic  relationships  among  the  ten  described  New  World  species  are  poorly  known.  Three  spe- 
cies occur  in  Nearctic  North  America:  the  widespread  punciella  Cramer,  the  quite  similar  exquisita  Busck 
which  was  formerly  known  only  from  northern  Mexico,  and  a  dissimilar  endemic  in  Florida,  floridana 
Neumoegen.  The  geographical  distribution  and  status  o{ exquisita  in  relation  io  punctella  are  analyzed.  The 
two  are  allopatric  with  a  meeting  and  possible  blend  zone  near  the  Rio  Grande.  The  northward  spread  of 
punctella  following  the  distribution  of  adventive  A  ilanthus  trees  is  documented.  The  biology  and  behavior  of 
exquisita.  based  primarily  on  a  recently  discovered  colony  in  southeastern  California,  and  the  relationship  of 
the  distribution  of  this  moth  to  various  Simaroubaceae  are  examined.  After  a  long  premating  period,  mating 
takes  place  at  the  onset  of  the  light  phase  of  the  diel  cycle;  oviposition  occurs  at  the  end  of  the  light  phase,  on 
fibrous  substrates,  probably  mainly  in  the  larval  webbing  in  the  field;  larvae  feed  gregariously  in  a  tent-like 
shelter,  using  flowers  and  seed  o{  Castela  emoryi  and  also  foliage  of  more  leafy  host  plants,  reaching  maturity 
in  about  50  davs  at  laboratory  temperatures;  pupation  occurs  in  a  frail  cocoon  within  the  larval  tent;  the  pupa 
remains  in  place  at  emergence  of  the  moth.  The  species  is  multivoltine.  apparently  without  a  diapause  period. 
The  egg,  larva,  and  pupa  are  described. 

Members  of  A  tleva  are  colorful,  rather  conspicuous  moths  that  are  encountered  during 
the  daytime  at  flowers  but  apparently  are  essentially  crepuscular  and  nocturnal  in  behavior. 
The  genus  is  primarily  Pan-Tropical  in  distribution,  consisting  of  some  50  species,  half  of 
which  are  described  from  the  Indo-Malayan  area.  About  ten  New  World  species  have  been 
described,  and  these  are  concentrated  in  the  Antillean  and  circum-Caribbean  region.  There 
are  three  species  represented  in  the  Nearctic:  the  v/idespread  punctella  (Cramer)  (  =  aurea 
Filch).,  floridana  Neumoegen  in  Florida  and  exquisita  Busck,  previously  known  only  from 
northern  Mexico.  Despite  the  relatively  large  size  of  the  individuals  compared  to  most  so- 
called  Microlepidoptera,  and  their  conspicuousness,  no  satisfactory  taxonomic  treatment 
exists  for  New  World  species.  Most  of  the  described  Neotropical  species  have  been  in- 
adequately sampled  to  enable  firm  conclusions  on  geographical  variation  and  relation- 
ships. 

In  1966  our  attention  was  called  to  the  occurrence  ofanAtteva  near  Coyote  Wells,  Im- 
perial County,  in  the  Colorado  Desert  area  of  southern  California,  when  larvae  were  col- 
lected by  R.  V.  Moran.  Records  at  the  California  Department  of  Agriculture,  Sacramento, 
showed  that  this  moth  had  been  discovered  in  California  by  R.  A.  Flock  of  the  University  of 
California,  Riverside,  who  collected  larvae  "near  Seeley,"  Imperial  County  in  November, 
1964.  Probably  the  actual  source  was  the  same  colony  from  which  our  collections  were 
made.  Moths  reared  from  the  Moran  collection  were  determined  asAtteva  exquisita  Busck 
(1912),  described  from  Mobano,  Coahuila,  Mexico.  This  species  has  received  little  notice 
since  its  original  description,  and  apparently  it  was  not  collected  again  until  recently. 

The  California  population,  which  is  located  in  an  isolated  grove  of  Castela  (Hola- 
cantha)  emoryi  Gray  (Simarubaceae)  seven  airline  miles  southeast  of  Coyote  Wells,  Impe- 
rial County,  was  used  as  the  principal  source  of  material  in  our  biological  study.  The  locality 
was  revisited  by  Calvert  Norland  and  Powell  in  June,  1966;  by  Harbison  in  December, 
1966;  and  by  Powell  in  October,  1967,  and  June,  1968,  to  obtain  additional  information  on 
the  habits  of  this  moth.  Our  observations,  together  with  data  based  on  scattered  collections 
made  in  Baja  California,  Mexico,  should  prove  of  value  in  assessment  of  comparative  biol- 
ogy when  a  comprehensive  study  of  relationships  in  American  Atteva  is  realized.  Various 
aspects  of  the  bionomics  of  Atteva  punctella  have  been  recorded  from  the  eastern  United 
States,  and  that  species  recently  has  been  exten.sively  studied  in  Connecticut  (Taylor,  1966, 
1967).  Nothing  has  been  reported  previously  on  the  biology  of  exquisita. 

SAN  DIEGO  SOC.  NAT.  HIST.  TR.ANS.  17(  13):  175-186.  14  MAY  1973 


176 


Figures  1-10.  Adults  of /I //eva:  \,punctella  female.  5  mi.  W.  Cave  City,  Ky.  VIII-3/4-71  (J.  Powell)  1,  punctella 
male,  Alexandria,  Va.  IX-17-70  (J.  Powell).  3,  exquisita  female,  3  mi.  E.  Galeana.  N.L..  Mex.  VIII-7/9-63  (Duck- 
worth &  Davis).  4,  excjuisiia  male,  7  mi.  SE.  Coyote  Weils,  Calif.  V 1-25-66,  r.  f.  Holacantlui  emoryi  (J.  Powell  No. 
66F13)  5,  exqiiisita  female,  isla  San  Francisco,  Goifo  de  California.  Mex.  IV- 17-62  (Harbison),  b.exquisiia  female. 
21  mi.  W  La  Paz.  Baja,  Calif.  Mex.  V 11-9-66 (J.  A.  Chemsak).  7,  ^. punctelltiXexc/uisiia {pufMive  blend  zone  pop- 
ulation) female  and  male,  20  mi.  S.  Sabinas  Flidaigo.  N.L..  Mex.  Vll-7-66  (Buckett  &  Ciardner).  9.  aberrant  fe- 
male same  data.  10,  exquisita  aberrant  female,  same  data  as  fig.  4. 


177 


PHENOTYPIC  VARIATION 

Atteva  exquisita  differs  from  A.  punctella,  a  widespread  Neotropical  species  originally 
described  from  Surinam,  and  other  similar  described  species  (Walsingham,  1914)  by  having 
the  yellow  transverse  bands  of  the  orange  forewing  paler  and  relatively  unbroken  by  dark 
lines.  In  particular,  the  submedian  band  is  composed  of  about  4  to  7  separate  pale  spots  in 
punctella  and  2  to  4  in  exquisita;  the  postmedian  band  is  a  granulated-appearing  patch  of  12 
to  16  more  or  less  distinct  spots  in  punctella,  whereas  in  exquisita  these  spots  are  partially 
confluent,  numbering  about  6  to  8  (Figs.  1-6).  Although  both  punctella  and  exquisita  are 
variable  in  details  of  forewing  markings,  neither  varies  to  an  extent  that  field  collected  series 
of  one  include  individuals  with  the  wing  pattern  characteristic  of  the  other. 

To  what  degree  these  differences  reflect  different  environmental  effects  acting  directly 
on  individuals  rather  than  expression  of  genetic  characteristics  of  the  populations  is  un- 
known. O.  R.  Taylor  (in  litt.)  has  shown  extreme  variability  in  laboratory  stocks  and  be- 
lieves there  are  many  temperature  labile  genes  in  punctella.  Color  features,  including  the 
quality  of  orange,  amount  of  melanic  reticulation  on  the  transverse  yellow  areas  of  the  fore- 
wing, and  paleness  of  the  yellow  vary  with  temperature.  Taylor  stated,  for  example,  that  the 
melanic  reticulations  all  but  disappear  in  the  two  proximal  bands  at  high  temperatures.  The 
existence  of  reduced  melanic  lines  in  exquisita  in  widespread  desert  areas  would  seem  to 
corroborate  this  correlation,  although  this  feature  may  be  genetically  fixed.  In  addition, 
frequent  occurrence  of  striking  aberrations  (Figs.  9,  10)  both  in  the  field  and  in  moths 
reared  from  field  collected  larvae  suggests  caution  should  be  exercised  in  forming  con- 
clusions about  genetic  relationships  reflected  by  the  various  phenotypic  expressions. 

California  exquisita  adults  vary  in  color,  both  in  the  quality  of  the  orange  and  in  the 
markings,  which  range  from  whitish  to  yellowish.  The  individuals  that  exhibit  the  palest 
markings,  and  therefore  approach  most  closely  the  type  of  exquisita,  are  those  reared  in 
January.  Those  emerging  in  both  spring  and  fall  have  pale  yellow  transverse  bands,  in- 
distinguishable from  field  collected  specimens  from  various  localities,  including  northern 
Mexico. 

During  this  study  no  morphological  differences  could  be  discovered  among  several  col- 
lections of  Atteva  punctella  from  southern  Mexico  and  the  eastern  United  States.  This  sup- 
ports the  opinion  that  aurea  (Fitch,  1857)  is  a  synonym  of  punctella,  as  was  indicated  by 
Zeller  (1871),  Walsingham  (1897),  and  Forbes  (1923).  Evidently  the  persistent  use  of  the 
name  aurea  for  Nearctic  populations  of  this  species  by  textbooks  and  most  lepidopterists  is 
due  to  their  reliance  on  Holland's  Moth  Book  and  McDunnough's  Check  List.  We  concur 
with  Taylor  ( 1967)  in  treating  aurea  as  a  synonym. 

The  separation  of  exquisita  as  a  species  is  also  suspect,  because  it  is  an  allopatric  coun- 
terpart in  western  arid  regions  and  is  also  very  similar  in  morphological  details  (including 
genitalia).  Probably  exquisita  should  be  treated  as  a  subspecies  of  punctella  pending  in- 
vestigation of  the  nature  of  the  color  differences. 

GEOGRAPHICAL  DISTRIBUTION 

Atteva  exquisita  is  widespread  in  desert  and  thorn  forest  areas  of  northern  Mexico  and 
southwestern  United  States  (Appendix;  Fig.  1 1 ).  In  addition  to  the  type  locality,  this  species 
has  been  taken  in  Nuevo  Leon  in  Northeastern  Mexico,  while  westward,  it  has  been  col- 
lected in  the  plateau  region  in  southern  Chihuahua,  and  at  a  number  of  stations  in  southern 
Baja  California  and  along  the  Gulf  of  California.  It  ranges  thence  northward  into  southern 
California  and  Arizona. 

Available  information  indicates  that  exquisita,  like  related  species,  is  restricted  to  Sim- 
arubaceae  for  larval  foodplants.  The  species  is  more  widespread  than  any  genus  of  Sim- 
arubaceae  in  this  part  of  the  continent  (Standley,  1923),  but  a  complex  of  essentially 
allopatric  members  of  the  family  comprise  a  distributional  pattern  corresponding  to  that  of 
exquisita.  Thus  the  host  in  Coahuila  and  Nuevo  Leon  presumably  is  Castela  (C.)  texana 
(Torrey  and  Gray),  while  Alvaradoa  amorphoides  Liebm.  is  available  in  southern  Chi- 
huahua; the  moths  have  been  associated  with  Castela  (C.)  peninsularis  Rose  in  southern 
Baja  California,  with  Castela  (Eremacantha) polvandra  Moran  and  Felger  ( 1968)  in  the  cen- 


178 


Figure  1 1 .  Geographical  distribution  of  A  tteva  exquisita  Busck,  according  to  material  cited  in  the  Appendix.  Half- 
closed  circles  indicating  localities  where  the  phenotype  of  samples  suggests  a  possible  blend  zone  with  A.  punctella 
to  the  north.  The  type  locality  (TL)  of /I.  exquisita,  Mobano,  Coahuila,  is  indicated,  but  this  place  has  not  been 
located  on  maps  we  examined. 

tral  Gulf  of  California  region,  and  with  Castela  (Holacantha)  emorvi  in  southern  California. 
This  last  plant  presumably  also  serves  as  the  foodplant  in  Arizona. 

The  most  commonly  encountered  and  best  known  species  of  American  A  tteva,  punc- 
tella (=  aurea),  is  widespread  in  the  eastern  United  States.  The  species  is  assumed  to  be 
adventitious  from  some  Neotropical  area  because  its  host  ipXdini,  A ilanthus  altissima  (Mill.) 
(Simarubaceae)  is  an  introduced  ornamental  tree  from  Asia.  The  plant  was  brought  to 
North  America  by  way  of  Europe  about  1784  (Davies,  1941),  and  it  adapted  and  voluntarily 
spread  so  that  by  the  middle  of  the  nineteenth  century,  when  the  A  tteva  was  first  formally 
noticed,  it  was  widespread  in  the  eastern  United  States. 

A  tteva  aurea  was  described  by  Fitch  (1857),  who  received  specimens  from  a  corre- 
spondent in  Savannah,  Georgia.  Clemens  ( 1861 )  redescribed  the  moth  (as  compta)  based  on 
specimens  from  Texas.  The  species  was  found  to  be  common  on  Ailanthus  in  Missouri  by 
Riley  ( 1869),  and  general  statements  in  the  literature  listed  >4.  punctella  (as  aurea)  from  the 
Gulf  States  and  thence  southwestward.  No  reports  of  its  presence  in  the  Washington,  D.C.— 
Pennsylvania  areas  were  made  by  Clemens,  Riley,  or  other  early  entomologists  of  the  re- 
gion. Later,  occurrence  of  the  species  was  recorded  at  Raleigh,  North  Carolina  (Brimley, 
1909),  at  Philadelphia,  Pa.  (Ilg,  1911),  and  at  Baltimore,  Maryland,  and  in  Illinois  (Gibson, 
1920).  The  distribution  was  summarized  as  New  York  to  Illinois  and  southward  by  Forbes 
(1923).  Thus  the  fragmentary  record  suggests  that  the  extension  to  approximately  42°  N 
latitude  was  the  result  of  spread  during  the  half  century  between  1870  and  1920.  The  range 
may  not  have  expanded  much  subsequently,  but  Taylor  (in  litt.)  has  seen  specimens  from 
Minnesota  (St.  Paul)  and  southeastern  South  Dakota  (Yankton).  We  have  specimens  from 
northern  Wisconsin  (Lake  Katherine,  Oneida  County,  H.  M.  Bower)  collected  in  1961, 
which  represent  the  northern  record  we  have  seen,  about  46°  N. 

Although  Ailanthus  is  the  only  host  over  most  of  eastern  North  America,  in  southern 
regions  three  native  species  of  Simarubaceae  are  available  (Small,  1903).  In  southern  Texas 
Castela  texana  (Torrey  and  Gray)  is  native,  while  in  Florida  both  Picrannia  pentantra  Sw. 
and  Simarubra  glauca  DC.  may  serve  as  hosts.  The  latter  is  used  by  a  related  species,  ^//eva 
floridana  Neumoegen,  but  is  not  known  to  be  a  host  plant  o^ punctella  (Dyar,  1897;  Kimball, 
1965). 

Thus  it  seems  likely  thai  punctella  was  not  introduced  into  the  United  States  by  man 


179 


but  merely  represents  a  northern  component  of  populations  that  were  native  in  the  West 
Indies,  or  Florida  and  (or)  southern  Texas.  A  spread  northward,  as  Ailanlhus  became  suffi- 
ciently abundant  to  support  populations,  presumably  occurred  from  the  nearest  geogra- 
phical areas  in  which  the  species  lived. 

As  noted,  the  close  similarity  and  allopatry  of punctella  and  exquisita  suggest  that  they 
are  geographical  components  of  a  single  species.  The  blend  zone  between  the  two,  or  pos- 
sible sympatric  occurrence,  is  to  be  expected  in  southern  Texas  or  areas  of  Mexico  near  the 
Rio  Grande.  Material  for  study  from  this  region  has  been  limited.  A  series  from  the  vicinity 
of  Galeana  in  southern  Nuevo  Leon  shows  a  phenotype  similar  to  the  type  of  exquisita  in 
reduction  of  dark  lines  in  the  forewing  pattern.  However,  a  single,  worn  specimen  from  Val- 
lecillo,  and  a  good  series  from  20  miles  south  of  Sabinas  Hidalgo  in  northern  Nuevo  Leon 
are  less  similar.  The  submedian  band  is  relatively  unbroken,  as  in  exquisita,  while  the  post- 
median  band  has  more  extensive  dark  lines,  differentiating  about  9  to  13  pale  spots,  an  in- 
termediate condition  between  exquisita  and  punctella  (Figs.  7,  8).  These  specimens  lend 
credence  to  the  supposition  that  the  populations  here  treated  as  exquisita  represent  a  west- 
ern, arid  country  subspecies  of  punctella. 

BIOLOGY  AND  BEHAVIOR 

Observations  were  made  on  plant  associations  of  the  moths  at  the  Imperial  County, 
California,  site  and  in  two  areas  of  Baja  California,  on  Isla  San  Francisco  in  the  Gulf  of 
California,  and  in  the  vicinity  of  La  Paz.  Behavior  of  the  adults  was  studied  in  the  labora- 
tory, using  reared  moths  from  Coyote  Wells  and  employing  glass  jar  breeding  cages  with  a 
screen  ceiling  of  nylon  (Powell,  1964)  or  a  portable  type  consisting  primarily  of  a  plastic 
cylinder  17  x  30cm.  Larval  and  pupal  habits  in  the  field  were  recorded  only  at  the  Imperial 
County  locality. 

Adult.— T)\xx\ng  the  daytime  adults  of  both  sexes  were  found  on  the  host  plant,  as  well 
as  at  flowers  of  other  plants.  Diurnal  visitation  of  various  flowers  has  also  been  recorded  for 
Atteva punctella  (Brimley,  1909;  Ilg,  1911;  Riley,  1869).  At  Isla  San  Francisco  .4.  exquisita 
was  taken  in  association  with  Castela peninsularis,  a  presumed  foodplant;  at  Coyote  Wells  a 
few  were  found  on  the  larval  webs  and  flowers  of  Castela  emoryi;  while  in  the  La  Paz  area 
adults  were  visiting  flowers  of  Wislizenia  refracta  Englem.  (Capparidaceae)  and  two  un- 
identified shrubs.  The  moths  were  observed  at  midday  but  were  not  witnessed  flying.  Sev- 
eral individuals  were  taken  at  light.  The  species  was  attracted  in  numbers  to  fluorescent 
blacklight  at  two  localities  in  Nuevo  Leon.  Atteva  punctella  is  also  commonly  collected  at 
lights. 

Under  laboratory  conditions,  several  groups  of  adults  (totaling  about  80  individuals) 
were  caged  during  a  sequence  of  nine  weeks  in  the  summer  of  1966  and  in  January,  1967. 
Cages  were  stored  in  one  of  three  conditions:  a)  at  outdoor  temperatures  (which  in  inland 
Contra  Costa  County,  California,  are  lower,  especially  the  nightly  minima,  than  would  be 
expected  at  the  Coyote  Wells  site);  b)  at  variable  room  temperature  ( 15-20  C);  and  c)  in  a 
temperature  controlled  laboratory  at  20  ±1  C  Atteva  exquisita  proved  to  be  a  hardy  moth, 
relative  to  many  Microlepidoptera,  and  successful  mating  and  oviposition  were  obtained  in 
all  three  situations.  Individual  moths  lived  4  to  36  days  and  averaged  about  15  days.  Mating 
and  oviposition  took  place  primarily  during  the  first  few  days  after  caging  and  neither  oc- 
curred after  the  tenth  day.  Females  survived  in  dry  vials  up  to  four  days,  but  the  moths  were 
observed  to  take  v^ater  readily,  even  during  midday,  especially  after  periods  when  none  had 
been  available. 

The  moths  appear  to  be  primarily  crepuscular  in  activity,  but  some  phases  of  behavior, 
notably  mating,  apparently  consistently  occur  at  other  times  in  the  diel  rhythm.  Under  nat- 
ural lighting  conditions  the  period  of  greatest  activity  of  caged  adults  was  about  1730  to 
2000  PST,  from  about  1.5  hours  before  sunset  to  1  hour  or  more  after  sunset.  Possibly  tem- 
perature was  a  critical  factor  in  masking  normal  activity  periods,  since  evenings  were  cool, 
usually  below  15  C.  by  nightfall  or  shortly  afterwards.  During  the  late  afternoon  and  dusk 
period,  most  individuals  actively  crawled  about  the  screen  ceiling  of  the  cage  and  occasion- 
ally flew.  At  other  times  of  day  only  occasional  moths  moved;  a  reaction  to  the  observer 
appeared  to  be  a  factor.  Artificial,  overhead  lighting  aff'ected  diurnal  activity  of  exquisita. 


180 


Individuals  exposed  to  this  light  condition  (in  a  temperature  controlled  laboratory)  sporad- 
ically moved  about  without  apparent  external  stimulus,  but  neither  mating  nor  oviposition 
was  observed  under  these  circumstances.  With  the  lights  off,  the  same  motljs  became  less 
active  during  mid  afternoon  than  they  had  been  while  exposed  to  artificial  lighting. 

Pronounced  activity  at  the  side  of  the  cage  towards  lights  was  also  noted  at  night. 
Therefore,  nocturnal  observations  were  made  by  means  of  a  red-covered  flashlight,  which 
did  not  seem  to  affect  the  moths.  In  outdoor  temperature  conditions  they  became  inactive 
by  2030  and  2130,  with  the  temperature  at  16  and  13  C.  on  different  evenings,  while  once 
when  the  temperature  remained  at  19  C.  at  2130,  the  moths  were  still  slowly  crawhng.  At 
2300  with  the  temperature  13  C.  there  was  no  activity,  even  when  a  flashlight  was  directed 
onto  the  moths. 

Mating  by  six  pairs  was  observed,  at  least  once  in  each  of  the  three  cage  situations. 
Pairs  copulated  on  the  second  to  seventh  day  following  emergence  (average  4.8  days). 
Under  controlled  conditions  of  regular  photoperiod  and  constant  temperature  (about  22 
and  28  C),  Taylor  (1967)  found  a  long  premating  period  also  characteristic  in  punctella: 
isolated  pairs  of  virgin  moths  usually  did  not  mate  until  the  third  to  sixth  day  after  eclosion. 
In  each  case  our  exquisita  pair  was  witnessed  at  the  onset  of  morning  observations,  usually 
at  0700,  but  once  at  0510  (after  daybreak),  and  the  moths  remained  apparently  inactive, 
in  copulo  during  the  morning  hours.  Separation  usually  occurred  between  1000  and  1 100. 
In  once  instance  the  pair  was  first  observed  at  1054,  and  they  separated  at  1 148.  In  at  least 
three  examples  the  moths  were  known  to  have  been  not  in  copulo  late  the  previous  evening, 
after  apparent  activity  had  ceased.  Mating  did  not  take  place  during  the  crepuscular  height 
of  individual  movement,  and  no  copulation  was  observed  in  the  field. 

Taylor  (1967)  found  tha{  punclella  males'  responsiveness  and  mating  occurred  chiefly 
during  the  first  30  minutes  of  the  light  period.  Evidently  a  similar  diel  rhythm  obtains  in  the 
mating  behavior  of  exquisita. 

Oviposition  by  exquisita  took  place  in  late  afternoon.  Females  engaged  in  a  character- 
istic behavior  pattern,  walking  slowly,  with  the  abdomen  extended  and  curved  ventrad.  The 
extended  ovipositor  could  be  seen  to  press  against  the  twig  or  protrude  through  the  nylon 
mesh  of  the  cage  ceiling.  Oviposition  was  a  slow  process,  often  requiring  two  to  three  min- 
utes at  one  egg  site,  and  no  female  was  observed  to  deposit  a  second  egg  without  moving. 
Our  observations  corroborate  those  of  Taylor  (1967)  on  punctella  in  a  controlled  environ- 
ment. He  reported  that  oviposition  generally  began  1  to  3  hours  before  the  end  of  the  light 
period  and  continued  into  the  dark  period. 

Females  of  exquisita  consistently  selected  fibrous  or  pitted  surfaces  for  egg  laying. 
Counts  were  not  made  of  the  various  substrates  used  for  oviposition  in  the  cages,  but  nearly 
all  eggs  were  deposited  on  the  screen,  on  the  foodplant,  especially  in  partiaUy  eaten  flowers 
(Figs.  12,  13),  or  on  the  cotton  used  to  hold  the  foodplant  bouquet.  A  few  were  placed  on 
the  rough  wooden  floor  of  the  cylinder  cage.  None  were  laid  on  the  smooth  walls  of  the 
breeding  cages,  where  the  moths  spent  most  of  their  time  crawling. 

Ilg  (191 1)  stated  that  eggs  of  Atteva punctella  (=  aurea)  were  distributed  through  the 
communal  web.  We  provided  females  with  Castela  emoryi  which  had  been  cleaned  of 
nearly  all  the  silk  webbing.  Selection  of  the  cotton  and  nylon  fibers  for  oviposition  suggests 
that^.  exquisita  uses  the  larval  webbing  in  the  field. 

Riley  (1881)  reported  that  the  egg  of^.  punctella  was  sometimes  laid  on  the  web,  but 
generally  was  attached  to  the  side  of  the  mid-rib  of  the  new  leaves  of  Ailanthus,  where  it 
caused  a  well  defined  swelling  of  the  leaf  vein.  Riley  attributed  this  to  a  toxic  substance 
which  he  supposed  was  secreted  during  oviposition. 

Egg.— Eggs  of  exquisita  required  8  to  9  days  for  development  at  20  ±1  C.  Groups  of 
eggs  were  stored  in  a  refrigerator  at  4  C  five  or  six  days  (third  to  eighth  day  and  about  fifth  to 
eleventh  day)  and  their  development  took  15  to  16  days.  Eggs  stored  in  a  dry  refrigerator  for 
several  weeks  did  not  survive. 

Larva.— A  tendency  for  larvae  of  various  ages  to  live  gregariously  and  occupy  a  com- 
mon web  is  characteristic  for  Atteva  {e.g.,  punctella,  (Ilg,  191 1)  and  fabriciel la  (Swed.)  in 
India  (Fletcher,  1 920)).  Larvae  of  exquisita  produce  copious  amounts  of  silk,  even  in  the  early 
instars,  and  at  the  Coyote  Wells  site  large  inflorescences  of  the  host  plant  were  enmeshed  in 


181 


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JM 

W\             ^N        ^9^ 

^^mSii-               .^m 

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Figures  12,  13.  Eggs  of  A i leva  exquisita  Busck.  12,  (left)  eggs  deposited  on  flat,  plastic  surface.  13,  (right)  eggs 
deposited  in  the  laboratory  in  partially  eaten  Holacanlha  flowers. 

fine  webbing.  Larvae  of  various  sizes  were  present  in  the  same  web  in  June.  Since  Castela 
emorvi  is  essentially  leafless,  most  of  the  feeding  took  place  on  the  flowers  and  developing 
seed.  Both  male  and  female  flowers  of  the  dioecious  plant  were  used,  but  in  late  June  larvae 
were  more  abundant  on  the  staminate  inflorescences.  Later  in  the  season,  after  the  flowers 
had  dried,  overwintering  individuals  were  found  only  among  the  seed-bearing  in- 
florescences, where  they  fed  primarily  on  the  seed  covers.  Moran  and  Felger  (1968)  found  a 
similar  situation  with  Castela  polyandra  in  Baja  California,  where  at  each  of  their  localities 
at  flowering  time,  larvae  of  exquisita  "festoon  the  flowering  branches  with  spidery  webs." 
They  noted  that  larvae  also  ate  the  leaves  and  bark. 

In  the  laboratory  newly  hatched  larvae  were  able  to  establish  and  survive  on  25  to  40 
day  old  Castela  stems  with  remnants  of  flowers  eaten  by  preceding  generation  larvae.  The 
bark  was  skeletonized  and  at  times  whole  twigs  were  girdled.  Similar  feeding  behavior  was 
reported  for  Atteva punctella  on  Ailanthus,  a  leafy  plant,  by  Riley  (1869).  Although  the  Cas- 
tela branches  were  kept  in  water,  they  appeared  dry  by  the  time  the  first  instar  larvae  began 
hatching  during  our  study.  When  first  and  second  instar  larvae  were  placed  in  vials  contain- 
ing Castela  twigs  and  fresh  terminal  leaflets  of  Rhus  typhina  (Anacardiaceae)  from  the  Uni- 
versity of  California,  Berkeley,  campus,  only  the  Castela  was  accepted.  A  small  amount  of 
feeding  occurred  on  the  Rhus,  but  no  larvae  successfully  established  on  it.  A  few  larvae  sur- 
vived to  the  penultimate  instar  on  the  dry,  skeletonized  Castela  branches,  but  none  reached 
maturity. 

Other  first  instar  larvae  were  ofl'ered  only  fresh  leaflets  of  Ailanthus  altissima  from  the 
University  of  California,  Berkeley,  Botanical  Garden.  Development  on  Ailanthus  was  only 
partially  successful,  with  a  small  percentage  of  first  instar  estabhshment.  Once  estabhshed, 
growth  occurred  at  about  the  same  rate  as  in  larvae  feeding  on  Castela  stems.  The  few  sur- 
viving individuals  on  Ailanthus  did  not  reach  maturity,  but  rearing  conditions  other  than 
the  food  may  have  been  critical.  Larvae  died  in  the  penultimate  and  antepenultimate  in- 
stars,  when  15  to  20  days  old. 

Larger  larvae  taken  from  Castela  in  May  accepted  Ailanthus  foliage  from  San  Diego 
and  development  proceeded  successfully.  However,  penultimate  and  final  instar  larvae  col- 
lected in  December  failed  to  accept  seeds  of  Ailanthus  from  Contra  Costa  County  even 
when  these  were  sliced  longitudinally,  exposing  the  soft,  inner  tissue. 

An  attempt  was  made  to  determine  the  number  of  instars  through  head  capsule  meas- 
urements. However,  data  are  mconclusive,  owing  to  lack  of  material  representing  the  inter- 
mediate instars.  Probably  five  or  six  instars  are  normal,  and  it  is  likely  that  the  number 
diff"ers  depending  upon  circumstances.  Larvae  subjected  to  adverse  conditions,  such  as  dur- 
ing winter,  may  undergo  an  additional  moult.  The  largest  head  capsule  measurements  orig- 
inate from  larvae  collected  in  October  and  December.  There  is  considerable  overlap  in  the 
size  of  the  final  two  instars  when  individuals  representing  various  seasons  are  considered 


182 


15 


10 


0 


mm       .34 


.68 


1.02 


1.36 


.70 


2.04 


Figure  14.  Distribution  of  measurements  of  head  capsule  width  in  la.TVd\  Aiieva  exquisiia  Busck;  based  on  samples 
taken  in  various  seasons  from  one  site  near  Coyote  Wells,  California.  Filled  squares  indicate  head  capsules  of 
preserved  larvae;  cross-hatched  squares  indicate  head  capsules  shed  by  pre-final  instar  larvae;  open  squares  in- 
dicate head  capsule  widths  of  final  instar  exuviae  (estimated  byconversion  from  measurements  of  frontal  triangle). 

together  (Fig.  14). 

Pupa —At  maturity  larvae  construct  a  frail  silken  network  in  which  they  suspend  for 
pupation.  In  the  field  pupae  were  found  within  the  inflorescence,  particularly  in  lower  por- 
tions of  the  webbing  which  the  larval  colony  inhabited.  The  extremely  thin  cocoon  appears 
to  be  a  biological  feature  associated  with  a  behavioral  tendency  to  remain  in  the  communal 
shelter,  and  a  tendency  to  wander  in  the  laboratory  probably  is  abnormal.  Cocoons  also 
occur  in  the  larval  webs  in punctella  (Ilg,  1911)  and  A  tfevafloridana  (Dyar,  1897). 

Construction  of  the  cocoon,  a  quiescent  prepupal  period,  and  transformation  to  the 
pupa  required  about  48  hours.  Cocoons  spun  in  isolation  from  other  larval  webbing  were  so 
thin  as  to  be  almost  invisible.  They  were  rather  flat,  roughly  oval,  measuring  about 
25  X  35mm  in  outline,  with  an  irregular  interior  network  and  a  slightly  heavier  inner  cocoon 
some  15mm  in  length  in  which  the  pupae  was  suspended,  held  about  2mm  from  the  sub- 
strate. 

Metamorphosis  in  the  pupa  required  nine  days  at  room  temperatures  (approximately 
18  to  22  C.  daily  range). 

LIFE  HISTORY 

There  was  no  indication  in  the  laboratory  of  either  obligate  or  facultative  diapause. 
Evidently  there  are  at  least  two  or  three  annual  generations,  with  emergence  of  adults  in 
late  May  or  June,  again  in  July,  and  probably  at  least  once  more  in  late  summer.  Probably 
extreme  summer  conditions  reduce  longevity  of  adults  compared  to  that  in  confinement, 
but,  even  so,  flight  periods  of  generations  overlap.  By  mid-June  all  stages  were  present  at 
the  Coyote  Wells  site,  and  well-defined  generations  probably  are  not  exhibited  from  that 
time  on  through  the  season. 

Our  observations  indicate  a  developmental  period  of  about  40  to  50  days.  Thus  adults 
emerging  in  May  could  produce  a  third  generation  by  late  August  or  early  September.  Sur- 
vey in  October  and  in  late  December  revealed  only  larger  larvae.  No  adults,  eggs,  or  pupae 
could  be  located.  When  brought  into  laboratory  temperatures  these  larvae  continued  devel- 
opment, feeding  on  nearly  dry  seed  covers,  and  produced  adults  within  a  month.  Appar- 
ently individuals  resulting  from  eggs  deposited  in  fall  begin  feeding  and  enter  a  quiescent 
state,  possibly  growing  slowly  by  feeding  during  warm  spells  in  winter,  and  reach  maturity 
by  the  time  Castela  blooms  again  in  spring. 


183 


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Figures  15-17.  Structures  of  last  instar  larva  oi  Aitexa  exquisita  Busck.  15,  setal  arrangements  on  segments  1  and  II 
of  thorax,  and  1,  2,  6,  7,  8,  and  9  of  abdomen;  SP  =  spiracle,  arrows  indicate  variable  loci.  16,  frontal  view  of  head 
capsule,  showing  color  pattern,  black  ventral,  red-brown  median,  and  orange  dorsal.  17,  crotchets  of  abdominal 
proleg. 

Riley  (1869)  believed  that  adults  o{  Atteva  punctella  overwintered,  because  he  was 
unable  to  obtain  oviposition  from  moths  emerging  in  September  and  October.  Taylor  (in 
litt.)  has  no  data  indicating  appreciable  cold-hardiness  in  any  stage  o^ punctella  and  be- 
lieves that  the  species  does  not  overwinter  in  the  northern  part  of  its  range. 

DESCRIPTION  OF  EARLY  STAGES 

f^g'.— Appearing  whitish  when  first  deposited,  turning  pale  yellow  within  48  hours  and  gradually  deeper 
yellow  prior  to  darkening  of  the  embryo.  Pliable  when  deposited  and  assuming  variable  shapes  depending  in  part 
upon  substrate;  on  flat  surfaces,  oval,  flattened,  somewhat  produced  towards  micropylar  end  (Fig.  12),  varying 
from  about  1.05  X  0.60mm  to  1. 15  X  0.65mm;  on  fibrous  and  other  irregular  surfaces,  usually  somewhat  thickened 
and  less  regular  in  outline  (Fig.  13).  Dorsal  surface  more  or  less  regularly  sculptured  with  round,  shallow  pits. 

Riley  (1881)  mentioned  the  variable  form  of  the  eggs  xn  punciella  and  indicated  that  one  end  frequently  was 
produced  into  a  "short  neck."  Riley  gave  the  length  as  0.9mm,  and  Peterson  ( 1967),  who  published  a  photograph 
of  eggs  of//,  punctella  (  =  aurea).  gave  0.9  X  0.5mm.  slightly  smaller  than  anv  observed  in  our  study. 

La/Tfl-.— First  Instar:  Head  capsule,  width  0.27  to  0.31mm;  pale  when  teneral,  becoming  dark  brown,  slightly 
paler  towards  frontal  triangle.  Body,  length  about  2.3  mm  when  teneral,  3.0  to  3.4  mm  when  fully  fed;  integument 
unpigmented  at  first,  becoming  lightly  mottled  with  reddish  grav  specks,  without  a  distinct  pattern.  Thoracic  shield 
scarcely  discernible;  anal  shield  unpigmented.  Setae  and  pinacula  minute  and  colorless;  arrangement  apparently 
very  similar  to  that  of  final  instar.  Abdominal  proleg  crotchets,  6.  in  an  irregular  circle;  anal  proleg  crotchets,  8  or  9. 

Second  Instar:  Head  capsule,  width  0.41  to  0.58mm  (possiblv  to  0.68mm).  dark  brown  with  well  defined  pale 
spots  on  front  and  crown,  corresponding  to.  but  proportionately  larger  than,  those  of  last  instar.  Body,  length  about 
3.4  to  4. 1  mm;  integument  heavily  mottled  with  brownish  gray,  tending  to  form  dorsolateral  bands  which  contrast 
with  the  relatively  large,  unpigmented  DL  pinacula.  Thoracic  shield  brown,  well  defined.  Setae  relatively  elongate, 
unpigmented.  apparently  arranged  as  in  final  instar.  Abdominal  proleg  crotchets,  10  to  12,  uniordinal,  in  a  regular 
circle;  anal  proleg  crotchets,  about  14,  irregularly  biordinal. 

'Description  based  on  specimens  from  La  Paz  area  of  Baja  California;  females  collected  in  August,  1966. 
■Based  on  specimens  from  the  Coyote  Wells  site  in  California;  distended  in  K.A.A.D.;  first  two  instars  from  eggs 
deposited  by  females  reared  in  July;  late  instars  from  larvae  collected  in  May.  June,  and  December. 


184 


Figure  18.  Larva  and  pupa  of  Aiieva  exquisiia  Busck.  A,  Penultimate  instar,  dorsal  aspect.  B.  penultimate  instar. 
head  capsule,  anterior  aspect.  C.  pupa,  dorsal  aspect.  D,  pupa,  ventral  aspect.  E,  pupa,  caudal  area,  showing  cre- 
master  structure. 


Penultimate  Instar:  (Figs.  ISA,  B).  Head  capsule,  width  (possibly  from  1.05  mm)  1.20  to  1.60  mm;  nearly 
unicolorous,  black,  at  times  deep  red-brown  at  crown;  round  unpigmented  spots  surrounding  setal  bases,  appear- 
mg  white  on  the  living  larva.  Body,  length  about  1 1  to  19  mm;  longitudinally  striped  with  orange,  black,  and  white, 
color  pattern  variable,  consisting  of  small  spots,  blotches  and  irrorations;  a  broad  dorsal  band  of  orange  or  och- 
reous-orange(  absent  on  prothora.x),  sprinkled  with  white  and  dark  orange  spots,  enclosing  a  thin,  mid-dorsal  whit- 
ish line,  margined  by  a  fine,  broken,  black  edging;  an  irregular  dorsolateral  band  of  black,  subtended  by  a  lateral 
band  of  blackish  which  is  heavily  sprinkled  with  white  dots  and  irregular  spots;  a  narrt)w  spiracular  band  of  orange 
runnmg  the  length  of  abdomen,  lacking  on  thorax,  venter  mottled,  black  and  white.  The  relatively  small  prolegs 
banded  yellowish  and  black.  Setal  arrangement  not  differing  from  final  instar.  Thoracic  legs  heavily  sclerotized, 
black.  Abdominal  proleg  crotchets,  about  26  to  34,  in  an  irregular  biordinal  tt)  triordinai  circle;  anal  proleg  crot- 
chets, about  24  to  28,  more  or  less  evenly  biordinal. 

FINAL  INSTAR:  Head  capsule,  width  1.43  to  2.00  mm;  black  bek)w  middle,  red-brown  to  orange  at  crown, 
the  pattern  and  extent  of  black-to-brown  marking  variable  (Fig.  16),  spots  surrounding  setal  bases  unpigmented, 
appearing  white  on  the  living  larva.  Body,  length  about  16  to  25  mm;  color  pattern  variable,  similar  to  penultimate 
instar,  generally  less  black  pigment,  with  corresponding  brighter  orange  and  paler  dark  bands.  Spiracles  on  ab- 
dominal segments  I  to  7  very  small,  scarcely  larger  than  base  of  DL  seta.  Setal  arrangements  as  in  Figure  15;  pina- 
culi  moderately  strongly  upraised,  whitish;  setae  unpigmented,  mostly  elongate,  L  setae  on  abdominal  segments 
very  small.  Abdominal  proleg  crotchets,  about  36  to  40,  an  irregular  arrangement  of  18  to  20  large  inside  a  pe- 
ripheral circle  of  18  to  19  small  (Fig.  17);  anal  proleg  crotchets,  about  32  to  35,  a  marginal  row  of  14  to  16  small 
spurs  followed  posteriorly  by  irregularly  scattered  large  crotchets. 


185 


Setal  characteristics  ofiwc/uisiia  dii  nut  ditTcr  appreciahlv  from  those  given  by  Mathur  ( 1960)  in  his  excellent 
description  of  the  larva  offahriciella  (Swed.).  This  and  the  partial  description  ol'punciella  given  by  F\'terson  ( 1956) 
indicate  a  close  similarity  in  larvae  of  various  species  in  this  genus.  The  abdominal  proleg  crotchets  apparently  are 
more  irregularly  arranged  in  fabriciella.  and  the  anal  crotchets  are  more  numerous  (  ±43)  in  that  species.  Although 
variable  in  placement  and  number  to  some  extent,  the  abdominal  proleg  crotchets  are  more  numerous  than  those 
of  the  anal  proleg  in  all  instars  of  exquisiia.  whereas  the  reverse  is  true  in  fabriciella  according  to  Mathur. 

Pupa.— (Figs.  18C-E)  Length  about  12  mm:  greatest  width  3  mm.  Subfusiform.  head  round  anteriorly,  cauda 
regularly  tapering.  Front  and  sides  of  head  v\  ith  a  number  of  hooked  anchor  setae:  front  with  twt)  small,  whitish 
nodules,  each  with  two  hooked  setae.  Antennae  and  maxillae  reaching  wing  tips,  darker  than  other  appendages. 
General  color  light  brown,  with  darker  shading  on  several  structures,  as  illustrated:  wings  with  several  veins  darker 
than  ground  color.  Dorsum  with  the  longitudinal  white  lines  of  larva,  considerably  obscured;  a  black  longitudinal 
line  lateral  to  each  white  line,  discontinuous  on  anterior  portion  of  abdomen,  unbroken  on  caudal  segments.  Cre- 
master{Fig.  18E)  with  numerous,  fragile,  hooked  anchor  setae,  easily  broken:  apparently  showing  no  regularity  in 
placement. 

Details  of  the  pupal  structure  of  punciclla  have  been  illustrated  by  Mosher  ( 1916). 

ACKNOWLEDGMENTS 

Our  sincere  thanks  are  extended  to  the  following,  who  assisted  in  gathering  field  data  during  the  present 
study:  J.  A.  Chemsak  and  J.  T.  Doyen,  University  of  California,  Berkeley:  R.  V.  Moran.  who  pointed  out  the 
Coyote  Wells  colony,  and  R.  P.  Phillips,  San  Diego  Museum  of  Natural  History:  and  C.  E.  Norland,  San  Diego 
State  College. 

W.  D.  Duckworth.  U.  S.  National  Museum,  provided  assistance  with  the  identification  of  .4 1 leva  excpdsiia. 
and  Paddy  McHenry,  Burbank,  California,  furnished  excerpts  of  some  of  the  literature.  F.  L.  Blanc,  State  Depart- 
ment of  Agriculture.  Sacramento,  supplied  early  records  of  exquisita. 

O.  L.  Taylor.  Jr..  LIni\ersit\  of  Kansas.  Lawrence,  reviewed  a  draft  of  the  manuscript  and  offered  many  help- 
ful suggestions  along  with  unpublished  data  on  various  aspects  of  the  bionomics  of  .Aiieva punciella. 

Joachim  Wolf,  assistant  on  the  National  Science  Foundation  project  (GB-4014)  which  supported  part  of  the 
research  program,  carried  out  some  of  the  maintenance  and  surveillance  of  the  laboratory  colonies,  and  coopera- 
tion by  Anton  Crist,  University  of  California,  Berkeley,  enabled  use  of  plant  materials  from  the  Botanic  Garden. 

Photographs  of  the  eggs  were  made  by  A.  A.  Blaker,  Scientific  Photographic  Laboratory,  University  of  Cali- 
fornia, Berkeley. 

This  research  was  supported  in  part  by  grants  from  the  National  Science  Foundation  (NSF-GB-4014,  NSF- 
GB-6813X). 

APPENDIX 

Data  from  specimens  examined  representing /I //eva  exquisiia  and  populations  of  possible  blend  with/1.  /ji^/jc- 
tella. 
MEXICO 

Baja  California.  Norte:  Bahia  de  Los  Angeles.  VI 1-2-66.  on  Castela  polyandra  (R.  P.  Phillips). 

Baja  California.  Territorio  Sur:  Isla  San  Francisco.  Golfo  de  California.  IV- 17-62  (C.  F.  Harbison).  9  mi.  SW 
La  Paz.  VIIl-10-66.at  fis.nisli:eiiiarefracta(i.  T.  Doyen.  J.  Powell):  VIII-14-66(J.  Powell).  21  mi.  W  La  Paz,  Vlil- 
9-66.  at  fls.  legume  shrub  (J.  A.  Chemsak).  26  mi.  W.'La  Paz.  Vlil- 10-66  (J.  A.  Chemsak).  7  mi.  S  San  Pedro.  Vlll- 
10-66.  at  light  (J.  Powell).  Todos  Santos.  VII-14-57  (D.  Spencer.  R..  J.  &  A.  Ryckman).  1  mi.  SW  Punta  Palmilla. 
IX- 14-67,  at  Bl.  &  white  lights  (J.  A.  Chemsak).  3  mi.  N  San  Jose  del  Cabo,  IX-10.  1 1-67  (J.  A.  Chemsak). 

Chihuahua:  8  mi.  NE  Hidalgo  del  Parral,  VII- 13-64,  at  light  (J.  A.  Chemsak,  J.  Powell). 

Coahuila:  Mobano  (R.  Muller).  Vallecillo.  VI-2-51  (P.  D.  Hurd). 

Nuevo  Leon:  20  mi.  S  Sabinas  Hidalgo.  VII-7-66  (J.  S.  Buckett,  M.  Gardner).  3  mi.  E  Galeana,  VlII-7/9-63 
(Duckworth  &  Davis). 
UNITED  STATES 

Arizona:  24  mi.  SE  Parker,  Yuma  Co.,  IX-5-64  (J.  Haddock). 

California:  Hiway  98,  7  airline  mi.  SE  Coyote  Wells,  Imperial  Co.,  V-22-66,  reared  from  Holacantha  emoryi 

[(R.  V.  Moran):  VI-1 1-66,  reared  fVom  //.  emorvi  (C.  E.  &  B.  Norland):  VI-25-66,  on  H.  emoryi  (J.  Powell):  reared 

from//,  ewonv,  emgd.  VI-29to  V11-13-66(J.  Powell-66FI3);  XI 1-25-66,  reared  from //.  ewon;,  emgd.  1-18,  19-67 

l(C.  F.  Harbison:  JAP-67A6):  X-5-67,  reared  from  //.  emorvi,  emgd.  X-1 1-67  (P.  A.  Opler.  J.' Powell.  P.  A.  Rude: 

JAP-67K68). 


LITERATURE  CITED 

Brimley,  C.  S. 

1909.  List  of  moths  observed  at  Raleigh,  N.Carolina.  Ent.  News  20:  33-41. 

Busck.  A. 

1912.  New  Microlepidoptera  from  Mexico.  Proc.  Ent.  See.  Washington  14:  83-87. 

Clemens,  B. 

1861.  Contributions  to  American  Lepidopterology.  No.  7.  Proc.  Acad.  Nat.  Sci.,  Philadelphia,  1860:  522-547. 


186 


Davies,  P.  A. 

1941.  The  history,  distribution,  and  value  of  Ailanthus  altissima  in  North  America.  Trans.  Kentucky  Acad. 
Sci.  9:  12-124. 

Dyar,  H.  G.  * 

1897.  Oetafloridana  Neumoegen.  J.  New  York  Ent.  Soc.  5:  48. 

Fitch,  A. 

1857.  Third  report  on  the  noxious  and  other  insects  of  the  State  of  New  York.  Trans.  N.Y.  State  Agr.  Soc, 
1856,  16:315-490. 

Fletcher,  T.  B. 

1920.  Life  histories  of  Indian  insects.  Microlepidoptera.  Mem.  Dept.  Agr.  India,  Ent.  Series  6  (1-9),  217  p. 

Forbes,  W.  T.  M. 

1923.  Lepidoptera  of  New  York  and  neighboring  states.  Primitive  forms,  Microlepidoptera,  Pyraloids, 
Bombyces.  Cornell  Agr.  Exp.  Sta.,  Mem.  68,  729  p. 

Gibson,  A. 

1920.  Note  on  the  distribution  of  Altera  aurea  Fitch.  Canadian  Ent.  52:  15. 

Ilg,  C. 

1911.  The  Ufe  history  ofAtteva  aurea  Fitch.  Ent.  News  22:  229. 

Kimball,  C.  P. 

1965.  Arthropods  of  Florida  and  neighboring  land  areas.  Vol.  1.  The  Lepidoptera  of  Florida,  an  annotated 
checklist.  Div.  Plant.  Indust.,  Florida  Dept.  Agr.,  Gainesville. 

Mathur,  R.  N. 

1960.  Setal  arrangement  of  y4//evayaZ>n'aW/aSwederus(Yponomeutidae,  Lepidoptera).  Indian  J.  Ent.  21:1-5. 

Moran,  R.  and  R.  Felger 

1968.  Castela polyandra,  a  new  species  in  a  new  section;  union  of  Holacantha  with  Castela  (Simarubaceae). 
Trans.  San  Diego  Soc.  Nat.  Hist.  15:  31-40. 

Mosher,  E. 

1916.  A  classification  of  the  Lepidoptera  based  on  characters  of  the  pupa.  Bull.  Illinois  State  Lab.  Nat.  Hist. 
12:  17-159. 

Peterson,  A. 

1956.  Larvae  of  Insects.  Lepidoptera  and  Hymenoptera.  Part  1.  3rd  ed.  Publ.  by  author,  Columbus,  Ohio. 

Peterson,  A. 

1967.  Some  eggs  of  moths  from  several  families  of  Microlepidoptera.  Florida  Ent.  50:  125-132. 

Powell,  J.  A. 

1964.  Biological  and  taxonomic  studies  on  tortricine  moths,  with  reference  to  the  species  in  California  (Lepi- 
doptera: Tortricidae).  U.  CaUfomia  Publ.  Ent.,  vol.  32.  318  p. 

Riley,  C.  V. 

1 869.  First  annual  report  on  the  noxious,  beneficial,  and  other  insects  of  the  State  of  Missouri.  Missouri  State 
Board  Agr.,  Jefferson  City,  181  p. 

Riley,  C.  V. 

1881.  Lepidopterological  notes.  Papilio  1 :  106-110. 

SmaU,  J.  K. 

1903.  Flora  of  the  southeastern  United  States.  Publ.  by  author,  New  York  City,  1370  p. 

Standley,  P.  C. 

1923.  Trees  and  shrubs  of  Mexico.  Oxalidaceae-Tumeraceae.  Contr.  U.  S.  Natl.  Herbarium  23:  517-848. 

Taylor,  O.  R. 

1966.  A  study  of  genetics,  sperm  precedence,  and  causes  of  multiple  mating  iwAtteva  punciella  (Cramer) 
(Yponomeutidae,  Lepidoptera).  Unpubhshed  M.S.  thesis,  U.  Connecticut,  Storrs. 

Taylor,  O.  R. 

1%7.  Relationship  of  multiple  mating  to  fertility  in  Atteva punctella  (Lepidoptera:  Yponomeutidae).  Ann. 
Ent.  Soc.  Amer.  60:  583-590. 

Walsingham,  T. 

1897.  Revision  of  the  West  Indian  Microlepidoptera,  with  descriptions  of  new  species.  Proc.  Zool.  Soc.  Lon- 
don, 1897:  54-182. 

Walsingham,  T. 

1914.  Lepidoptera:  Heterocera,  Vol.  4.  Biol.  Centralia  Americana:  327-332.  lij 

Zeller,  P.  C. 

1871.  "First  annual  report  on  the  noxious,  beneficial,  and  other  insects  of  the  State  of  Missouri,"  by  C.  V. 
Riley,  1869  [review].  Stett.  Ent.  Zeit.  32:  178. 


Powell:  Department  of  Entomological  Sciences,  University  of  California,  Berkeley 
94720.  Comstock:  Deceased  subsequent  to  the  writing  of  the  original  draft.  Harbison:  Natural 
History  Museum,  San  Diego,  California  92112. 


MUS.  COMP.  ZOOU 
LIBRARY 

SEP30\974 

HARVARD 
UNIVERSITY 
LIFE  HISTORY  OF  THE 

WESTERN  NORTH  AMERICAN  GOBY, 

CORYPHOPTERUS  NICHOLSII  (BEAN) 


JAMES  W.  WILEY 


TRANSACTIONS 

OF  THE   SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  14  30  OCTOBER  1973 


LIFE  HISTORY  OF  THE 

WESTERN  NORTH  AMERICAN  GOBY, 

COR  YPHOPTER  US  NICHOLSII  ( BEAN ) 

JAMES  W.  WILEY 


ABSTRACT. -The  life  history  of  Coryphopterus  nicholsii  was  investigated  by  field  and  laboratory 
studies  based  primarily  on  a  population  at  Laguna  Beach,  California.  The  species  occurs  in  depths  of 
61T1  to  more  than  60m  on  rock  reefs,  where  it  utilizes  holes  and  undercuts  in  the  rock  for  shelter. 
Crustaceans  (amphipods  and  copepods)  are  the  major  food  item;  mollusks  are  also  taken  in  significant 
numbers.  Echinoderms,  annelids,  mollusks,  bryozoans  and  various  eggs  are  more  important  food  items 
during  fall  and  winter  seasons.  Pelagic  prejuveniles  feed  only  on  copepods.  In  the  study  area  the  sex 
ratio  is  1.7  females  :  1  male.  Juveniles  were  first  observed  on  the  reefs  in  February  1967.  Length- 
frequencies  and  observed  scale  age  groups  show  4  to  5  age  groups.  Paralahrax  nebulifer  and  Lythrypnus 
dalli  prey  on  C.  nicholsii.  Sexual  dimorphism  occurs  in  the  genital  papilla,  size,  length  of  dorsal  and 
anal  fins,  and  nuptial  color  of  pelvic  fins.  Breeding  extended  from  mid-February  to  late  August  in 
1967.  Males  became  ripe  at  55  mm,  in  age-group  II  and  III.  Females  became  ripe  at  about  47  mm,  in 
age-group  II.  Ripe  ovaries  contain  two  egg  groups:  ripe  and  unripe.  The  total  number  of  ripe  eggs  in 
4  individuals  ranged  from  3274  to  4788.  The  spindle-shaped  fertilized  eggs  are  attached  directly  to  the 
overhanging  rock  surface  of  the  nest.  The  larvae  are  pelagic  and  prejuveniles  have  been  taken  far  from 
shore.  Juveniles  as  small  as  21.8  mm  were  found  on  the  reefs  from  February  through  August,  1967. 
The  male  prepares  and  guards  the  nest.  In  courtship  the  male  rushes  at  the  female  and  also  rises  off 
the  bottom  a  few  centimeters  with  the  fins  spread,  before  it  settles  back  down.  This  species  is  terri- 
torial.  A  hierarchy  is  established  in  the  laboratory  aquarium. 

The  gobies  constitute  a  widely  divergent  group  of  fishes,  the  suborder  Gobioidei.  Tliey 
occur  in  the  tropicaL  temperate,  and  subboreal  zones  throughout  the  world,  avoiding  only  the 
polar  regions.  Most  gobioids  are  marine,  but  some  inhabit  fresh  water,  including  a  few  in  tor- 
rential streams. 

Gobies  of  the  New  World  genus  Coryphopterus  are  common  in  the  inshore  waters  of  both 
the  tropical  western  Atlantic  and  eastern  Pacific,  inhabiting  holes  in  shallow  water  coral  reefs  or 
rocks.  Nine  species  have  been  described  in  the  tropical  and  subtropical  western  Atlantic.  Of 
the  two  eastern  Pacific  species,  C  urospihis  is  tropical,  but  C  nicholsii  (Bean),  the  subject  of 
this  paper,  ranges  widely  from  subtropical  to  subboreal  waters. 

STUDY  AREA 

An  intensive  field  study  of  Coryphopterus  nicholsii  was  conducted  at  Laguna  Beach, 
Orange  Co.,  California  from  September  1966  through  January  1968.  The  study  area  consisted 
of  two  rock  reefs:  one  a  shallow,  breaking  reef  which  extends  down  to  10m  is  approximately 
90m  offshore;  the  other  varies  from  15  to  25m  in  depth,  and  is  approximately  1200m  from  the 
shallow  reef,  and  800m  offshore.  Each  is  approximately  45m  long,  and  is  surrounded  by  sand 
bottom. 

METHODS  AND  MATERIALS  EXAMINED 

Tlie  population  at  Laguna  was  sampled  at  weekly  or  biweekly  intervals  with  the  use  of 
SCUBA.  Most  specimens  were  collected  with  a  slurp  gun,  although  some  were  taken  with 
"Chem-Fish."  Other  collections  were  made  in  California  at  Malibu  and  Palos  Verdes,  Los 
Angeles  Co.;  at  Cameo  Shores  and  Aliso  Creek,  Orange  Co.;  and  at  La  Jolla,  San  Diego  Co.;  and 
in  Baja  California,  Mexico,  at  Punta  Banda  (SW  side). 

Methods  of  counting  serial  parts  and  taking  measurements  follow  those  of  Hubbs  and 
Lagler  (1958),  except  that  the  caudal  ray  counts  follow  the  methodology  of  Ginsburg  (1945). 
Tlie  last  two  ray  bases  of  the  dorsal  and  anal  fins  were  counted  as  one  ray.  All  measurements 
were  taken  with  dial  calipers  to  the  nearest  0.1  mm.  Proportions,  obtained  arithmetically,  are 
presented  as  ranges  and  means.    All  measurements  of  body  length  are  standard  lengths  (S.L.). 

SAN  DIEGO  SOC.  NAT.  HIST.,  TRANS.  17(14):  187-208,  30  OCTOBER  1973 


188 

Museum  specimens  examined  were  from  the  following  collections:  Stanford  University 
(SU);  California  Academy  of  Sciences  (CAS);  Scripps  Institution  of  Oceanography  (SIO);  Uni- 
versity of  California,  Los  Angeles  (UCLA);  Los  Angeles  County  Museum  of^Natural  History 
(LACM);and  California  State  College,  Long  Beach  (CSCLB). 

BLUESPOT  GOBY 

Coryphoptems  nicholsii  (Bean) 

Gobius  nicholsii. -Bean,  1881:  469  (original  description;  type  locality,  Departure  Bay,  British  Columbia; 
20  fm.).  Jordan  and  Evermann,  1898:  2218  (specimens  recorded  from  coast  of  British  Columbia). 
Halkett,  1913:  30,  95  (listed;  coast  of  British  Columbia).  Fowler,  1923:  293,300  (Malibu  Cove,  Point 
Firmin,  Newport,  Catalina,  Isthmus  Harbor,  Cataling  Harbor,  Avalon,  Santa  Cruz,  and  La  Jolla,  California). 
Clemens  and  Wilby,  1946:  168  (description  of  type). 

Gobius  nicholsi.  -Jordan  and  Gilbert,  1882:  946  (coast  of  British  Columbia;  description).  Jordan  and  Eigen- 
mann,  1886:  489,  494,  516,  517  (coast  of  British  Columbia;  analysis;  listed).  Jordan,  1885:  893  (105) 
(listed).  Eigenmann  and  Eigenmann,  1888:  59  (California;  listed).  Eigenmann  and  Eigenmann,  1892: 
354  (San  Diego,  California).  Eigenmann,  1892:  130,  159  (Point  Loma,  California).  Jordan  and  Starks, 
1895:  838  (Vancouver  Island,  British  Columbia;  listed).  Jordan  and  Evermann,  1896:  456  (coast  of 
British  Columbia;  listed).  Gilbert  and  Starks,  1904:  176  (mentions  Gobius  nicholsi  in  comparing  dermal 
fold  of  Microgobius  emblematicus).  Starks,  1911:  211  (listed;  San  Juan  Islands,  Washington).  Bean  and 
Weed,  1919:  79  (3  specimens  -  33,  43,  and  47  mm  long;  taken  at  Ucluelet,  Vancouver  Island,  British 
Columbia,  during  low  tide;  June-July,  1909). 

Gobius  nicholsoni  (sic). -Eigenmann,  1890:  66  (taken  in  deep  water  by  the  Albatross  off  Point  Loma,  Cali- 
fornia).  Eigenmann,  1909:  65  (off  Point  Loma,  California). 

Rhinogobius  nicholsii. -Starks  and  Morris,  1907:  223  (San  Pedro,  California).  Starks  and  Mann,  1911:  16 
(San  Diego,  California;  50  fm.). 

Rhinogobius  nicholsi. -Snyder,  1913:  459  (Pacific  Grove,  Cahfornia;  description;  taken  from  10-15  fm.). 
Gilbert,  1915:  359  (abundant  in  harbor  at  Avalon,  Catalina  Island;  taken  in  shallow  water  at  Monterey, 
California).  Kincaid,  1919:  40  (San  Juan  Islands,  Washington). 

Rhinogobiops  nicholsii. -Huhhs,  1926:  2  (type  of  genus;  description;  Santa  Barbara  Channel  and  southwest 
of  Newport,  California).  Hubbs,  1928:  15  (listed).  Ulrey  and  Greeley,  1928:  20  (Catalina  Island,  Hunt- 
ington Beach,  Malibu,  Newport,  Point  Firmin,  and  Santa  Cruz,  Cahfornia).  Jordan,  Evermann,  and  Clark, 
1928:  440  (coast  of  British  Columbia,  south  to  southern  California).  Ulrey,  1929:  10  (listed).  Wismer 
and  Swanson,  1935:  34 3;  Table  19  (San  Juan  Channel,  Washington;  depth  8-1 2m;  estimate  of  numbers  of 
R.  nicholsii  at  17  fish/2000m2  on  dredge  and  trawl  catches).  Schultz,  1936:  122,  191;  fig.  16  (key;  figure 
showing  ventral  side;  British  Columbia  to  southern  California).  Barnhart,  1936:  81;  fig.  245  (description; 
San  Clemente  Island  to  British  Columbia;  sometimes  taken  in  more  than  2100  feet).  Schultz  and  DeLacy, 
1936:  137,  213  (British  Columbia  to  southern  Cahfornia;  Hood's  Canal  near  Holly,  Washington;  marine; 
not  rare;  San  Juan  Island,  Washington).  Clemens  and  Wilby,  1946:  5,  29,  167-168;  fig.  103  (key;hsted; 
range,  English  and  Nanoose  bays,  Barkley  Sound  at  Ucluelet,  Esperanza  Inlet  on  west  coast  of  Vancouver 
Island,  Skidegate  Channel,  Queen  Charlotte  Islands;  20  fm.  or  more;  description).  Limbaugh,  1962:  552 
(La  Jolla,  California;  colonizing  newly  exposed  reefs). 

Coryphoptems  nicholsii. -Ginshurg,  1938:  113  (no  locahty;  differences  from  Coryphoptems  urospilus). 
Ginsburg,  1945:  136,  137  (C  nicholsii  used  in  study  of  fin-ray  count  methodology).  McAllister,  I960: 
38  (listed).  Miller  and  Lea,  1972:  186  (figure;  description;  key;  range  -  south  of  Point  Rompiente,  Baja 
California,  to  Skidegate  Channel,  Queen  Charlotte  Island,  British  Columbia;  depth  5  to  80  feet). 

Coryphoptems  nicholsi. -Huhhs  and  Follett,  1953:  34  (listed).  Limbaugh,  1955:  26,  35,  120  (southern  Cali- 
fornia; observed  in  "sand-bottom  holdfast  biotope"  and  "kelp  rock-bottom  biotope";  preferred  southern 
Cahfornia  habitat  sand  near  rocks;  1-180  feet;  description;  observed  at  following  localities  -  Pacific  Grove, 
Yankee  Point,  Goleta,  Point  Dume,  EI  Segundo,  Rocky  Cove,  Newport  Beach,  San  Clemente,  and  La 
Jolla;  and  at  San  Miguel,  Santa  Rosa,  Santa  Cruz,  Anacapa,  Santa  Catalina,  Los  Coronados,  and  San 
Martin  islands).  Bohlke  and  Robins,  I960:  103,  105  (key;  characteristics;  discussion  of  genus).  Ebert  and 
Turner,  1962:  249-252  (ecology;  breeding  habits;  behavior;  description  of  eggs  and  embryos).  Pequegnat, 
1964:  272  (Corona  del  Mar,  California;  abundance  on  reef).  Carlisle,  Turner  and  Ebert,  1964:  15,  28,  73, 
77  (listed;  observed  on  artificial  reefs  and  offshore  oil  installations,  and  in  Santa  Monica  Bay,  California; 
spawned  on  Redondo  Beach,  California  artificial  reef;  listed  -  Seal  Beach,  Rincon,  Summerland,  Redondo 
Beach,  and  Paradise  Cove,  California).  Best  and  Oliphant,  1965:  101  (listed;  Point  Arguello,  California). 
Turner,  Ebert  and  Given,  1965:  109,  112  (listed,  San  Elijo  Lagoon,  San  Diego  County,  California).  Berry 
and  Perkins,  1966:  676  (distribution  of  pelagic  prejuveniles).  Turner,  Ebert  and  Given,  1966a:  16,  17,  18, 
19,  26-27,  29;  tables  1-4  (abundance;  distribution  on  benthic  quadrats;  listed  as  abundant;  Point  Loma, 
California).  Turner,  Ebert  and  Givfen,  1966b:  40,  47  (distribution  around  Orange  Co.,  California  sewer 
outfall  pipeline;  relative  abundance  of  goby  in  vicinity  of  outfall).  McCart,  1967:  433-434  (scale  regener- 
ation). Fitch,  1967:  4,  16;  fig.  3  (lower  Pleistocene  otoliths).  Fitch,  1968:  2,  21-22;  fig.  2s  (early  Pleis- 
tocene otoliths).  Turner,  Ebert  and  Given,  1969:  185  (habits  on  southern  California  artificial  reefs). 
Macdonald,  1972:  91  (cephalic-lateralis  system).  Quast,  no  date  (a):  4,  6;  table  1  (listed  among  species 
ranging  from  boreal  into  temperate  waters;  British  Columbia  through  north  temperate;  0-130  feet;  listed 
as  member  of  southern  California  rocky-inshore  zone  fauna). 


189 


Bohlke  and  Robins  (1%0)  referred  to  this  species  as  Coryphopterus  nicholsi.  Although  it 
stands  alone  in  the  group,  Bohlke  and  Robins  accept  it  in  Coryphopterus.  Tliey  indicated  that 
should  C.  nicholsii  be  treated  as  subgenerically  distinct,  the  name  Rhinogobiops  Hubbs  would 
apply. 


Figure  1.  Coryphopterus  nicholsii.   Adult  male,  86.5  mm  in  standard  length,  from  Laguna  Beach,  Orange  Co., 
California. 

Diagnosis. -Coryphopterus  nicholsii  (Fig.  1)  is  easily  distinguished  from  all  other  species 
of  Coryphopterus  by  having  more  soft  dorsal,  anal,  and  pectoral  rays,  more  scales,  and  in  having 
a  narrow  wedge  of  scales  reaching  a  point  above  the  anterior  margin  of  the  opercle. 

Description. -h  has  an  elongate,  moderately  stout  body.  Greatest  body  depth  4.1-6.1 
(5.17)  in  standard  length.  Body  slightly  compressed,  width  5.7-10.6  (7.26)  in  standard  length. 
Head  moderate,  wider  than  deep;  head  width  1.2-2.1  (1.50)  in  head  length;  head  length  3.0-4.2 
(3.56)  in  standard  length.  Cheeks  not  tumid.  Mouth  small  and  terminal.  Lower  jaw  projecting. 
Maxilla  not  reaching  to  point  below  anterior  margin  of  eye,  1.9-3.2  (2.58)  in  head  length.  Jaw 
teeth  conical,  in  bands,  enlarged  in  both  an  outer  and  inner  row.  Tongue  truncate  at  tip.  Eye 
directed  superolaterally  and  large  (diameter  2.5-4.3  (3.42)  in  head  length).  Bony  interorbital 
very  narrow.  A  high,  thin  and  nearly  vertical  crest  on  top  of  head  from  behind  eyes  to  origin  of 
spinous  dorsal  fin.  Rows  of  papillae  on  sides  of  head  moderately  developed.  No  barbels  (Fig. 
12).  Slit  behind  fourth  gill-arch  reduced;  pseudobranchiae  exposed.  Branchiostegals  5  (1  on 
epihyal).  Pelvic  fins  fully  united,  free  from  belly,  each  with  1,4  rays.  Pectoral  fins  with  21-24 
(22.3)  rays,  none  silky.  Dorsal  fins  barely  separated,  VI-I,  12-15  (13.8).  Caudal  fin  broadly 
rounded,  17:  12  segmented,  branched  rays  and  a  variable  number  of  segmented,  unbranched 
rays  and  simple  (procurrent)  rays.  Some  specimens  with  2  above  and  1  below  segmented,  un- 
branched rays,  plus  2  simple  rays  below  occur  about  as  frequently  as  those  with  2  above  and 
none  below  segmented,  unbranched  rays,  plus  3  simple  rays  —  1  above,  2  below.  Rarely  the 
caudal  elements  composed  of  1 2  branched,  segmented  rays  with  above  and  below  1  unbranched, 
segmented  ray,  plus  1  above  and  2  below  simple  rays.  Anal  1,  1 1-14  (12.1).  Body  completely 
scaled  except  in  predorsal  midline.  Head  scaleless.  Scales  of  sides  each  with  a  comb-like  row  of 
marginal  spines,  a  submarginal  focus,  and  basal  radii.  Scales  in  oblique  rows,  23-28  (25.6)  at 
midline.  No  lateral  line.  Shoulder  girdle  without  papillae.  Color:  pale  orange-olive  or  light 
yellow,  with  irregular  vertical  purplish  brown  streaks  developed  at  time  of  death  or  in  social 
interaction.  Body  irregularly  flecked  with  metallic  blue-green.  Iridescent  stripe  below  eye, 
giving  rise  to  vernacular  name  -  bluespot  goby.  Tip  of  first  dorsal  jet  black.  Pelvic  fin  of 
breeding  male  black. 


OCCURRENCE 

Coryphopterus  nicholsii  ranges  from  Point  Rompiente  (27°N),  Baja  California  to  Skide- 
gate  Channel  (530N),  Queen  Charlotte  Island,  British  Columbia  (Miller  and  Lea,  1972;McCart, 
1967).   It  is  common  in  this  area  and  its  range  may  be  more  extensive  than  is  currently  reahzed. 


190 

Otoliths  of  C.  uichohii  have  been  found  in  many  southern  CaHfornia  Pliocene  and  Pleis- 
tocene deposits.  More  than  1 ,700  have  been  recovered  from  the  Lomita  marl  (Pliocene)  at  San 
Pedro,  and  others  have  come  from  Timms  Point  silt,  San  Pedro  sand  (Lower  I^eistocene)  and 
Baldwin  Hills,  Los  Angeles  (Upper  Pleistocene)  (Fitch,  1967,  1968). 

GENERAL  ECOLOGY  AND  NATURAL  HISTORY 

Coryphopterus  nicholsii  inhabits  shallow  water,  from  6m  to  more  than  60m.  After  a 
pelagic  oceanic  existence  as  larvae  and  early  juveniles,  individuals  assume  a  benthic  habit.  The 
preferred  habitat  appears  to  be  a  rock-reef  area,  but  small  groups  of  rocks  on  the  open  sand  are 
also  inhabited.  The  greatest  concentrations  occur  near  the  periphery  and  in  the  channels  of  the 
reef,  where  the  rock  meets  the  surrounding  sand  bottom;  in  these  areas  there  are  numerous 
undercuts  and  holes,  into  which  this  goby  can  find  safety.  It  is  seldom  found  more  than  25  cm 
from  protective  cover,  but  is  common  on  the  open  sand  or  rubble  bottom  in  front  of  its  shelter. 
The  open  area  around  the  hole  serves  as  a  feeding  and  display  site. 

Coryphoptenis  nicholsii  is  also  found  over  the  tops  of  reefs,  particularly  in  areas  having 
many  holes,  ledges,  or  thick  gorgonian  cover.  Using  the  protective  cover  of  the  gorgonian 
canopy,  individuals  may  venture  more  than  a  meter  from  their  home  shelter. 

MORPHOMETRICS 

The  meristics  of  C.  nicholsii  from  the  sampled  areas  are  rather  uniform  (Table  1 ).  How- 
ever, pectoral  ray  counts  from  British  Columbia  are  lower  than  those  from  the  southernmost 
locality  (Isla  San  Martin). 

Pelagic  prejuveniles  were  sampled  in  southern  California  from  San  Pedro  Basin,  San  Juan 
Seamount,  and  between  Anacapa  and  Santa  Cruz  Islands.  Variation  in  the  pelagic  prejuveniles 
was  similar  to  that  in  the  juvenile-adult  sample  with  the  exception  of  the  first  dorsal  spines  and 
scale  counts.  For  the  scales  the  explanation  is  one  of  delayed  development.  Variation  in  dorsal 
spines  between  the  age  groups  may  be  the  result  of  delayed  development  or  perhaps  some  spines 
were  overlooked. 

Morphological  data  for  three  localities  were  considered  in  size  groups  in  Table  2.  Although 
adequate  samples  were  not  available  from  Baja  California  or  British  Columbia,  no  notable  trend 
of  morphological  variation  was  evident.  Differences  in  proportions  were  observed  in  fin  ray 
lengths.  As  C.  nicholsii  exhibits  sexual  dimorphism  of  some  body  parts  (particularly  fins) 
morphological  comparisons  should  be  made  on  the  basis  of  individual  sex,  but  sample  sizes 
were  inadequate  for  such  analyses. 

FOOD  HABITS 

Food  studies  were  conducted  to  identify  the  major  foods,  to  determine  if  differences  in 
seasonal  utilization  occurred,  and  to  disclose  any  variation  in  the  foods  taken  by  fish  of 
different  sizes.  Tliree  methods  of  analysis  were  used:  numerical,  volumetric,  and  frequency-of- 
occurrence.  Of  the  106  stomachs  examined,  two  were  empty,  and  were  not  included  in  the 
calculations. 

It  appears  that  crustaceans  are  the  principal  food  item  of  C.  nicholsii  (Fig.  2);  they  were 
found  in  nearly  all  stomachs  examined,  generally  in  the  greatest  numbers,  and  formed  the  bulk 
of  the  volume.  Numerically  crustaceans  comprised  90.9%  of  the  food  items;  mollusks  (4.8%) 
were  second.  Other  food  items,  such  as  annelids,  echinoderms,  etc.,  were  usually  found  in  small 
numbers. 

By  volume  crustaceans  made  up  79.0%  of  the  diet;  mollusks  again  were  second  (4.6%). 
Other  foods  composed  a  small  portion  of  the  diet. 

Crustaceans  occurred  in  97.0%>  of  the  stomachs  analyzed  (Fig.  2).  Although  found  in  small 
quantities  numerically  and  volumctrically,  mollusks  occurred  in  63%  of  stomachs  examined. 
Bryozoans,  which  accounted  for  only  0.4%  of  total  food  volume,  were  found  in  37%  of 
stomachs  examined.  As  this  goby  was  not  observed  to  selectively  bite  off  pieces  of  bryozoans 
on  the  reefs  it  is  possible  that  these  organisms  were  picked  up  incidentally  along  with  the  pre- 
ferred bottom-dwelling  food  items.  Bryozoans  are  relatively  indigestible  and  are  probably  re- 
tained in  the  gut  for  some  time,  which  may  account  for  their  apparent  abundance. 


191 


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Table  2.   Comparison  of  morphometries  among  populations  of  Coryphopterus  nicholsii 


Punta  Banda, 
Baja  California 


Laguna  Beach, 
California 


Hunt  Island, 
British  Columbia 


Proportions 


S.L.         N     Range      Mean  N     Range     Mean  N     Range     Mean 


standard  length 


head  length 

22-39mm 

11 

3.0-3.4 

(3.30) 

39 

3.0-3.6 

(3.30) 

40-55 

17 

3.1-3.8 

(3.39) 

56 

3.1-3.7 

(3.40) 

56-75 

18 

3.2-4.0 

(3.60) 

86 

3.3-3.9 

(3.46) 

9 

3.6-3.9 

(3.75) 

76-92 

26 

3.3-4.2 

(3.65) 

17 

3.3-4.0 

(3.76) 

standard  length 

body  width 

22-39mm 

11 

5.7-8.4 

(7.46) 

37 

7.0-10.6 

(8.48) 

40-55 

9 

6.7-7.8 

(7.33) 

45 

6.1-8.7 

(7.25) 

56-75 

17 

6.6-8.1 

(7.35) 

51 

5.9-8.5 

(6.60) 

76-92 

28 

5.9-8.8 

(7.28) 

standard  length 

length  base  1st  dorsal 

22-39mm 

11 

5.7-6.4 

(6.17) 

39 

5.3-7.2 

(6.28) 

40-55 

14 

4.7-5.8 

(5.30) 

51 

4.9-7.1 

(5.67) 

56-75 

18 

5.1-6.1 

(5.59) 

75 

4.9-6.2 

(5.41) 

10 

5.0-6.0 

(5.50) 

76-92 

23 

4.6-6.1 

(5.20) 

17 

4.8-6.1 

(5.25) 

standard  length 

22-39mm 

11 

3.4-4.0 

(3.80) 

39 

3.4-4.1 

(3.82) 

length  base  2nd  dorsal 

40-55 

15 

3.4-3.8 

(3.58) 

54 

3.0-3.8 

(3.56) 

56-75 

18 

3.4-4.2 

(3.74) 

76 

3.3-3.9 

(3.51) 

10 

3.5-3.8 

(3.64) 

76-92 

23 

3.2-4.0 

(3.54) 

17 

3.3-4.2 

(3.67) 

standard  length 

length  base  anal 

22-39mm 

11 

3.9-4.9 

(4.69) 

38 

3.4-4.8 

(4.78) 

40-55 

13 

4.3-4.9 

(4.54) 

49 

3.9-5.1 

(4.61) 

56-75 

18 

4.4-4.8 

(4.56) 

79 

4.1-5.0 

(4.69) 

10 

4.4-4.8 

(4.56) 

76-92 

24 

4.0-4.9 

(4.52) 

17 

4.2-5.0 

(4.54) 

standard  length 

22-39mm 

11 

3.0-3.6 

(3.40) 

38 

3.1-3.9 

(3.51) 

length  of  longest  pectoral  ray 

40-55 

16 

3.3-3.9 

(3.46) 

52 

3.1-4.3 

(3.60) 

56-75 

18 

3.1-4.0 

(3.51) 

70 

3.1-4.0 

(3.69) 

3 

3.1-3.2 

(3.17) 

76-92 

21 

3.2-3.9 

(3.55) 

17 

2.8-3.5 

(3.22) 

standard  length 

22-39mm 

11 

2.4-2.8 

(2.53) 

28 

2.4-2.5 

(2.45) 

length  of  longest  2nd  dorsal  ray 

40-55 

16 

3.2-3.7 

(2.74) 

11 

2.1-2.5 

(2.32) 

56-75 

18 

2.0-2.2 

(2.15) 

34 

1.9-2.4 

(2.23) 

3 

2.0-2.1 

(2.03) 

76-92 

18 

1.9-2.3 

(2.10) 

17 

1.7-2.0 

(1.89) 

standard  length 

body  depth 

22-39mm 

11 

4.4-5.4 

(4.97) 

39 

4.5-5.9 

(5.18) 

40-55 

16 

4.5-5.5 

(4.98) 

50 

4.4-5.9 

(5.35) 

56-75 

18 

4.6-5.6 

(5.00) 

73 

4.1-6.1 

(5.05) 

3 

4.9-5.3 

(5.10) 

76-92 

23 

4.5-6.1 

(5.27) 

17 

4.8-5.9 

(5.39) 

standard  length 

22-39mm 

11 

4.0-4.4 

(4.21) 

39 

4.1-4.7 

(4.40) 

length  of  longest  pelvic  ray 

40-55 

3 

4.3-4.5 

(4.37) 

11 

4.1-5.1 

(4.48) 

56-75 

4 

3.9-4.6 

(4.38) 

28 

3.5-5.1 

(4.55) 

3 

4.2-4.8 

(4.44) 

76-92 

19 

4.3-4.8 

(4.66) 

17 

3.9-4.5 

(4.25) 

standard  length 

22-39mm 

3 

2.8-3.4 

(3.03) 

35 

2.9-3.0 

(2.97) 

length  of  longest  anal  ray 

40-55 

3 

2.6-2.8 

(2.74) 

12 

2.7-3.4 

(2.89) 

56-75 

4 

2.6-2.8 

(2.75) 

28 

2.4-3.0 

(2.74) 

3 

2.5-2.7 

(2.56) 

76-92 

17 

2.3-2.9 

(2.56) 

16 

2.1-2.7 

(2.18) 

head  length 

22-39mm 

4 

2.5-3.2 

(2.65) 

35 

2.4-3.2 

(2.82) 

length  upper  jaw 

40-55 

16 

2.3-3.1 

(2.66) 

56 

2.3-3.2 

(2.86) 

56-75 

7 

2.4-2.9 

(2.64) 

77 

2.1-3.2 

(2.55) 

3 

1.9-2.8 

(2.40) 

76-92 

22 

2.2-2.7 

(2.44) 

16 

2.3-2.7 

(2.45) 

head  length 

head  width 

22-39mm 

4 

1.6-1.7 

(1.62) 

36 

1.5-2.1 

(1.75) 

40-55 

16 

1.3-1.7 

(1.49) 

59 

1.4-1.8 

(1.51) 

56-75 

7 

1.3-1.5 

(1.43) 

77 

1.2-1.8 

(1.42) 

3 

1.4-1.5 

(1.43) 

76-92 

24 

1.2-1.6 

(1.37) 

17 

1.3-1.7 

(1.54) 

head  length 

snout  length 

22.39mm 

4 

3.5-4.9 

(4.06) 

36 

3.3-4.5 

(3.84) 

40-55 

15 

3.2-4.0 

(3.51) 

56 

3.2-3.9 

(3.42) 

56-75 

7 

3.1-3.9 

(3.34) 

78 

2.9-4.0 

(3.57) 

3 

2.9-4.5 

(3.63) 

76-92 

22 

3.1-4.0 

(3.45) 

17 

3.5-4.4 

(3.99) 

head  length 

length  of  eye 

22-39mm 

4 

3.2-4.3 

(3.52) 

35 

3.1-3.8 

(3.30) 

40-55 

15 

2.9-3.8 

(3.04) 

54 

2.9-3.9 

(3.36) 

56-75 

7 

3.1-3.9 

(3.46) 

78 

2.9-4.2 

(3.52) 

2.5-3.3 

(2.97) 

76-92 

24 

3.3-4.0 

(3.61) 

17 

3.1-3.6 

(3.45) 

193 


Echinoderms  (mainly  sea  urchin  spines  and  tests)  were  also  found  in  relatively  high  fre- 
quency (26%),  probably  for  the  reasons  discussed  for  bryozoans.  The  nutritional  importance  of 
bryozoans  and  echinoderms  is  seemingly  small. 


CRUSTACEANS 


MOLLUSCS 


ANNELIDS 


UNIDENTIFIED 
EGGS 


ECHINODERMS 


BRYOZOANS 


Si9^^$^s$C!$S!$?s$s^s^$s^ff^^ 


i 


k 


^ 


[ 


S  NUMERICAL  7o 
■  VOLUMETRIC  % 
n  OCCURRENCE  % 


— r— 
25 


50 


— r- 
75 


100 


Figure  2.    Analysis  of  106  stomach  contents  of  Coryphoptems  nicholsii  taken  at  Laguna  Beach,  California, 
from  October  1966  to  September  1967. 


CRUSTACEANS 
COPEPODS 
ISOPOOS 
AMPHIPODS 
OSTRACODS 
DECAPODS 
CUMACEANS 
UNIDENTIFIED 

MOLLUSCS 
PELECYPODS 
GASTROPODS 

ANNELIDS 

ECHINODERMS 

PORIFERA 

BRYOZOANS 

COELENTERATES 

VERTEBRATES 

MISC.  EGGS 


Figure  3.    Further  analysis  of  the  stomach  contents  of  Coryphoptems  nicholsii  expressed  as  numerical  and 
frequency-of-occurrence  percentages. 


194 


Measurements  and  counts  from  the  moUusks  were  taken  with  the  shell  intact.  However, 
for  accurate  evaluation,  only  the  digestible  parts  should  be  considered.  This  was  not  possible, 
because  of  the  small  size  of  the  food  items.  Because  mollusk  shells  are  relatively  undigestible, 
and  they  may  be  retained  in  the  stomach,  data  for  this  element  are  probably  biased.  Some 
shells  may  have  been  picked  up  incidentally  along  with  the  substrate  as  the  goby  grabbed  for  a 
desired  bottom-dwelling  organism.  Empty  gastropod  shells  could  also  be  the  shelter  of  hermit 
crabs,  an  important  part  of  the  decapod  element  of  the  diet.  The  crabs  are  presumably  digested 
rapidly,  whereas  the  shells  may  accumulate  in  the  gut.  These  emptied  hermit  crab  shelters 
would  then  be  categorized  as  a  mollusk  element  even  though  the  food  item  selected  by  the  goby 
was  a  crustacean. 

Amphipods  (39.9%  numerically)  and  copepods  (19.6%  numerically)  were  the  most  abun- 
dant crustaceans  taken  by  C.  nicholsii  (Fig.  3).  One  stomach  contained  256  amphipods.  Iso- 
pods  (3.6%)  and  decapods  (6.8%)  also  made  up  an  important  part  of  the  diet. 

Amphipods  were  found  in  88%  of  the  stomachs  examined.  Copepods  (74%),  decapods 
(73%)  and  isopods  (55%)  were  also  found  in  most  stomachs  (Fig.  3).  Although  pelecypods  and 
gastropods  composed  only  2.5%  and  2.3%  of  the  total  number  of  food  items  respectively,  pele- 
cypods were  found  in  49%  and  gastropods  in  47%  of  all  stomachs  examined  (Fig.  3). 


CRUSTACEANS 

.SSQS8S!3< 

KiSSS^SSS 

>^^^\^^\\^\^^l 

■■■■■■1 

MOLLUSCS 

ANNELIDS 

UNIDENTIFIED 
EGGS 

ECHINODERMS 
BRYOZOANS 

1 

1 

k- 

] 

S  NUMERICAL  % 

UJ 
QQ 

C9 

^^ 

■  VOLUMETRIC  % 
n  OCCURRENCE  % 

y 

wm 

c 

Sv:::. 

( 

25 

50                            75 

10 

CRUSTACEANS 

MOLLUSCS 

UNIDENTIFIED 
EGGS 

BRYOZOANS 

K^«S?Nfi?SISS«J2«5?je^;SS»5^^ 

OS 

i^:::;;:iii;;;;;;:;:i;h;;::;;  ■                                  j 

ea 

UJ 

F 

UJ 

:;:::S---::i 

CO 

1 

^ 

1 

a. 

? 

c 

1 

25 

1                         1 

50                              T> 

U)l 

Figure  4.  Seasonal  analxsis  of  the  stomach  contents  of  Coryphopterus  nicholsii. 


Coryphoptenis  nicholsii  exhibited  seasonal  variation  in  food  utilization  (Fig.  4).  Crus- 
taceans were  the  major  food  item  in  both  the  October-March  sample  (numerical:  87. 7%; volu- 
metric: 66.0%)  and  the  April-September  sample  (numerical:  94.0%;  volumetric:  95.5%). 
Mollusks  apparently  became  more  important  in  October-March  (numerical:  4.5%;  volumetric: 
4.6%)  as  compared  to  April-September  (numerical:  5.3%;  volumetric:  1.5%).  The  other  food 
classes,  e.g.,  echinoderms,annelids,'etc.,  also  became  more  important  during  the  Fall  and  Winter. 


195 


The  frequency  analysis  also  showed  a  similar  pattern  of  seasonal  food  use  (Fig.  4).  Crus- 
taceans were  found  in  86%  of  the  October-March  stomachs  and  98%  of  the  April-September 
stomachs.  Mollusks  were  found  in  70%  of  the  October-March  samples,  but  in  only  58%  of  the 
April-September  samples.  Thirty-two  percent  of  the  October-March  stomachs  contained  echino- 
derms,  but  only  18%  of  the  April-September  stomachs  contained  this  food. 

Benthic  organisms,  such  as  mollusks,  echinoderms,  annelids  and  bryozoans,  were  found 
more  frequently  in  the  October-March  period.  Also  in  the  October-March  sample  the  crustacean 
element  shifted  somewhat  to  more  benthic  forms  such  as  decapods,  cumaceans,  and  ostracods, 
with  concomitant  decreases  in  copepods  and  other  swimming  and  planktonic  forms.  Tliis  shift 
in  diet  perhaps  reflected  changes  in  populations  of  the  latter  groups. 

Copepods  composed  100%  (N  =  47)  of  the  food  of  four  pelagic  postlarvae  (18.7-21.1  mm) 
from  the  San  Juan  Seamount,  California.  One  stomach  contained  a  single  scale.  No  gross  dif- 
ferences in  diet  were  noted  between  size  classes  or  sexes  once  the  gobies  had  assumed  a  benthic 
existence. 

POPULATION  STRUCTURE 

Tlie  Laguna  population  showed  a  sex  ratio  of  1.7  females  to  1  male  (165  females  and  96 
males).  The  sex  ratio  seemingly  fluctuated  somewhat,  but  the  paucity  of  data  for  some  months 
precluded  a  complete  analysis. 

Scales  provided  satisfactory  material  for  aging  C.  nichohii,  and  scale  analysis  demonstrated 
the  relation  between  age  and  size  (Fig.  5).  Modes  in  the  length-frequency  distributions  agree 
well  with  the  observed  ages,  although  overlap  occurs  due  to  the  differential  growth  and  pro- 
longed spawning  period. 


V- 


IV- 


«  III- 

UJ 

C9 


II- 


I- 


STANDARD  LENGTH,  MM 


Figure  5.  Empirical  growth  rate  of  Coryphopterus  nicholsii  from  Laguna  Beach,  California,  based  on  scale 
analysis  of  134  specimens  collected  from  October  1966  through  January  1968.  Vertical  line  represents  mean; 
horizontal  line,  the  range  of  variation;  longer  rectangle,  one  standard  deviation  on  either  side  of  the  mean; 
shorter  rectangle,  2  standard  errors  on  either  side  of  the  mean. 


196 


Young  gobies  were  first  observed  on  the  Laguna  reefs  in  February.  Tliese  individuals, 
which  ranged  from  21.8  to  26.2  mm  in  standard  length,  were  probably  ones  that  had  hatched 
the  previous  year.  Growth  during  the  pelagic  period  is  considerable  as  the  newly-hatched  larvae 
measure  under  3.0  mm  in  total  length. 

Scales  of  C.  nicholsii  form  during  the  pelagic  period.  One  specimen  showed  some  well 
developed  scales  at  19.5  mm  on  28  December  1966.  However,  the  majority  of  specimens  did 
not  show  scales  until  they  had  attained  a  length  of  21 .1  mm  or  greater.  This  is  the  approximate 
size  at  which  C.  nicholsii  settled  on  the  reefs  in  1967.  Tlie  first  annulus  is  not  laid  down  until 
the  following  winter. 

Tlie  observed  length-frequencies  of  C  nicholsii  coWeciQd.  at  Laguna  Beach  (Fig.  6)  usually 
corresponded  with  the  observed  age-groups  indicated  by  scales  in  that  five  (sometimes  four) 
frequency  groups  were  represented  in  the  samples.  Males  of  any  frequency  group  were  rela- 
tively larger  (as  indicated  by  mean  standard  length)  than  females  of  the  same  group. 


r^1n  II       r^rlnrn     n  1^  n 


JAN. 


r^    nn   n    n  h 


EL 


^^^ 


IL 


FEB. 


AFInnnn       nr^       A       nP-n     P^^^  h  I ^n       F^  n 'f\ 


MAR. 


n^n  n^^r^rnnTiJi^  n./\ 


JUNE 

_L1 


pJl^r^n    n^^hn    h^nn 


RJl 


JULY 


f^npJI     . 


IL 


rXnrnnr^^ 


AUG. 


5- 


0 
5 
0 
10 
5H 


Ik 


h 


JhL 


SEPT. 


A 


^ 


OCT. 


^nnnn   jl 


^   ySF^ 


^ 


j^ 


NOV. 


X 


I 


r^ ,  n  K^^^^  r^  .  /h 


DEC. 


20  30  40  50  60  70 

STANDARD  LENGTH.  MM 


80 


90 


Figure  6.     Length-frequency  measureihents  of  1325  specimens  of  Coryphopterus  nicholsii  taken  at  Laguna 
Beach,  Cahfornia,  from  October  1966  through  September  1967. 


PREDATION 

One  incident  of  predation  on  C.  nicholsii  was  observed  on  the  shallow  Laguna  reef.  A 
small  sand  bass  {Paralahrax  nebulifer)  caught  and  ate  a  Coryphopterus  after  I  flushed  the  goby 
away  from  the  shelter  of  the  reef  onto  the  surrounding  open  sand.    Subsequent  collections  of 


197 


22  sand  bass  on  the  reef  revealed  that  7  (32%)  had  fed  on  C.  nichohii.  Turner,  Ebert  and  Given 
(1969)  did  not  find  C.  nichohii  in  the  stomach  of  large  predaceous  fish  collected  on  reefs  be- 
tween 1960  and  1963,  although  this  goby  was  abundant  and  appeared  to  be  a  suitable  food 
item.  Smith  (1970)  and  Quasi  (n.d.b.)  list  Gobiidae  as  a  food  of  kelp  bass  (Paralabrax 
clathratus). 

Stomach  analyses  of  bluebanded  gobies,  Lythrypnus  dalli,  another  common  inhabitant  of 
the  southern  California  rock  reef,  revealed  larval  C.  nicholsii  in  2  of  42  (4.8%)  stomachs. 
Investigations  were  not  made  on  the  food  habits  of  other  possible  predators  on  the  reefs 
inliabited  by  C.  nichohii. 

Turner,  Ebert  and  Given  (1969)  observed  a  3  mm  larval  C.  nichohii  entrapped  in  the 
hydranth  of  an  Obelia  on  an  artificial  reef  off  southern  Cahfornia. 

SEXUAL  DIMORPHISM 

Sexual  dimorphism  of  the  genital  papilla,  is  usually  evident  in  the  Gobiidae  (Dotu,  1957a, 
1958a,  1961a;  Miller,  1963;  Springer  and  McErlean,  1961 ;  Tavolga,  1954;  Smith,  1964;Weisel, 
1947).  Dimorphism  has  also  been  noted  in  fin  size  (Hildebrand  and  Cable,  1938;  Baird,  1965; 
Dotu,  1958b,  1959,  1961b),  pigmentation  (Heincke,  1880;  Tavolga,  1954;Ninni,  1938;D6tu, 
1958c,  1961c),  shape  of  mouth  (Baird,  1965),  and  size  and  shape  of  the  body  (Breder  and 
Rosen,  1966;  Baird,  1965;  Dotu,  1957b,  1958c,  1961c). 

Four  hundred  specimens  of  C.  nicholsii  from  9.6  to  88.0  mm  in  standard  length  were 
examined  for  external  sex  identification.  The  genital  papilla  was  found  to  be  sexually  dimorphic 
(Fig.  7).  Specimens  shorter  than  25  mm  standard  length  could  not  be  accurately  sexed  extern- 
ally, but  those  larger  than  25  mm  were  correctly  sexed  by  examination  of  the  papilla,  which 
is  elongate  and  pointed  in  the  male  and  is  broad  and  truncate  in  the  female. 

During  the  breeding  season  the  males  of  C.  nichohii  are  easily  distinguished  from  the 
females  by  their  black  pelvic  fins.  These  fins  remain  light  grey  throughout  the  year  in  the 
females. 


83.0  mm.  S.L. 


Gl.Omm.S.L. 


Figure  7.  Sexual  dimorphism  in  the  genital  papilla  of  Coryphoptems  nicholsii. 

Dimorphism  is  also  discernable  in  the  maximum  sizes  of  the  sexes.  Males  attain  a  greater 
length  than  females.  The  largest  male  examined  was  90.0  mm  in  standard  length,  whereas  the 
largest  female  was  76.0  mm.  Larger  size  may  accord  the  nest-guarding  male  greater  success  in 
the  protection  of  the  eggs. 

Examinafion  of  the  length  of  the  second  dorsal  and  anal  fins  of  mature  gobies  also  re- 
vealed sexual  dimorphism.  Both  fins  were  found  to  be  relatively  longer  in  the  male  (Fig.  8). 
Measurement  was  from  the  front  of  the  base  of  the  first  element  to  the  distal  end  of  the  last 
ray.  Tliis  dimorphism  was  not  expressed  in  meristic  differences. 


198 


REPRODUCTION 

The  Laguna  population  was  sampled  at  weekly  or  biweekly  intervals  from  October  1966 
to  September  1967  to  determine  the  length  of  the  breeding  season.  The  presence  of  eggs  and 
individuals  of  both  sexes  in  breeding  condition  was  used  as  evidence  of  reproductive  activity. 


CO 


2.0        2.1         2.2       2.3        2.4 

2nd  Dorsal  in  Standard  Length 


Oct 
09 


23       24        25       2.6       2.7       2.8 

Anal  Fin  in  Standard  Length 


Figure  8.    Sexual  dimorphism  in  length  of  the  second  dorsal  and  anal  fins  of  Coryphoptenis  nicholsii,  from 
anterior  of  base  to  posterior  tip  of  last  ray;  expressed  as  ratio  of  length  of  fin  into  standard  length. 

Females  of  C.  nicholsii  were  found  to  be  mature  at  47.3  mm  or  larger  (Fig.  9).  This  size 
corresponds  to  age-group  11  as  observed  from  scale  and  length-frequency  analysis.  Mature  males 
shorter  than  55  mm  were  not  found.  This  size  corresponds  with  the  last  of  age-group  III.  How- 
ever, mature  males  were  found  with  two  or  more  annuli  (age-groups  III  througli  V).  Ripe 
females  were  found  in  age-groups  II  througli  V. 


o 
a> 


4- 
3- 
2 
1 


0 


cr 


• — Mature 
*— Immature 


• — •• 


• — • — •' 


— I    '  I      I      I      I 

20  30  40 


7 


I    ■  '    I 
50 


— p— 
60 


70 


80 


Standard  Length  in  Millimeters 


Figure  9.     Standard  lengths  of  Coryphoptenis  nicholsii  at  sexual  maturity  based  on  gonad  development  of 
22  males  and  43  females  from  Laguna  Bcach,  California. 


199 


Tlie  first  ripe  females,  measuring  47.3  to  73.5  mm  in  standard  length,  were  taken  on 
10  February  1967.  Tlie  first  ripe  males  appeared  slightly  earlier  on  2  February  1967;  they 
ranged  from  72.1  to  83.0  mm  in  standard  length.  No  ripe  gobies  of  either  sex  were  found  after 
26  August  1967  (Fig.  10). 


c»o»-«  ©•o»o»- o»  o  w^  o  •  o  • 


'  Mar.  '  Apr.  '  May  '  June  '  July  '  Aug  '  Sep. '  Oct  '  Nov  '  Dec  ' 


Jan    '  Feb 


MONTH 


Figure  10.  Monthly  percentages  of  ripe  Coryphoptents  nicholsii  from  Laguna  Beach,  CaHfornia.  Based  on 
1134  specimens  taken  from  Laguna  Beach,  California  from  October  1966  through  September  1967.  Solid 
dots  (solid  line)  represent  males,  circles  (broken  line)  represent  females. 


The  mature  ovaries  contained  two  egg  groups,  one  ripe  and  one  unripe.  The  ripe  egg 
group  seemed  to  be  spawned  at  one  time.  The  number  of  ripe  ovarian  eggs  (in  both  ovaries) 
was  found  in  four  individuals  to  range  from  3274  to  4788.  However,  the  assessment  of 
fecundity  in  a  species  which  may  spawn  several  times  over  an  extended  period,  has  little  bio- 
logical value  per  se.    I  could  not  determine  how  many  times  this  species  spawned  in  a  season. 

Ripe  ovarian  eggs  are  orange  and  round,  and  measure  0.4  to  0.7  mm.  Unripe  eggs  range 
in  diameter  from  0.05  to  0.2  mm. 

Eggs  were  found  on  the  Laguna  reefs  from  1 1  April  to  5  August  1967.  However,  because 
it  is  difficult  to  locate  the  nests,  I  do  not  think  that  these  dates  wholly  encompass  the  spawning 
period.  Ebert  and  Turner  (1962)  observed  nests  off  Hermosa  Beach  and  Santa  Monica,  Cali- 
fornia, from  April  through  October. 


200 


Fertilized  eggs  (Fig.  1 1)  have  the  spindle  shape  characteristic  of  gobies.  They  are  attached 
directly  to  the  rock  surface  but  have  no  adhesive  threads.  Ebert  and  Turner  (1962)  found  that 
mature  eggs  averaged  2.10  mm  long  by  0.48  mm  wide.  The  embryo  within  eqph  mature  egg 
averages  2.97  mm  in  length  and  its  head  is  directed  opposite  (downward  from)  the  pole  of 
attachment  on  the  lower  surface  of  the  nest.  Ebert  and  Turner  described  the  developing 
embryo. 


Fertilized  Egg  Mass 
Section 


A.. 

Fertilized  Egg 


2.2  mm. 


Figure  1 1 .  Fertilized  eggs  of  Coryphopterus  nicholsii. 

Tlie  prejuvenile  of  C.  nicholsii  (Fig.  12)  is  pelagic.  Specimens  ranging  in  size  from  15.5  to 
29.0  mm  standard  length  have  been  taken  560  km  off  San  Francisco  and  260  km  off  Santa 
Barbara.  One  individual  was  taken  on  Davidson  Seamount,  97  km  southwest  of  Point  Sur, 
California  (Berry  and  Perkins,  1966).  These  individuals  have  been  described  by  Berry  and 
Perkins  as  pelagic,  oceanic,  protracted  prejuvenile  stages.  Specimens  from  the  vicinity  of  San 
Diego  ranging  from  9.6  to  22.6  mm  in  standard  length,  have  been  examined.  These  were  dis- 
tinct from  the  adults  in  having  vertical  bars  which  are  burnt  orange  in  life.  These  bars  become 
light  brown  in  alcohol. 

Juveniles  of  C.  nicholsii  were  found  to  assume  a  benthic  habit  on  the  Laguna  study  reefs 
at  21.8  mm  standard  length.  Specimens  as  short  as  24.0  mm  were  found  on  other  reefs.  Fish 
of  this  size  were  found  on  the  Laguna  reefs  in  February  through  August  1967. 


Figure  1  2.   Pelagic  prejuvenile  of  Coryphopterus  nicholsii,  1 9.4  mm  in  standard  length,  from  San  Pedro  Basin, 
California. 


201 

FEEDING  BEHAVIOR 

Coryphoptems  nicholsii  has  three  feeding-behavior  patterns.  The  most  frequent  was 
swimming  off  the  bottom  for  a  distance  of  about  8  cm,  grabbing  a  small  crustacean  and  then 
settling.  Another  pattern  involved  picking  a  benthic  organism  from  the  substrate.  The  third 
involved  picking  up  a  mouthful  of  the  loose  bottom,  spitting  it  out,  and  then  selecting  the 
desired  food  as  it  fell  through  the  water.  By  the  latter  two  methods  such  items  as  decapods, 
annelids,  and  echinoderms  were  taken.  These  items  were  probably  detected  visually,  although 
this  species  does  have  moderately  developed  papillae  (Macdonald,  1972)  which  may  function 
in  sensing  burrowing  organisms. 

I  noted  one  exception  to  the  general  benthic  habit  on  the  southwest  coast  of  Punta  Banda, 
Mexico,  where  there  are  many  narrow  canyons  through  which  strong  upwelling  currents  flow. 
Gobies  there  hovered  0.5  to  1.0m  off  the  bottom,  using  their  pectoral  fins  as  the  means  to 
counter  the  current,  as  they  fed  on  the  plankton  that  drifted  up  the  canyons. 

In  the  aquarium,  C.  nicholsii  was  fed  on  frozen  brine  shrimp.  Usually  the  gobies  grabbed 
the  shrimp  in  midwater,  and  swallowed  them  whole.  If  the  shrimp  was  picked  off  the  bottom 
the  goby  would  spit  it  out  with  the  associated  rubble  and  then  once  again  grab  the  shrimp  as  it 
drifted  downward.  When  fed  on  chunks  of  frozen  smelt,  which  were  too  large  to  be  swallowed 
whole,  the  gobies,  which  have  moderately  developed  teeth,  bit  off  large  pieces. 

BREEDING  BEHAVIOR 

Gobies  and  species  of  various  other  groups  endowed  with  adherent  eggs  generally  seek 
some  rocky  crevice,  shell,  or  other  hard  object  suitable  for  egg  attachment.  As  is  typical  of 
gobioid  fishes,  and  most  territorial  fishes,  the  male  C.  nicholsii  selects  and  prepares  the  nest 
(Tavolga,  1954;  Dotu,  1958a;  Guitel,  1893).  Nest  building  is  intermittent,  alternating  with 
courtship  and  other  phases  of  social  behavior.  The  male  enters  and  leaves  his  shelter  frequently. 
The  duration  of  the  stay  within  the  shelter  is  highly  variable. 

Nest  preparation  consists  of  several  cleaning  movements,  similar  to  those  described  by 
Tavolga  (1954)  for  Bathygobius  soporator:  fanning,  rubbing,  scooping,  and  nibbling.  Fanning 
the  most  frequent  act,  consists  of  the  vigorous  waving  of  the  body  and  pectoral  fins,  sending  up 
a  cloud  of  sand  for  several  seconds.  This  appears  to  be  the  most  efficient  type  of  digging  activ- 
ity for  this  fish.  Rubbing  consists  of  brushing  the  body  against  the  algae-covered  surface  of  the 
nest,  apparently  to  dislodge  this  material.  Scooping  is  accomplished  by  taking  mouthfuls  of 
sand,  small  shells,  or  other  debris  and  carrying  it  away  from  the  nest.  Nibbling  may  occur  if  the 
shelter  has  algae  or  other  material  clinging  to  its  surface.  Nest  cleaning  is  not  thorough  and  the 
adhesive  algae  and  other  organisms  are  not  cleaned  off  completely. 

Essentially  the  same  nest-preparation  movements  observed  in  C.  nicholsii  have  been 
described  for  other  species  of  gobies,  as  well  as  for  most  teleosts  that  construct  any  sort  of 
hollow  in  sand  substrates.  The  fanning  method  of  nest  formation  is  probably  the  most  wide- 
spread of  the  nesting  behaviors  in  fishes  and  has  been  described  for  the  Centrarchidae  (Breder 
and  Rosen,  1966),  certain  cichlids  (Baerends  and  Baerends-Van  Roon,  1950),  and  Clinocottus 
and  other  cottids  (Breder  and  Rosen,  1966). 

When  courting,  each  male  rose  a  few  centimeters  off  the  bottom,  spread  his  fins  fully,  and 
settled  back  to  the  substrate.  After  one  to  several  of  these  displays,  he  swam  back  to  his  nest, 
where  he  continued  nest  construction  for  a  short  time,  and  then  resumed  his  courting. 

Intermittently  the  male  swam  swiftly  toward  the  female  and  then  quickly  returned  to  his 
starting  place,  apparently  in  an  attempt  to  stimulate  the  female  and  to  attract  her  to  his  nest 
site.  If  the  female  reacted  negatively,  the  male  followed  and  continued  courting,  often  nipping 
and  chasing  her.  Early  in  courtship  the  female  simply  darted  away  into  other  shelters.  Prior  to 
spawning  the  female  often  took  shelter  in  the  male's  nest,  from  which  she  was  chased.  However, 
the  male,  upon  seeing  the  female  within  the  nest,  approached  with  courting  movements  before 
entry  and  chasing.  Apparently  the  male  faced  a  conflict  situation  between  defense  of  his  nest 
and  enticing  the  female  to  remain  and  spawn.  Occasionally  a  female  approached  a  male  that 
was  courting  her.  The  female  then  slowly  undulated  the  body,  gaped,  and  spread  the  fins. 

Some  form  of  courtship  behavior  is  exhibited  by  males  of  most  nesting  species  of  fishes. 
There  is  considerable  interspecific  variation  in  the  details  of  this  behavior  within  gobiids.  The 


202 

courting  male  of  Gobius  munitus  exhibits  body  tremors  and  rapid  breathing  movements  while 
approaching  the  female  with  short  hops,  his  fins  bristling,  head  raised,  throat  puffed,  and 
mouth  agape  (Guitel,  1892).  Males  of  Gobiosoma  approach  the  female  with  short  darts,  with 
fins  widely  spread  (Breder,  1942).  Courting  males  of  Brachygobius  xanthozdnus  swim  back 
and  forth  in  front  of  the  female  (Field,  1945).  Tlie  courtship  display  of  a  male  Elacatinus 
oceanops  consists  of  violent  swimming  while  he  clings  to  the  substrate  with  the  pelvic  cup;  he 
then  butts  the  female  in  the  head  and  genital  regions  with  his  nose,  and  slaps  her  on  the  head 
with  his  caudal  fin  (Feddern,  1967).  The  male  of  Batliygobius  soporator  slowly  approaches  the 
female  and  positions  himself  beside  or  in  front  of  her;  he  then  waves  his  body,  tail,  and  pec- 
torals in  a  manner  similar  to  that  involved  in  nest  cleaning;  if  the  female  moves  away,  the  male 
follows  and  continues  courting,  often  chasing  and  nipping  her  (Tavolga,  1954). 

A  change  in  the  color  of  the  pelvic  fin  of  Coryphoptenis  nicholsii  males  during  the  breed- 
ing season  presumably  stimulates  courtship  response  by  the  female.  Color  changes  have  been 
noted  in  various  fins  of  other  gobiid  fishes  during  the  breeding  season  (Dotu,  1956;Kinzer, 
1960).  Tavolga  (1954)  showed  that  Bathygobius  soporator  females  can  at  times  "recognize" 
males  by  coloration,  in  the  absence  of  any  courtship  activity.  Tliat  the  male's  black  pelvic  fin 
may  provide  adequate  stimulus  for  sexual  recognition  was  suggested  in  an  experiment  wherein 
an  adult  female  with  artificially  blackened  pelvic  fins  was  approached  by  a  ripe  female  showing 
definite  courtship  behavior. 

The  female  C.  nicholsii  attaches  her  eggs  to  the  underside  of  the  nest.  Tlie  deposited  egg 
masses  average  10  cm  in  diameter,  are  roughly  circular  and  are  made  up  of  a  single  layer  of  eggs. 
The  male  is  intermittently  present  in  the  nest  at  the  time  of  oviposition,  where  he  frequently 
circles  the  female,  butting  and  bitting  her.  He  often  passes  his  turgid  genital  papilla  over  the 
surface  of  the  eggs.  Fertilization  apparently  takes  place  during  and  immediately  after 
oviposition. 

Among  the  gobies  whose  nesting  habits  have  been  described,  oviposition  on  the  underside 
of  shelters  is  quite  common.  Fishes  that  hide  and  nest  under  shelters  are  likely  to  be  confronted 
with  a  nest  floor  which  consists  of  sand,  mud,  rubble,  or  other  irregular  surfaces,  whereas  the 
ceiling  will  probably  present  a  hard  surface  more  suitable  for  the  adherence  of  eggs.  Also  such 
shelters  offer  a  degree  of  safety  against  egg  predators. 

Tavolga  (1950)  found  that  unless  artificially  fertilized  eggs  of  Bathygobius  soporator  ?lxq 
placed  in  a  hanging  position  many  of  the  embryos  do  not  rotate  properly  within  their  elongate 
egg  cases.  Such  individuals  develop  with  their  heads  pointed  toward  the  attached  end  of  the 
shell  and  are  thus  unable  to  hatch. 

Spawning  completed,  the  male  of  C  nicholsii  defends,  cleans,  and  fans  the  eggs  until  they 
hatch.  He  fans  the  eggs  by  intermittently  waving  his  body  and  pectoral  fins.  This  movement 
resembles  that  of  nest  preparation,  and  produces  a  strong  current  of  water  over  the  eggs.  Tlie 
residing  male  rushes  out  to  chase  away  any  goby  or  other  organism  that  approaches  the  nest,  as 
well  as  a  slurp  gun  placed  in  front  of  the  nest. 

The  male  guards  the  eggs  in  the  majority  of  nesting  fishes  (Breder  and  Rosen,  1966). 
Among  the  gobies,  only  Typhlogobius  is  reported  as  an  exception,  in  that  both  sexes  guard  and 
fan  the  spawn  (MacGinitie,  1939).  Brood  care  by  the  male  of  C  nicholsii  consists  almost  exclu- 
sively of  fanning  with  some  or  all  of  the  fins.  Brood  care  is  practiced  by  most  nesting  fishes. 
Tlie  function  of  the  brooding  is  three-fold:  circulation  of  water  for  respiration,  prevention  of 
bacterial  and  fungal  growth,  and  defense  of  the  eggs.  Tavolga  (1954)  proposed  another  func- 
tion in  that  the  fanning  activity  of  the  male  in  some  way  prevents  the  abnormal  positioning  of 
the  embryos  within  the  eggs. 

SOCIAL  BEHAVIOR 

Coryphopterus  nicholsii  is  a  bottom  dweller.  Its  swimming  activity  is  confined  to  short, 
quick  spurts  for  feeding,  for  territory  defense,  and  for  escape  to  shelter.  Although  it  is  able  to 
change  color  and  pattern  according  to  the  habitat,  its  light  color  is  generally  retained  even  in 
dark,  rocky  areas.  This  light  color  blends  well  with  the  sand  bottom,  which  appears  to  be  the 
preferred  substrate  type  of  this  goby.  Tlie  black  eyes  and  tip  of  the  first  dorsal,  although  con- 
spicuous to  the  human  observer,  may  serve  as  disruptive  markings,  breaking  up  the  shape  of  the 
goby  before  a  predator. 


203 


Like  many  other  gobies  (Stebbins  and  Kalk,  1961;  Tavolga,  1954),  C.  nicholsii  exhibits 
territorial  behavior.  It  is  a  solitary  species  that  sets  up  a  territory  that  includes  a  shelter  and  a 
feeding-display  area  in  front  of  it.  Tlie  juxtaposition  of  the  territories,  which  may  be  spaced 
less  than  25  cm  between  shelter  centers,  induces  numerous  encounters  between  neighboring 
individuals,  hi  the  aquarium  any  available  shelter,  including  a  tank  corner,  was  utilized  and 
fought  over.  The  species  was  quite  aggressive  in  a  tank  containing  several  fish;  there  was  almost 
continuous  nipping  and  chasing. 


Figure  13.  Changes  in  coloration  of  Coryphopterus  nicholsii  during  social  interaction.  A.  Normal  light  color- 
ation of  undisturbed  or  dominant  individual.  B.  Intermediate  darkening  of  submissive  goby.  C.  Final  darkened 
coloration  of  submissive  goby.  Note  light  spot  under  eye.  This  spot  is  blue  in  life  and  gives  rise  to  the  ver- 
nacular name,  biuespot  goby. 


204 

A  variable  nip  order  was  established  which  was  somewhat,  but  not  absolutely,  correlated 
with  size  (see  also  Bopp,  1957;  Tavolga,  1954).  Social  orders  were  established  very  quickly. 
During  the  breeding  season,  spawning  males,  which  were  the  largest  individuals,  appeared  to  be 
dominant.  Subdominant  members  retreated  to  higher  levels  of  the  aquarium,  hanging  on  the 
tank  walls  and  corners  by  means  of  continuous  swimming  movements  of  the  fins  and  by  the 
suction  of  their  fused  pelvic  fins.  Some  attempted  to  jump  out  of  the  tank. 

The  fundamental  color  of  C.  nicholsii  is  uniform  pale  yellow,  but  the  coloration  is  vari- 
able, changing  rapidly  in  response  to  different  social  situations.  During  social  interactions  the 
subordinate  animal  usually  becomes  much  darker,  and  is  mottled  (Fig.  13).  One  subordinate 
individual  which  had  been  displaced  from  the  bottom  of  the  tank  took  up  a  position  three- 
fourths  of  the  way  up  the  tank  wall  next  to  the  filter  siphon  which  had  evenly  spaced  holes 
with  growths  of  dark  algae.  The  color  pattern  of  the  fish  quickly  matched  that  of  the  siphon 
holes.  This  individual  held  this  position  and  retained  this  color  pattern  for  several  days. 


i^.^ 


Figure  14.  Combat-threat  posturing  behavior  of  adult  males  of  Coryphopterus  nicholsii  in  aquarium. 

The  highest  degree  of  aggressive  behavior  was  observed  in  encounters  between  mature  fish 
of  about  equal  size.  The  two  gobies  approached  each  other  with  slow  undulations  of  the  body, 
and  with  all  fins  stiffly  erected.  They  positioned  themselves  next  to  one  another,  directly  head 
on  (Fig.  14),  or  head  to  tail.  The  mouth  was  then  widely  gaped,  with  the  throat  expanded  and 
the  head  elevated.  The  two  gobies  displayed  either  alternately  or  simultaneously.  The  "loser" 
assumed  the  mottled  color  pattern  and  dashed  to  the  safety  of  shelter. 

Interactions  between  fish  of  different  sizes,  or  after  a  hierarchy  has  been  established, 
usually  involve  a  quick  dash  by  one  of  the  fish,  the  more  dominant  one,  with  the  other  fish 
retreating.  The  more  dominant  fish  nips  the  fins  and  scales  of  the  retreating  individual.  In  the 
laboratory,  the  fish  of  the  higher  rank  continually  attacked  the  subordinate  intruders  if  the 
tank  was  not  large  enough  to  provide  adequate  territories  for  the  gobies  present. 

Pugnacity,  a  feature  common  to  gobies  and  most  territorial  fishes,  is  expressed  in  C. 
nicholsii  as  a  simple  type  of  biting  and  pursuit  behavior  between  combatants.  Guitel  (1892) 
described  similar  darkening,  throat  puffing,  gaping,  and  fin  stiffening  in  Gobius  minutus.  Breder 
(1942)  reported  that  males  of  Gobiosoma  robustum  exhibit  darkening  and  fin  spreading  as 
intimidation  mechanisms.  Tavolga  (1954)  found  Bathygobius  sporator  also  exhibits  color 
changes  correlated  with  fighting  and  with  reproductive  behavior,  especially  in  males,  and  that 
extreme  darkening  is  characteristic  of  fighting  males;  this  occurs  together  with  throat  puffing, 
gaping,  quivering,  butting,  and  biting  movements.  Weisel  (1947)  found  that  intimidation  be- 
havior o[  Gillichthys  mirabilis  consists  almost  entirely  of  the  display  of  its  huge  gape. 

ACKNOWLEDGMENTS 

I  am  grateful  to  Dr.  David  W.  Greenfield,  my  major  sponsor,  for  his  assistance  and  stimulation  in  every 
phase  of  this  study.  I'or  help  in  the  field,  I  should  like  to  thank  Jack  C.  Turner,  David  M.  Wildrick,  and 
Dannie  A.  Hensley.  Dr.  Philip  Adams  deserves  special  thanks  for  his  assistance  in  the  laboratory  and  field. 
Robert  J.  Lavenberg  provided  the  specimens  of  pelagic  prejuveniles.  I  wish  to  thank  Dr.  Bayard  Brattstrom 
and  Dr.  Kenneth  L.  McWilliams  for  their  suggestions  and  critical  reading  of  an  early  draft  of  the  manuscript. 
Dr.  Carl  L.  Hubbs  ottered  many  helpful  suggestions  for  the  manuscript.  Terry  Greenfield  typed  the  manu- 
script and  helped  in  the  laboratory,  for  which  I  am  very  grateful.    My  wife,  Beth,  provided  the  illustrations. 


205 

LITERATURE  CITED 

Baerends,  G.  P.,  and  J.  M.  Baerends-Van  Roon. 

1950.  An  introduction  to  the  study  of  the  ethology  of  cichlid  fishes.  Behaviour,  Suppl.  No.  1:  1-242. 

Baird,  R.  C. 

1965.  Ecological  implications  of  the  behavior  of  the  sexually  dimorphic  gohy  Microgobius  gulosus 
(Girard).  Inst.  Mar.  Sci.,  Texas  10:  1-8. 

Barnhart,  P.  S. 

1936.  Marine  fishes  of  southern  Cahfornia.   Univ.  California  Press,  Berkeley.   209p. 

Bean.T.  H. 

1881.  Notes  on  a  collection  of  fishes  made  by  Henry  E.  Nichols,  U.S.N.,  in  British  Columbia  and  south 
Alaska,  with  description  of  a  new  species  and  a  new  genus.    Proc.  U.S.  Natl.  Mus.  4:  463-474. 

Bean,  B.  A.,  and  A.  C.  Weed 

1919.  Notes  on  a  collection  of  fishes  from  Vancouver  Island,  B.C.  Trans.  Royal  Soc.  Canada,  ser.  3, 
13  (4):  69-83. 

Berry,  F.  H.,  and  H.  C.  Perkins 

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1964.  The  epifauna  of  a  California  siltstone  reef.   Ecology  45(2):  272-283. 
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1936.  Keys  to  the  fishes  of  Washington,  Oregon  and  closely  adjoining  areas.  Univ.  Washington  Publ. 
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1936.  Fishes  of  the  American  northwest:  a  catalog  of  the  fishes  of  Washington  and  Oregon,  with  dis- 
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1964.  Tacnioides  limicola,  a  new  goby  from  Guam,  Marianas  Islands.   Micronesica  1 :  145-150. 

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California  State  University,  Fullerton,  California  92631.   Present  address:  16341  Skymeadow 
Drive,  Placentia,  California  92670. 


.^^ 


MUS.  COM  P.  ZOOU 
LIBRARY 

SeP30S74 

HARVARD 
UNIVERSITY. 


ANE\N  PLATYDOR/S  (GASTROPODA:    NUDIBRANCHIA) 
FROM  THE  GALAPAGOS  ISLANDS 


DAVID  K.  MULLINER  AND  GALE  G.  SPHON 


TRANSACTIONS 

OF  THE  SAN  DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  15  12  APRIL  1974 


A  NEW  PLATYDORIS  (GASTROPODA;  NUDIBRANCHIA) 
FROM  THE  GALAPAGOS  ISLANDS 

DAVID  K.  MULLINER  AND  GALE  G.  SPHON 


ABSTRACT. — Platvdoris  carolynaen.  sp.  is  described  from  the  Galapagos  Islands  and  compared  with 
the  two  eastern  Pacific  species  of  PlatyJoris  and  with  P.  scabra.  the  only  member  of  this  genus  with 
wide  distributional  limits.  Platydorids  are  rasping  sponge  feeders  that  live  in  tropical  and  temperate 
oceans.  The  distribution  and  nomenclature  of  the  36  known  species  is  reviewed  briefly. 

The  nudibranch  fauna  of  the  Galapagos  Islands  has  been  neglected  by  previous 
workers.  Apparently,  only  two  species,  Doris  peruviana  Orbigny  1837  and  Onchidium 
k'slici  Stearns  1893,  have  been  reported  (Pilsbry  and  Vanatta,  1902:  556;  Stearns,  1893: 
383).  Yet,  in  March  1971  members  of  the  Ameripagos  Expedition  to  the  Galapagos 
Islands  collected  at  least  15  species  of  nudibranchs,  some  of  them  fairly  common,  at 
various  localities  in  the  islands  (Sphon  and  MuUiner,  1972).  Included  among  these  was  a 
previously  undescribed  species  of  Plat}'doris  that  was  found  at  several  localities,  and  which 
may  be  endemic  to  these  islands.  In  this  paper,  we  describe  this  new  species,  and  briefly 
review  the  distribution  and  nomenclature  of  Pkitydoris. 

BIOGEOGRAPHY 

Members  of  the  genus  Pkitydoris  are  sluggish,  retiring  invertebrates  that  cling  tightly 
tc  crevices  on  the  underside  of  rocks  and  coral  heads.  They  are  found  in  tropical  and 
temperate  waters  from  40°  N  latitude  to  32°  S  latitude.  All  but  one  of  the  thirty-six  known 
species  have  limited  ranges,  usually  consisting  of  one  shoreline,  one  island  chain,  or  one 
location  (Fig.  1).  Platydoris  scabra  (Cuvier,  1804)  is  the  exception,  ranging  in  tropical 
waters  from  38°  E  longitude  to  155°  W  longitude. 

The  majority  of  the  platydorids  are  found  in  the  Indo-Pacific.  They  are  rasping 
sponge-feeders,  and  the  great  abundance  and  diversity  of  sponges  may  account  for  the 
large  number  of  platydorids  found  in  these  seas  as  compared  to  the  Atlantic  or  Eastern 
Pacific. 

SYSTEMATICS 

Order  Nudibranchia 

Family  Dorididae 

Genus  Pkity^doris  Bergh,  1877 

Definition. — The  body  is  leathery,  flattened,  and  oval  with  a  coarse  to  smoothly 
granular  mantle.  The  foot  is  notched  anteriorly.  The  branchial  aperture  is  oval  and  six- 
lobed.  There  is  no  labial  armature,  and  the  radula  consists  of  numerous  hamate  teeth. 
The  stomach  is  large;  the  penis  is  armed  with  small  spines,  and  the  vagina  has  a  thick 
cuticular  lining  (translated  and  modified  after  Bergh,  1877). 

Type  species. — Platydoris  argo  (Linnaeus,  1758),  by  original  designation. 

Platydoris  carolynae  n.  sp. 

Type  locality. — Docking  area,  Charles  Darwin  Research  Station,  Santa  Cruz  Island, 
Galapagos  Islands,  Ecuador,  (0°  45'  05"  S,  90°  15'  38"  W). 

Description.  —The  ground  color  of  the  animal  is  cream,  the  entire  dorsum  mottled 
with  black  or  brown  blotches.  The  ventral  side  of  the  mantle  is  also  cream  with  black  or 
brown  spotting,  each  spot  made  up  of  multiple  fine  cross-hatched  lines.  The  rhinophores 
are  tan-colored  with  dark  brown  spots.  The  branchiae  are  translucent  with  dark  brown  or 


SAN  DIEGO  SOC.  NAT.  HIST.,  TRANS  17(15):  209-216.  12  APRIL  1974 


210 


140"  120°  IOC  80° 60° 40* ?0°  W  0°  E  20'  40° W 


100°  120°  140°  160°        E    180°  W        160°  140° 


Figure  1.  Distributional  records  for  Platydoris  scabra  are  indicated  by  the  solid  dots.  Numerals  represent  the 
number  oi  Platydoris  species  found  at  each  locality. 

black  specks  (Fig.  2). 

The  rhinophores  are  perfoliate  with  twenty-four  leaves.  They  are  completely  retract- 
able and  set  in  a  rhinophoral  pit  with  a  seven-lobed  margin.  The  branchiae  are  completely 
retractable,  tripinnate,  six  in  number  and  divided  into  two  groups  of  three.  The  anterior 
end  of  the  branchial  opening  forms  a  crenulate  lobule.  The  pharynx  extends  for  approxi- 
mately half  the  distance  from  the  anterior  end  of  the  foot  to  the  edge  of  the  dorsum. 

Two  small  head  tentacles  are  attached  to  the  head  between  the  mantle  and  the  body. 
No  eyespots  are  visible.  The  foot  is  bilabiate  anteriorly  to  just  behind  the  foot  corner, 
notched  medially.  The  radula  is  approximately  heart-shaped  with  76  longitudinal  rows  of 
hamate  teeth.  No  rachidian  teeth  are  present.  The  radula  formula  is  76  x  70.0.70  (Figs.  3, 
4).  In  the  reproductive  system  the  spermatocyst  is  elongated,  connecting  directly  into  the 
mucus  and  albumen  gland.  A  short  convoluted  tube  connects  to  the  oval  spermatheca. 
The  prostate  is  large  and  globular.  The  penis  is  armed  with  erect,  slightly  curved  spines. 
The  vagina  is  lined  with  thick  cuticle-bearing  folds  (Fig.  5). 

Ety^mology. — This  species  is  named  for  Carolyn  Stover,  a  member  of  the  Ameripagos 
Expedition. 

Type  material. — Holotype,  California  Academy  of  Sciences,  Invertebrate  Zoology 
Type  Series  No.  303.  Photographs  of  the  living  animal  are  deposited  in  the  CASIZ  slide 
collection  as  Nos.  153-155.  The  specimen,  which  is  46.4  mm  long  and  32.5  mm  wide  was 
collected  by  Andre  DeRoy  on  13  February  1964,  intertidally  at  the  Charles  Darwin 
Research  Station  dock. 

Paratypes  (7). — One  specimen  deposited  at  the  Charles  Darwin  Research  Station, 
collected  intertidally  in  shallow  pools  on  Santa  Cruz  Island.  One  specimen  deposited  at 
the  Los  Angeles  County  Museum  of  Natural  History,  Invertebrate  Zoology.  Type  Collec- 
tion No.  1619,  collected  intertidally  at  Duncan  Island,  by  the  Ameripagos  Expedition  on 
26  March  1971.  Two  specimens  deposited  at  the  San  Diego  Natural  History  Museum, 
Department  of  Marine  Invertebrates:  SDSNH  No.  62826,  collected  from  10m  at  Jervis 
Island  by  the  Ameripagos  Expedition  on  24  March  1971;  Radula  slide  and  dissected 
animal  SDSNH  No.  62827,  collected  from  6m  off  Punta  Alfaro,  Isabella  Island,  by  the 
Ameripagos  Expedition  on  25  March  1971.  One  specimen  deposited  at  the  American 
Museum  of  Natural  History,  Department  of  Living  Invertebrates,  AMNH  No.  173729, 
collected  from  10m,  off  Jervis  Island,  by  the  Ameripagos  Expedition  on  24  March  1971. 
One  specimen  deposited  at  the  United  States  National  Museum  of  Natural  History,  type 
Collection  No.  735349,  collected  from  6m  off  Punta  Alfaro,  Isabella  Island,  by  the 
Ameripagos  Expedition  on  26  March  1971.  One  specimen  deposited  at  the  Delaware 
Museum  of  Natural  History,  No.  64524,  collected  at  Long  Beach  on  the  northern  coast  of 
Santa  Cruz  Island  by  Sue  Andrews  on  21  December  1972.  The  paratypes  range  in  size 
from  19.5  mm  long  and  14.2  mm  wide  to  42.3  mm  long  and  28.0  mm  wide. 

Discussion. — The  only  species  of  Platydoris  known  from  the  eastern  Pacific  are  P. 
tnacfarhmdi  Hanna,  1951,  and  P.  punctatella  Bergh,  1898.  The  three  species  are  separ- 


211 


Figure  2.  Platydoris  carolynae,  dorsal  (top)  and  ventral  (bottom)  views. 

able  by  external  appearance  and  geographical  range.  Platydoris  macfarlatidi  is  known 
only  from  the  type  lot  of  four  specimens  dredged  from  172m  off  Pismo  Beach,  San  Luis 
Obispo  County,  California.  It  is  dark  red,  velvety  smooth,  with  no  spots  or  markings  on 
the  surface.  The  foot  tapers  to  a  point  posteriorly.  Platydoris  punctatella  is  from  "Isia  de 
Pajargo",  Chile  (?  =  Isla  de  Pajaros,  Chile,  ca.  26°  S.  lat.).  It  is  pale  yellow.  The  rhino- 
phores  and  the  anterior  margin  of  the  foot  are  bright  yellow.  The  back  has  a  few  scattered 


212 


Figure  3.  Radula  with  offset  drawings  of  individual  teeth. 

light  brown  spots  and  flecks.  The  underside  of  the  mantle  is  smooth  and  taint  yellow  with 
no  markings. 

Platydoris  carolytuie  is  known  only  from  the  Galapagos  Islands.  The  foot  is  round 
posteriorly  in  contrast  to  P.  nnicfurlandi.  The  color  is  cream  or  white  with  brown  or  black 
mottling  on  the  dorsum.  Each  of  the  spots  on  the  ventral  side  of  the  mantle  is  made  up  of 
fine  cross-hatched  lines.  Platydoris  punctatella  has  no  ventral  markings. 

Internal  differences  were  noted  in  P.  carolynae.  The  radula  formula  is  76  x  70.0.70 
for  a  46  mm  animal,  whereas  the  radula  of  a  50  mm  P.  scuhra  is  less  elongated,  49  x 
103.0.103.  The  vas  deferens  in  P.  scuhra  is  long  and  coiled,  whereas  it  is  short  and 
straight  in  P.  carolynae. 


THE  SPECIES  OF  PLATYDORIS 

We  have  been  able  to  find  46  specific  names  described  as.  or  later  assigned  to  Platy- 


213 


doris  in  the  literature.  Of  these,  one  (P.  variolata)  has  been  shifted  to  Anisodoris;  eight 
are  currently  considered  synonyms;  and  two  are  nomina  nuda.  The  remaining  names  are 
listed  alphabetically  in  Table  1.  Synonyms  are  cited  chronologically  under  the  currently 
accepted  name.  We  have  also  indicated  the  distribution  of  each  species  and  have  added 
the  two  nomina  nuda  at  the  end  of  the  list,  as  they  both  appear  to  be  undescribed  species. 

TABLE  1.  Currently  recognized  species  of  Platydoris  and  their  distributions. 


SPECIES 


DISTRIBUTION 


P.  angustipes  (Morch,  1863) 

Synonyms: 

P.  angustipes  alaleta  (Bergh.  1877a) 

P.  rubra  White.  1952 
P.  argo  (Linnaeus,  1767)  (Type  species  of  the  genus) 

P.  canariensis  (Orbigny,  1839) 

P.  capricomensis  Allan,  1932 

P.  carinata  Risbec,  1928 

P.  cruenta  (Quoy  and  Gaimard,  1832) 

Synonym: 

P.  arrogans    Bergh,  1877a 
P.  dura  Pruvot-Fol,  1951 
P.  ellioti  (Alder  and  Hancock,  1864) 
P.  flammulata  Bergh,  1905 
P.  formosa  (Alder  and  Hancock,  1864) 
P.'galhanus  Burn,  1958 
P.  hi'patica  (Abraham,  1877) 
P.  herdmani  Farran,  1905 
P.  immonda  Risbec,  1928 
P.  incena  Eliot.  1904 
P.  inframaculata  (Abraham,  1877) 
P.  infrapicta  (Smith.  1884) 
P.  laminea  Risbec,  1928 
P.  macfarlandi  Hanna,  1951 
P.  murrea  (Abraham,  1877) 
P.  noumeae  Risbec,  1928 
P.  papiUata  EHot.  1904 
P.  philippi  Bergh.  1877a 
P.  pukhra  Eliot.  1904 
P.  punctata  (Orbigny,  1839) 
P.  punctatella  Bergh,  1898 
P.  sanguinea  Bergh,  1905 
P.  scahra  (Cuvier,  1804) 
Synonyms: 

P.  coelestis  (Kelaart,  1858) 

P.  I'urychlamys  Bergh.  1877a 

P.  coriacea  (Abraham,  1877) 

P.  vicina  Bergh,  1880 

P.  in-dalci  Allan.  1932 
P.  sordida  (Quoy  and  Gaimard.  1832) 
P.  speciosa  (Abraham.  1877) 
P.  spinulosa  Farran.  1905 
P.  spongllla  Risbec.  1928 
P.  striata  (Kelaart,  1858) 
P.  townscndi  EWot.  1905 
P.  variolata  (Orhigney.  1837) 

See:  Anisodoris  variolata  (Bergh.  1898) 
P.  varicgata  Bergh,  1880 

NOMINA  NUDA 
P.  hnumca  Bergh.  1877a 
P.  niarniorata  Bergh.  1877b 


Southern   Florida, 
Bahi'a,  Brazil. 


through   Caribbean,    and   south    to 


Mediterranean  Sea.  Also,  a  questionable  report  from 
the  East  Indies. 
Canary  Islands. 

Capricorn  Group,  Queensland,  Australia. 
New  Caledonia. 

Western  Pacific,  Japan,   Philippine  Islands,   and   East 
Indies. 


Mediterranean  Sea. 

Indian  Ocean  and  southeast  coast  of  India. 

East  Indies. 

Eastern  Indian  Ocean;  also,  reported  from  Hawaii. 

Southern  Australia. 

Riciniola      (Pacific  Ocean). 

Ceylon. 

New  Caledonia. 

Zanzibar. 

Ceylon  and  East  Indies. 

Queensland,  Australia. 

New  Caledonia. 

Central  California. 

Mauritius,  Indian  Ocean. 

New  Caledonia. 

Eastern  Africa. 

Mediterranean  Sea. 

Eastern  Africa. 

Canary  Islands. 

Isla  de  Pajaros,  Chile. 

East  Indies. 

Indian  and  western  Pacific  Oceans. 


Mauritius,  Indian  Ocean. 

Western  Pacific  Ocean. 

Ceylon. 

New  Caledonia. 

India  and  Japan. 

India. 

Central  Chile. 

Tahiti. 


Although  this  is  the  type  locality  o\  P.  hcpatica.  as  given  by  Abraham  (1877).  we  have  been  unable  to  locate 
such  a  locality  from  available  gazetteers. 


214 


Figure  4.  Scanning  electron  micrographs  from  a  section  of  the  radula  at  3a.  Left,  X400;  right,  XQOO. 


_--  — P*" 


mg  — 


a-- 


1.0mm 


Figure  5.  Camera-lucida  drawing  of  reproductive  organs  with  offset  of  a  cirral  hook,  hd-hermaphrodite  duct, 
sc-spermatocyst.  st-spermatheca.  pr-prostate,  vd-vas  deferens,  v-vagina,  p-penis,  od-oviduct,  mg-mucus  gland, 
a-albumen  gland,  am-anipulla,  eg-external  genital  opening. 


215 


ACKNOWLEDGEMENTS 

We  wish  to  thank  the  Charles  Darwin  Research  Station  and  its  Director,  Peter  Kramer,  for  making  it 
possible  for  the  Ameripagos  Expedition  to  collect  in  the  Galapagos  Islands. 

We  also  wish  to  thank  Allyn  G.  Smith,  Department  of  Invertebrate  Zoology,  California  Academy  of  Sciences 
for  lending  specimens  and  photographs. 

Joe  Nakanishi  of  the  Los  Angeles  County  Museum  of  Natural  History  prepared  the  illustrations  of  the 
dorsal  and  the  ventral  views  of  the  holotype;  Anthony  D'Attilio  of  the  San  Diego  Natural  History  Museum  drew 
the  reproductive  organs  and  the  radula;  Michael  Featherby  made  the  SEM  photographs.  James  Lance  helped 
with  the  literature  search  and  offered  technical  advice.  George  E.  Radwin  read  the  manuscript,  offered  technical 
advice,  and  extracted  and  mounted  the  radula.  Thanks  are  also  due  to  the  Ameripagos  Expedition  members  for 
their  collecting  help  and  companionship. 

LITERATURE  CITED 

Abraham,  P.  S. 

1877.  Revision  of  the  anthobranchiate  nudibranchiate  Mollusca,  with  descriptions  or  notices  of  forty-one 
hitherto  undescribed  species.  Proc.  Zool.  Soc.  London,  1877,  p.  l%-269,  pis.  27-30. 
Alder,  J.  and  Hancock,  A. 

1864.  Notice  of  a  collection  of  nudibranchiate  mollusca  made  in   India.   Trans.   Zool.   Soc.   London  5: 
113-147,  pis.  28-33. 
Allan,  J. 

1932.  Australian  nudibranchs.  Australian  Zool.  7(2):  87-105,  pis.  4,  5. 
Bergh,  R. 

1877a  Malacologische  Untersuchungen,  Nudibranchiaten.  ///  C.  G.  Semper,  Reisen  im  Archipel  der  Phil- 
ippinen.  Sect.  2,  2(12):  495-546.  pis.  58-61. 

1877b  Kritische  Untersuchung  der  Ehrenberg'  schen  Doriden.  Jahrb.  Deutsch.  Malakoz.  Ges.  4:  45-76. 

1880.  Malacologische  Untersuchungen,  ///  C.  G.  Semper,  Reisen  im  Archipel  der  Phiiippinen.  Sect.  2,  4(1): 
Suppl.  (1)  1-78,  pis.  A-F. 

1898.  Die  Opisthobranchiata  der  Sammlung  Plate.  Zool.  Jahrb.  Suppl.  4(3):  481-582,  pis.  28-33. 

1905.  Die  Opisthobranchiata  der  Siboga-Expedition.  Monographic  50:  1-248,  pis.  1-20. 
Burn,  R. 

1958.  Further  Victorian  Opisthobranchia.  J.  Malacol.  Soc.  Australia  2:  20-36,  pis.  6,  7. 
Cuvier 

1804.  Suite  de  I'extrait  des  memoires  sur  les  mollusques.  Bull.  Sci.  Soc.  PhiJom.  Paris.  3(93):  254-256,  pi.  22. 
Eliot,  C. 

1904.  On  some  nudibranchs  from  East  Africa  and  Zanzibar.  Part  III.   Proc.  Zoo).  Soc.   London,   1903, 
p.  354-385,  pis.  22-24. 

1905.  Nudibranchs  from  the  Indo-Pacific.  1.  Notes  on  a  collection  dredged  near  Karachi  and  Maskat. 
J.  Conch.  11(8):  237-256.  pi.  5. 

Farran,  G. 

1905.  Report  on  the  opisthobranchiate  mollusca  collected  by  Professor  Herdman,   at  Ceylon,   in    1902. 
Report  to  the  Government  of  Ceylon  on  the  pearl  oyster  fisheries  of  the  Gulf  of  Manaar.   Suppl. 
Rep.  21:  329-364,  pis.  1-6. 
Hanna,  G.  D. 

1951.  A  new  west  American  nudibranch  mollusk.  Nautilus  65(1):  1-3. 
Kelaart,  E.  F. 

1858.  New  and  little  known  species  of  Ceylon  nudibranchiate  molluscs  and  zoophytes.  J.  Roy.  Asiatic  Soc, 
Ceylon  Branch,  Colombo  3(9):  76-111. 
Linnaeus,  C. 

1767.  Systema  Naturae,  12th  ed.,  pp.  1-1367. 
Marcus,  E.  and  E.  Marcus 

1960.  Opisthobranchia  aus  dem   Roten   Meer  und   von  den   Malediven.   Akad.   Wiss.   Lit.   Abh.   Math. 

Naturwiss..  12:  873-933. 
1967.  American  Opisthobranch  Molluscs.  Studies  in  Tropical  Oceanography  No.  6:  Inst.  Mar.  Sci.,  Univ. 
Miami,  Florida  pp.  i-viii  +  1-256,  1  pi. 
Morch,  O. 

1863.  Contributions  a  la  faune  malacologique  des  Antilles  Danoises.  J.  Conchyl.  (3)  11:  21-43. 
Orbigny,  A.  D. 

1835-46.  Voyage  dans  I'Amerique  Meridionale.  Mollusques  pp.  1-758,  pis.  1-85. 

1839.  Mollusques.  echinodermes,  foraminiferes  et  polypiers,  recueillis  aux  lies  Canaries  par  Mm.  Webb  et 
Berthelot.  Mollusques  1.  2(2):  1-117,  pis.  1-28. 
Pilsbry,  H.  A.  and  E.  G.  Vanatta 

1902.  Papers  from  the  Hopkins  Stanford  Galapagos  Expedition,  1898-1899.  XIII,  Marine  Mollusca.  Proc. 
Washington  Acad.  Sci.  4:  549-560. 
Pruvot-Fol,  A. 

1951.  Etudes  des  nudibranches  de  la  Mediterannee.  2.  Arch.  Zool.  Exper.  Gen.  Paris  88:  1-80.  pis.  1-4. 
Quoy,  J.  R.  and  Gaimard,  J.  P. 

1832.  Voyage  de  decouvertes  de  I' Astrolabe  .  .  .  pendant  les  annees  1826-1829  sous  le  commendemcnt  de 
M.  J.  Dumoni  D"Urville,  Zool.  2:  1-686,  pis.  1-26  (1833). 


216 


Risbec,  J. 

1928.  Contribution  a  I'etude  des  nudibranches  Neo-Caledoniens.  Faune  des  Colonies  Francaises  2(1):  1-328, 
pis.  1-12. 
Smith,  E.  A.  ♦• 

1884.  In:  MoUusca.  Report  on  the  zoological  collections  made  in  the  Indo-Pacific  Ocean  during  the  voyage 
of  H. M.S.  'Alert'  1881-1882.  Trustees  of  the  British  Museum.  London,  pp.  34-116,  pis.  4-7. 
Sphon,  G.  G.  and  D.  K.  Mulliner 

1972.  A  preliminary  list  of  known  opisthobranchs  from  the  Galapagos  Islands  Collected  by  the  Ameripagos 
Expedition.  Veliger  15(2):  147-152. 
Steams,  R.  E.  C. 

1893.  Scientific  results  of  explorations  by  the  U.S.  Fish  Commission  steamer  Albatross.  No.  XXV.  -Report 
on  the  mollusk-fauna  of  the  Galapagos  Islands  with  descriptions  of  new  species.  Proc.  U.S.  Natl. 
Mus.  16(942):  353-450,  pis.  51,  52. 
White,  K.  M. 

1952.  On  a  collection  of  molluscs  from  Dry  Tortugas,  Proc.  Malacol.  Soc.  London  29(2-3):  106-120,  pi.  6. 


Contribution  No.  153  of  the  Charles  Darwin  Foundation  for  the  Galapagos  Islands. 

Mulliner:  Department  of  Marine  Invertebrates,  Natural  History  Museum,  San  Diego, 
California  92112. 

Sphon:  Invertebrate  Zoology,  Los  Angeles  County  Museum  of  Natural  History,  Los 
Angeles,  California  90007. 


SAN 


MUS.  COMP.  ZOOL. 
LIBRARY 

SEP  Q      10"'/1 

HARVARD 
UNiVERSITY 


THE  DISTRIBUTION  AND  ECOLOGY  OF  MARINE  BIRDS 
OVER  THE  CONTINENTAL  SHELF  OF  ARGENTINA 
IN  WINTER 


JOSEPH  R.  JEHL,  JR. 


0^ 


I 


TRANSACTIONS 


OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 


28  JUNE  1974 


VOL.  17,  NO.  16 


THE  DISTRIBUTION  AND  ECOLOGY  OF  MARINE  BIRDS 
OVER  THE  CONTINENTAL  SHELF  OF  ARGENTINA 
IN  WINTER 

JOSEPH  R.  JEHL,  JR. 


ABSTRACT. — Quantitative  data  on  the  distribution  and  abundance  of  marine  birds  in  winter  were 
obtained  on  three  transects  of  the  coastal  shelf  of  Argentina  in  1971  and  1972.  On  the  basis  of  avifaunal 
assemblages,  the  shelf  waters  can  be  divided  into  two  zones,  the  boundary  occurring  near  the  southern 
edge  of  Golfo  San  Jorge  (47° S).  Spheniscus  magellanicus,  Puffinus  griseus.  and  Sterna  hirundinacea 
were  characteristic  of  the  northern  zone;  Fulmarus  glacialoides.  Eudyptes  crestatus,  Pelecanoides  magel- 
lani,  Diomedea  exulans/ epomophora.  and  Pachyptila  sp.  of  the  southern  zone.  Beyond  the  continental 
shelf  off  northern  Argentina  the  avifauna  was  similar  to  that  over  the  southern  shelf,  but  included  several 
additional  species:  Garroidia  nereis.  Procellaria  cinereus.  and  Pelecanoides  urinatrix. 

Winter  sea  bird  populations  along  the  coast  of  Argentina  appear  to  be  established  by  mid-June  and 
to  remain  stable  through  the  winter.  In  general,  abundances  seemed  low  and  concentrations  were  found 
mainly  in  areas  of  strong  mixing  or  upwelling.  The  winter  census  data  are  compared  with  those  from  a 
brief  summer  transect.  Despite  pronounced  shifts  in  the  ranges  of  individual  species,  there  was  little 
seasonal  difference  in  total  sea  bird  abundance.  A  preliminary  ecological  study  indicated  that  the  bulk  of 
the  sea  bird  biomass  through  the  entire  year  is  contributed  by  large  species  that  obtain  their  food  from 
the  upper  meter  of  the  sea,  mainly  by  surface  seizing.  However,  marked  seasonal  and  latitudinal  differ- 
ences in  patterns  of  resource  utilization  appear  among  divers,  plungers,  and  filter  feeders. 

Circumstantial  evidence  suggests  that  oil  pollution  is  a  major  cause  of  sea  bird  mortality,  particularly 
over  the  northern  shelf. 


Although  sea  birds  are  among  the  most  conspicuous  inhabitants  of  the  ocean,  their 
role  in  marine  ecosystems  has  received  little  attention  (Ashmole,  1971).  Basic  information 
such  as  population  sizes  at  different  seasons,  species  composition  of  sea  bird  flocks,  and 
periods  of  migration  are  prerequisites  for  ecological  analysis.  While  such  data  are  slowly 
beginning  to  accumulate  (e.g.,  King,  1970),  they  are  inadequate  if  not  entirely  lacking  for 
most  parts  of  the  world. 

In  the  austral  winters  of  1971  and  1972,  the  R/V  Hero,  a  research  vessel  of  the 
National  Science  Foundation  was  engaged  in  oceanographic  research  along  the  coast  of 
Argentina  (Cummings  et  al.,  1971;  Jehl,  1973b).  One  objective  was  to  obtain  data  on  the 
distribution,  abundance,  and  ecology  of  sea  birds  over  the  continental  shelf.  General  pat- 
terns of  sea  bird  distribution  in  this  area  have  been  presented  by  Murphy  (1936), 
Escalante  (1970).  and  others  (references  in  Cooke  and  Mills,  1972).  Watson  et  al.  (1971) 
have  mapped  the  distribution  of  those  antarctic  and  subantarctic  species  that  occur  there. 
Yet.  detailed  data  are  scarce  and  pertain  mostly  to  observations  made  in  spring,  summer, 
or  autumn.  With  the  exception  of  Cooke  and  Mills'  (1972)  report  on  a  brief  summer 
transect  between  Buenos  Aires  and  Tierra  del  Fuego  there  seem  to  be  no  precise  quanti- 
tative data  for  shelf  waters  at  any  season.  Tickell  and  Woods  (1972)  discussed  sea  bird 
abundance  between  Montevideo,  Uruguay,  and  the  Falkland  Islands  on  the  basis  of  17 
transects  in  the  period  November-May  1954-64;  however,  their  transect  route  was  largely 
beyond  the  continental  shelf  and  their  quantitative  data  are  too  simplified  for  detailed 
analysis. 

This  paper  deals  mainly  with  quantitative  data  obtained  on  three  transects  of  the 
Argentine  coastal  shelf:  one  in  June  1971  between  the  Strait  of  Magellan  and  Bahi'a 
Blanca;  the  second  in  July  1971  on  the  return  voyage;  and  the  third  in  July-August  1972 
between  Buenos  Aires  and  the  Strait  of  Magellan.  The  dates  of  the  several  transect  periods 
were  far  enough  apart  that  distributional  changes  through  the  austral  winter  could  be 
determined.  The  present  data  with  those  of  Cooke  and  Mills  (1972)  also  allow  a  prelimi- 
nary comparison  of  summer  and   winter  differences   in  abundance,   distribution,    and 

SAN  DIEGO  SOC.  NAT.  HIST..  TRANS.  17  (16):  217-234.  28  JUNE  1974 


218 


ecological  impact  of  the  sea  bird  fauna. 

The  sun'ey  area. — The  coast  of  Argentina  is  bordered  by  a  broad,  shallow  continental 
shelf,  which  extends  offshore  for  about  180  km  at  the  latitude  of  Mar  del  Plata,  500  km 
near  Bahia  Blanca,  and  800  km,  to  the  Falkland  Islands,  off  the  Strait  of  Magellan.  Over 
most  of  the  shelf  depths  are  less  than  60  fathoms,  and  along  the  transect  routes  depths 
over  40  fathoms  were  uncommon. 

The  shelf  waters  are  derived  from  the  subantarctic  waters  of  the  Falkland  (Malvinas) 
Current,  which  flows  northward  along  the  edge  of  the  continental  slope.  They  can  be  sep- 
arated into  two  zones.  South  of  Golfo  San  Jorge  (ca.  47°  S)  strong  westerly  winds  prevail 
for  most  of  the  year,  forcing  surface  waters  seaward,  and  causing  upwelling.  In  that  area 
surface  waters  are  cold  and  rich  in  dissolved  oxygen,  nitrates,  and  phosphates.  North  of 
Golfo  San  Jorge,  surface  waters  are  warmer  and  levels  of  dissolved  nutrients  are  much 
lower.  Beyond  the  continental  shelf  off  northern  Argentina,  the  Falkland  Current  brings 
subantarctic  waters  into  sharp  juxtaposition  with  the  warmer  shelf  water.  Although  condi- 
tions there  are  similar  to  those  prevailing  over  the  southern  shelf,  it  is  useful  to  recognize 
a  third  zone  beginning  about  30  km  landward  of  the  continental  slope. 

The  northern  terminus  of  the  Falkland  Current  varies  seasonally.  In  August- 
September  waters  beyond  the  continental  slope  retain  a  subantarctic  character  to  about 
36°30'  S.  There  they  meet  and  mix  with  warmer  waters  moving  northeastward  off  the 
continental  shelf  and  with  subtropical  waters  of  the  southward-flowing  Brazil  Current. 
This  area  of  confluence,  which  is  often  realized  near  the  mouth  of  the  Rio  de  la  Plata, 
creates  rich  feeding  conditions  for  a  wide  variety  of  sea  birds  (Murphy,  1936;  Escalante, 
1970;  Cooke  and  Mills,  1972).  There  is  a  pronounced  faunal  shift  there,  with  warm  water 
species  reaching  their  southern  limits  and  cold  water  species  their  northern  limits  over  the 
shelf. 

Detailed  oceanographic  information  on  the  region  may  be  obtained  in  the  extensive 
series  of  "Pesqueria"  reports  (Aragno,  1968;  Valdez,  1969;  Villanueva,  1969-1971).  Cooke 
and  Mills  have  summarized  some  of  these  data  that  pertain  to  the  summer  months. 

CRUISE  TRACKS  AND  METHODS 

In  1971  the  Hero  departed  Punta  Arenas,  Chile,  on  11  June  and  proceeded  northward 
over  the  continental  shelf  of  Argentina  to  Bahi'a  Blanca,  arriving  on  25  June  (Fig.  1).  In 
general  the  transect  route  lay  16  to  40  km  offshore,  although  we  cruised  within  several  km 
of  the  beach  in  Golfo  San  Jorge  and  Golfo  Nuevo.  Observations  were  made  in  Golfo  San 
Jose'  on  22  and  23  June.  We  left  Bahi'a  Blanca  on  28  June  for  Golfo  San  Jose,  remaining 
inside  the  gulf  until  8  July.  After  a  port  call  in  Puerto  Madryn  we  proceeded  southward  on 
12  July  along  a  route  similar  to  that  of  the  northward  transect,  except  for  crossing  Golfo 
San  Jorge  near  its  mouth.  The  cruise  terminated  in  Punta  Arenas  on  16  July. 

In  1972  the  Hero  left  Buenos  Aires,  Argentina,  on  26  July.  Between  28  and  30  July  we 
cruised  slowly  southwestward  between  37°07'  S  and  41°40'  S,  mostly  over  deep  water 
beyond  the  continental  shelf  but  occasionally  zigzagging  over  the  edge  of  the  shelf.  Late  on 
30  July  we  re-entered  shelf  waters  and  headed  to  Puerto  Madryn,  arriving  on  1  August. 
Late  on  3  August  we  departed  for  Golfo  San  Jose,  where  we  spent  the  period  4  to  19 
August.  Following  a  port  call  in  Bahi'a  Blanca,  we  departed  for  Punta  Arenas  on  22 
August,  arriving  there  on  30  August.  The  route  was  similar  to  that  of  the  southward 
transect  in  1971,  except  that  most  of  25  August  was  spent  in  Bahi'a  Concepcion  and  27 
August  in  Bahi'a  de  los  Nodales. 

In  1971  I  made  censuses  as  often  as  possible,  except  when  the  ship  was  at  anchor. 
The  duration  of  the  observations  depended  upon  weather  conditions  and  ship's  activities, 
and  varied  from  2  to  7  hours  per  day.  In  1972,  with  the  assistance  of  Jon  P.  Winter,  it  was 
possible  to  monitor  bird  populations  almost  continously.  Most  observations  were  made 
from  a  flying  bridge  7  m  above  the  waterline,  affording  good  visibility  in  all  directions.  All 
birds  were  counted,  but  for  ship-following  species  the  maximum  numbers  were  estimated 
hourly.  In  1971,  daily  counts  were  totalled,  whereas  in  1972,  for  increased  precision,  they 
were  divided  into  morning  and  afternoon  components.  Surface  water  temperatures  were 
taken  regularly  except  when  sea  conditions  precluded  work  on  deck.  Quantitative  data. 


219 


Figure  1.  Cruise  track  of  the  R/V  Hero  along  the  coast  of  Argentina  during  transects  reported  in  this  study: 
1972  (left),  1971  (right). 

precise  localities,  and  sea  surface  temperatures  are  given  in  Tables  1  and  2. 

In  this  paper  I  consider  only  those  species  that  regularly  occur  over  the  open  ocean, 
or  more  than  about  5  km  from  shore.  Information  on  Golfo  San  Jose'  is  presented  else- 
where (Jehl,  RumboU,  and  Winter,  1973).  Specimens  obtained  in  these  studies  are  depos- 
ited in  the  Natural  History  Museum,  San  Diego. 

In  the  following  species  accounts  I  follow  the  generic  classification  of  Procellariiformes 
of  Alexander  et  al.  (1965).  Otherwise,  classification  and  common  names  largely  follow 
Meyer  de  Schauensee  (1966).  The  exceptions  involve  my  strong  preference  for  retaining 
the  traditional  whalers'  names  for  certain  species.  In  my  opinion  the  use  of  such  prosaic 
names  as  Gray  Petrel  for  Pediunker  and  White-chinned  Petrel  for  Shoemaker  has  little,  if 
anything,  to  recommend  it. 


ANNOTATED  LIST  OF  SPECIES 

Rockhopper  Penguin  (Eudyptes  crestatus). — Rockhopper  Penguins  follow  the 
Falkland  Current  north  to  Uruguay  in  winter  (Escalante,  1970).  They  seem  restricted  to 
the  cold,  deep  waters  beyond  the  continental  shelf,  and  may  be  much  commoner  off 
northern  Argentina  than  the  literature  suggests.  Groups  of  up  to  15,  mostly  adults,  were 
common  between  36-40°  S  in  late  July  1972;  the  maximum  concentration  was  79/hour. 
Over  the  continental  shelf,  however,  Rockhoppers  were  rare  or  absent.  The  only  sighting 
in  1971  was  of  a  single  bird  near  40°  S  on  15  July.  In  1972  a  few  appeared  at  the  southern 
edge  of  Golfo  San  Jorge,  where  surface  temperatures  dropped  sharply  to  4.4°  C,  but  none 


220 


were  seen  in  colder  waters  farther  south.  In  summer  Cooke  and  Mills  (1972)  recorded  only 
one  Rockhopper  over  the  shelf  near  52°40'S. 

Magellanic  Penguin  (Spheniscus  mugellanicus). — In  summer  Magellanic  Penguins 
are  common  in  the  vicinity  of  nesting  colonies  in  southern  Argentina  (i.e.,  south  of  44°  S); 
in  winter  they  largely  abandon  these  areas  and  move  northward  as  far  as  Uruguay  and 
southern  Brazil.  We  found  them  fairly  common  between  Buenos  Aires  and  the  Valdes 
Peninsula,  uncommon  to  rare  southward;  in  all  areas  their  local  abundance  was  markedly 
reduced  by  turbid  water.  All  sightings  were  made  in  shelf  waters,  mostly  within  30  km  of 
shore  in  areas  where  surface  temperatures  exceeded  9°  C.  On  all  three  transects  we  found 
concentrations  15  km  off  the  Valdes  Peninsula;  maximum  densities  were  27/hour.  The 
largest  concentration,  300  birds,  150  km  east  of  the  peninsula  on  31  July  1972,  was  near 
the  area  where  Cooke  and  Mills  found  large  flocks  in  summer.  Penguin  flocks  were  usually 
accompanied  by  South  American  Terns  {Sterna  hirnndinacea)  and  Sooty  Shearwaters 
(Piifjinus  griseus),  which  feed  on  fish  that  penguins  drive  to  the  surface.  This  penguin- 
tern-shearwater  assemblage  is  the  most  conspicuous  and  characteristic  avian  grouping 
over  northern  shelf  waters. 

Wandering  Albatross  (Diomedea  exulans),  Royal  Albatross  (D.  epomophora) . — Rare 
over  the  northern  shelf  but  slightly  commoner  farther  south.  In  1971  we  saw  occasional 
great  albatrosses  as  far  north  as  the  Valdes  Peninsula  but  the  only  concentration,  20  birds, 
was  in  Bahi'a  Grande  on  15  July.  Small  numbers  near  the  Valdes  Peninsula  and  in  Golfo 
San  Matias  in  late  August  1972  indicate  a  northward  shift  of  the  population  later  in  win- 
ter. Beyond  the  continental  shelf  great  albatrosses  were  fairly  common  from  36-40°  S. 
They  appeared  as  soon  as  the  ship  crossed  into  deep  water  and  often  outnumbered  the 
Black-brows.  Their  abundance  declined  immediately  as  we  re-entered  the  shelf  waters  and 
en  route  to  Puerto  Madryn  none  was  seen  closer  to  land  than  200  km. 

On  29  July  1972  we  saw  over  110  great  albatrosses,  3  of  which  were  color  banded;  30 
were  with  a  large  tlock  of  Black-brows  at  39°22'  S;  50  more  along  with  other  seabirds  fed 
on  scraps  from  a  fishing  trawler;  and  another  30  were  scattered  along  the  route.  Most  of 
the  birds  in  the  first  group  of  30  were  photographed;  of  these,  at  least  4  were  Royals  (dark 
line  on  tomium  visible)  and  20  were  Wanderers.  Sight  records  suggest  a  similar  species 
composition  in  the  other  groups.  No  Royals  were  identified  over  shelf  waters  in  1972, 
although  two  birds  in  1971  were  thought  to  be  epomophora  (Cabo  Danoso,  15  July;  Bahi'a 
Engano,  18  June). 

Robertson  and  Kinsky  (1972)  showed  that  large  numbers  of  Royal  Albatrosses  use  the 
southwestern  Atlantic  as  a  major  feeding  area,  particularly  in  winter.  However,  their  sug- 
gestion (following  Dabbene,  m  Murphy,  1936)  that  it  is  the  common  species  of  great 
albatross  there  is  questionable.  The  present  data  indicate  that  Wanderers  greatly  out- 
number Royals  throughout  the  winter,  in  coastal  as  well  as  offshore  waters.  Robertson  and 
Kinsky  (1972)  also  found  that  about  55  per  cent  of  the  Royals  wintering  in  the  south- 
western Atlantic  are  three  years  old  or  less  and  about  70  per  cent  are  four  or  less.  Wan- 
dering Albatrosses  of  those  ages  retain  considerable  brown  in  their  plumage  and  should  be 
distinguishable  in  the  field  (see  Tickell,  1968:  fig.  12).  Yet,  only  one  of  over  145  great 
albatrosses  observed  in  this  study  was  in  the  brown  juvenile  plumage  of  exulans;  four  were 
in  the  adult  ''chionoptera'  stage  of  exulans;  and  the  rest  were  in  plumages  in  which  the 
two  species  are  usually  indistinguishable.  If  my  estimates  of  relative  abundance  are  accu- 
rate, it  would  appear  that  Wanderers  wintering  off  Argentina  average  several  years  older 
than  Royals  in  the  same  are.  This  is  a  potentially  important  biological  difference  between 
these  similar  species  that  requires  confirmation.  Furthermore,  since  mottled  immatures  of 
exulans  were  fairly  common  along  the  coast  of  Chile  in  the  winter  of  1970  (pers.  obs.),  the 
average  age  of  Argentine  Wanderers  may  be  greater  than  that  of  birds  wintering  off  the 
Pacific  coast  of  South  America. 

Black-browed  Albatross  (Diomedea  melanophris) . — Common  to  abundant  along  the 
entire  coast,  except  where  waters  are  excessively  turbid.  In  both  years  it  was  rather  regu- 
larly distributed  over  the  shelf  north  to  Bahia  Blanca,  and  concentrations  were  found  off 
the  Valdes  Peninsula  and  in  Golfo  San  Jorge,  although  in  1971  the  largest  numbers 
(150/hour)  were  seen  in  Bahi'a  Grande.  No  important  seasonal  or  yearly  differences  in 


221 


Figure  2.   Part  of  a  tlock  of  10,000  Black-browed  Albatrosses  and  other  sea  birds.  Coast  of  Argentina.  39°22'S. 
55°22'W,  29  July  1972. 


distribution  were  evident.  Tickell  and  Woods  (1972)  did  not  observe  seasonal  differences 
in  abundance  on  transects  between  Montevideo  and  the  Falkland  Islands. 

The  species  was  also  common  —  and  once  spectacularly  abundant  —  near  the  edge 
of  the  continental  shelf.  On  the  morning  of  29  July  1972  an  estimated  10,000  were  feeding 
with  other  sea  birds  and  a  large  pod  of  Pilot  Whales  {Globicephala  melaetia)  near  39°22' 
S,  55°22'  W  at  the  edge  of  the  shelf  (Fig.  2).  That  afternoon  an  additional  10.000  were 
feeding  on  scraps  from  a  large  trawler.  Over  the  entire  route  it  appeared  that  adults  out- 
numbered immatures  by  about  5  to  1,  although  immatures  seemed  more  likely  to  occur  in 
near-shore  waters  and  mouths  of  bays.  Many  birds  near  Buenos  Aires  were  heavily  oiled. 

Giant  Petrel  [Macronectes  gigatiteus). — Widespread  and  remarkably  uniformly  dis- 
tributed in  shelf  and  offshore  waters  through  the  year,  though  commoner  in  winter. 
Usually  three  or  four  followed  in  our  wake.  Over  500  were  scavenging  offal  near  a  fishing 
ship  at  39°40'  S  on  29  July  1972. 

In  1971  approximately  70  per  cent  of  the  birds  seen  were  immatures,  whereas  in  1972 
adults  slightly  outnumbered  immatures  over  the  continental  shelf.  This  age  distribution 
suggests  a  possible  influx  of  adults  later  in  the  winter.  Beyond  the  continental  shelf 
immatures  composed  over  70  per  cent  of  the  flocks,  and  in  harbors  and  waters  very  close 
to  shore  they  predominated  strongly.  Only  3  white-phased  birds  were  encountered,  one 
with  a  huge  flock  of  sea  birds  at  39°22'  S  in  1972,  and  two  well  inside  Golfo  Nuevo  in 
1971.  No  birds  suspected  of  being  Macronectes  halli  were  among  the  giant  petrels  flying 
near  the  ship  (see  Bourne  and  Warham,  1966,  for  characters  that  may  allow  these  similar 
species  to  be  identified  under  field  conditions). 

Southern  Fulmar  [Fulmtirus  glacialoides). — Common  to  abundant  over  southern 
shelf  waters  in  winter,  in  waters  colder  than  7°  C.  In  June  and  July  1971,  Southern 
Fulmars  were  common  to  about  49°  S,  but  disappeared  abruptly  in  warmer  waters  to  the 


222 


north.  On  the  southward  transect  in  August  1972  a  few  appeared  at  44°  S,  where  temper- 
atures dropped  under  7°  C,  but  none  was  seen  again  until  47°24'  (4.4°  C).  Fulmars  were 
uncommon  but  regular  beyond  the  continental  shelf  at  36-40°  S;  surface  temperatures 
there  were  less  than  6.7°  C.  Cooke  and  Mills  did  not  record  this  species  on  their  summer 
transect. 

Cape  Pigeon  {Daption  capense). —  Cooke  and  Mills  did  not  observe  this  species 
during  their  cruise.  In  June  1971  it  was  widely  distributed  but  uncommon;  in  July  1971  it 
was  seemingly  commoner,  especially  in  the  south;  and  in  August  1972  it  was  common 
over  much  of  the  shelf  and  in  offshore  waters.  These  data  suggest  a  shift  northward  as  the 
winter  progresses.  It  occurred  in  greatest  abundance  beyond  the  continental  shelf  on  29 
July  1972,  where  flocks  of  4000  and  6000  were  in  association  with  albatross  fiocks. 

Whale-birds  or  Prions  {Pachyptila  spp.). —  In  1971  scattered  prion  flocks  were  seen 
between  San  Julian  and  the  Valdes  Peninsula,  and  in  Golfo  San  Jose.  The  largest  concen- 
tration (up  to  150/hr.)  occurred  off  Rio  Chico  on  15  July.  In  August  1972  they  were 
uncommon  to  rare  over  shelf  waters,  except  inside  Golfo  San  Jose  (Jehl  et  al.,  1973). 
Prions  were  somewhat  commoner  offshore,  especially  near  41°40'  S,  where  we  found 
scattered  flocks  of  10-15  birds.  Although  P.  desolata  and  P.  belc fieri  are  said  to  occur  in 
this  general  area  (Escalante,  1970),  the  only  specimens  we  obtained  were  belcheri  (cT, 
109  g,  37°22'  S,  54°24'  W;  cf,  4r38'  S,  56°43"  W;  ??,  Golfo  San  Jose).  In  summer 
Cooke  and  Mills  observed  prions  only  south  of  50°  S,  near  presumed  breeding  grounds. 

Pediunker  {Procellaria  ciriereus). —  A  single  bird  made  several  passes  near  the  ship 
on  30  July  1972,  when  we  set  out  a  chum  slick  well  offshore.  This  was  our  only 
observation  of  the  species,  which  appears  to  avoid  shelf  waters  at  all  seasons.  Not 
recorded  by  Cooke  and  Mills. 

Shoemaker  (Procellaria  aequinoctialis). —  In  June  1971,  Shoemakers  were 
widespread  though  generally  uncommon  along  the  entire  coast,  whereas  a  month  later 
they  were  virtually  absent  south  of  43°  S.  In  August  1972,  too,  they  were  uncommon  in 
coastal  waters  north  of  44°  S.  and  much  rarer  to  the  south.  In  both  years  concentrations 
occurred  in  waters  adjacent  to  the  Valdes  Peninsula. 

Beyond  the  continental  shelf  Shoemakers  replaced  Sooty  Shearwaters  as  the 
dominant,  and  usually  only,  species  of  shearwater,  though  they  appeared  to  be  no 
commoner  there  than  in  coastal  waters.  The  limited  data  hint  that  this  species  may  be 
more  abundant  in  summer  than  in  winter. 

Greater  Shearwater  [Puffmus  gravis). —  Common  to  abundant  in  summer  but  rare  or 
absent  in  winter,  when  the  species  occurs  in  the  North  Atlantic.  Our  only  winter 
observations  were  in  mid-June  1971:  four  between  43°40'  S  and  42°00',  and  several  in 
Golfo  San  Jose.  These  were  apparently  late  stragglers  on  the  northward  migration. 

Sooty  Shearwater  (Puf'finus  griseus). —  In  winter  Sooties  are  largely  restricted  to  shelf 
waters  north  of  45°  S;  their  distribution  seems  to  be  strongly  affected  by  surface 
temperatures  for  they  are  rare  in  waters  cooler  than  9°  C.  On  the  northward  transect  in 
June  1971  none  were  seen  south  of  43°40'  S  (9.5°  C),  but  farther  north  they  were  common 
to  abundant,  particularly  near  the  Valdes  Peninsula  (maximum,  375/hr.).  On  the 
southward  transect  in  July  they  seemed  rarer.  Only  scattered  individuals  were  seen 
between  the  Valdes  Peninsula  and  Golfo  San  Jorge  and  the  only  bird  seen  farther  south 
(47°35'  S)  was  sick  and  emaciated  (specimen,  weight  563  g). 

A  similar  distribution  was  observed  in  August  1972,  with  concentrations  off  the 
Valdes  Peninsula  and  the  northeastern  corner  of  Golfo  San  Matias  (maximum  800/hr.), 
and  in  the  mouths  of  the  larger  bays.  Although  some  birds  occurred  as  far  south  as  the 
Strait  of  Magellan,  they  were  uncommon  south  of  45°  S.  One  hundred  and  fifty  km  off 
the  Valdes  Peninsula  we  observed  600  with  large  flocks  of  South  American  Terns  and 
Magellanic  Penguins.  Sooties  were  virtually  absent  from  waters  beyond  the  continental 
shelf. 

Manx  Shearwater  {Pujfinus  pujfimis). —  This  northerm hemisphere  migrant  winters 
commonly  off  the  northern  coast  of  Argentina  (Cooke  and  Mills,  1972),  but  leaves  the 
area  in  the  austral  winter.  Our  only  sightings  were  in  1972;  one,  50  km  S.  of  the  coast  of 
Uruguay,  27  July;  and  two  in  Golfo  San  Matias,  23  August. 


223 


Wilson's  Storm-Petrel  (Oceanites  occanicus). —  In  both  years  Wilson's  Storm-Petrels 
were  rare  over  the  continental  shelf  between  the  Strait  of  Magellan  and  Bahia  Blanca, 
and  the  only  area  of  local  abundance  (maximum  23/hr)  was  off  the  Valdes  Peninsula. 
Nearly  all  of  our  observations  were  made  north  of  44°  S,  where  surface  temperatures 
exceeded  9°  C.  The  similar  distribution  patterns  found  in  all  three  transects  indicate  that 
migration  is  largely  completed  by  June.  These  petrels  were  widespread  but  still 
uncommon  in  colder  waters  (<6°  C)  beyond  the  continental  slope,  Twenty- tlve,  in  a  small 
area  225  km  SE  of  Mar  del  Plata  on  29  July  9172,  comprised  the  only  significant 
concentration.  Surprisingly,  Wilson's  Storm-Petrel  seems  even  rarer  in  shelf  waters  in 
summer.  Cooke  and  Mills  saw  only  a  single  storm-petrel  (sp.?)  during  their  transect. 

Gray-backed  Storm-Petrel  (Garrodia  nereis). —  This  species  was  not  observed  by 
Cooke  and  Mills  (1971),  and  Escalante  (1970)  does  not  include  it  in  his  compilation.  We 
saw  only  one  bird  over  shelf  waters,  37  km  offshore  at  48°59'  S  on  15  June  1971.  Beyond 
the  shelf,  on  30  July  1972,  we  saw  tlve  birds  and  collected  one  (cr*,  wt.  33  g)  near  41°38'  S, 
56°43'  W.  They  were  associated  with  a  tlock  of  six  Wilson's  Storm-Petrels.  These  appear 
to  represent  the  northernmost  records  of  the  species  (cf.  Watson  et  al.,  1971;  Olrog, 
1958),  which  nests  on  the  Falkland  Islands. 

Megallanic  Diving-Petrel  (Pclccanoides  magellani) . —  This  was  the  only  species  of 
diving-petrel  that  could  be  identified  in  the  shelf  waters  of  southern  Argentina,  and  all 
observations  there  are  referred  to  it.  In  each  year  it  occurred  to  Golfo  San  Jorge,  which  is 
farther  north  than  the  range  given  by  Meyer  de  Schauensee  (1966),  though  it  was  regular 
only  south  of  49°  S  and  common  to  abundant  only  between  the  mouth  of  the  Rio  Chico 
and  the  Strait  of  Magellan.  The  largest  concentration  (85/hr.)  was  found  in  Bahia 
Grande  on  15  July  1971.  Although  these  diving-petrels  seem  to  prefer  waters  colder  than 
7°  C,  we  found  no  seasonal  or  yearly  differences  in  distribution  even  though  quite 
different  water  temperatures  prevailed  in  the  two  years.  In  summer  Cooke  and  Mills  saw 
a  few  diving-petrels,  presumably  magellani,  near  Bahia  Grande. 

Subantarctic  Diving-Petrel  (Pelecanoides  urinatrix). —  This  species,  which  nests  on 
the  Falkland  Islands,  ranges  north  to  Uruguay  in  winter  (Escalante,  1970).  Apparently,  it 
follows  the  Falkland  Current,  for  we  saw  scattered  diving-petrels,  all  presumably 
urinatrix,  well  offshore  in  late  July  1972.  Our  first  records  were  made  on  the  night  of  27 
July,  when  7  flew  aboard;  all  had  fed  on  small  crustaceans  8  to  10  mm  long.  Other 
sightings  were  made  between  35-42°  S,  mostly  near  the  edge  of  the  continental  shelf;  one 
bird  was  as  close  as  160  km  from  shore.  Weights:  2?:  148,  160  g;  4(^:  120,  144,  145,  145  g. 

Great  Skua  (Catharacta  skua). —  Skuas  are  rare  off  Argentina  in  summer  (Cooke 
and  Mills,  1972)  and  in  winter.  Our  few  sightings  in  1971  were  made  within  a  mile  or  so 
of  land,  generally  in  the  vicinity  of  bays  and  harbors,  and  the  only  concentration  included 
several  flocks  on  the  beach  at  San  Julian  on  14  June.  Only  9  skuas  were  seen  in  1972:  five 
off  Mar  del  Plata  on  27  July,  one  well  offshore  on  30  July,  and  three  in  near-shore  waters 
between  the  Valdes  Peninsula  and  Golfo  San  Jorge.  All  were  referable  to  C.  s.  chilensis 
except  for  one  near  Mar  del  Plata  which  was  probably  C.  s.  antarctica.  In  each  year 
several  skuas  wintered  in  Golfo  San  Jose;  some  of  these  did  not  appear  to  be  chilensis  and 
may  have  been  antarctica. 

Parasitic  Jaeger  (Stercorarius  parasiticus). —  One  record,  a  dark-phased  bird  near 
Golfo  Nuevo  on  18  June  1971. 

Kelp  Gull  {Larus  dominicanus). —  In  winter  Kelp  Gulls  disperse  widely  along  the 
Argentine  coast.  Apparently  their  post-breeding  movements  are  largely  completed  by 
June,  because  we  noted  no  important  distributional  differences  in  the  three  transects. 
Except  for  local  concentrations  in  bays  and  near  centers  of  human  habitations,  this  gull 
was  uncommon  within  30  km  of  the  coast,  and  was  virtually  absent  farther  offshore.  None 
were  seen  beyond  the  continental  shelf.  Well  over  90  per  cent  of  the  birds  were  adults. 
Cooke  and  Mills  encountered  the  species  rarely,  and  only  near  land.  Weights:  6</,  850- 
1130  (952)  g;  8$,  430  (starved),  680-1040  (865)  g. 

Brown-hooded  Gull  (Larus  maculipennis). —  Not  seen  at  sea,  except  for  three  in 
mid-Golfo  San  Matias  in  1971.  Fairly  common  in  large  harbors  north  to  Buenos  Aires. 


224 


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South  American  Tern  (Sterna  hintndinacea). —  After  the  nesting  season  South 
American  Terns  leave  southern  Argentina  and  migrate  north  to  Brazil.  They  were  rare  or 
absent  south  of  40°  S,  and  uncommon  south  of  Golfo  San  Jorge.  Farther  north  they  were 
common  to  abundant,  particularly  near  the  mouth  of  bays  and  near  the  Valdes 
Peninsula,  where  large  tlocks  were  present  each  year  (maximum  375/hr.).  We  saw  no 
terns  beyond  the  continental  shelf,  although  on  31  July  1972  we  observed  scattered  terns 
up  to  190  km  offshore,  and  140  km  offshore  1500  were  feeding  with  Sooty  Shearwaters 
and  Magellanic  Penguins.  All  seemed  to  be  hiniiiditiacea,  but  other  species  could  have 
been  overlooked.  Most  sighting  were  made  within  30  km  of  shore. 

ZONATION,  CONCENTRATIONS 

Each  of  the  three  oceanographic  zones  in  the  survey  area  contains  a  distinct  faunal 
assemblage.  Sphetiiscus  magellanicus,  Pujftnus  griseus,  Oceanites  oceanicus.  and  Sterna 
hinindinacea  were  largely  restricted  to  the  continental  shelf  north  of  Golfo  San  Jorge; 
surface  temperatures  there  were  greater  than  7°  C.  Farther  south,  particularly  south  of 
49°  S,  sea  temperatures  were  lower  and  those  species  were  rare  or  absent.  Fulmanis 
glacialoides.  Eudyptes  crestatus,  Pelecanoides  magellani  appeared  and  Diomedea 
exulans/ epomophora  and  Pachyptila  spp.  became  commoner.  In  cool  waters  beyond  the 
continental  shelf  off  northern  Argentina  the  avifauna  was  similar  to  that  of  the  southern 
shelf.  Immediately  as  we  passed  beyond  the  shelf  Puffinus  griseus,  Spheniscus 
magellanicus,  and  Sterna  hinindinacea  dropped  out  and  the  following  species  appeared 
or  occurred  in  greatly  increased  numbers:  Diomedea  exulans/ epomophora,  Fulmanis 
glacialoides,  Eudyptes  crestatus,  Pachyptila  spp.,  Garrodia  nereis,  Procellaria  cinereus, 
and  Pelecanoides  urinatrix.  Note  that  the  diving-petrel  of  deep  waters  (urinatrix)  is  not 
that  of  the  southern  shelf  (magellani). 

Precise  patterns  of  distribution  within  these  zones  are  strongly  affected  by  local 
conditions,  especially  turbidity.  Waters  in  many  near-shore  areas  (e.g.,  the  mouth  of  the 
Rio  de  la  Plata  nearly  to  Punta  del  Este;  much  of  the  north  shore  of  Golfo  San  Matias 
and  northward  within  15  km  of  shore  to  Bahia  Blanca;  the  mouth  of  the  estuary  near  San 
Julian)  are  heavily  laden  with  sediment.  This  reduces  underwater  visibility  and  precludes 
the  presence  of  divers  such  as  penguins;  it  also  reduces  feeding  opportunities  for  plungers 
such  as  terns  and  some  shearwaters.  Even  scavengers  are  scarce,  presumably  because 
increased  turbidity  also  reduces  the  size  of  tlsh  populations. 

Concentrations  of  sea  birds  were  found  in  few  localities,  and  indeed  the  general 
sparseness  of  sea  birds  over  shelf  waters  was  impressive.  Concentrations  seemed  to  occur 
mainly  in  areas  of  upwelling  or  strong  tidal  currents,  where  vertical  mixing  could  enrich 
surface  waters.  For  example,  in  each  year  flocks  of  Magellanic  Penguins,  Sooty 
Shearwaters,  and  South  American  Terns  were  at  the  mouth  of  Golfo  Nuevo  and  Golfo 
San  Jose  as  well  as  8-15  km  of  the  northeastern  corner  of  the  Valdes  Peninsula.  On  23 
August  1972  large  numbers  of  sea  birds  were  distributed  across  the  mouth  of  Golfo  San 
Matias,  but  greatest  abundances  were  realized  east  of  Punta  Rasa  and  near  the  tip  of  the 
Valdes  Peninsula.  Strong  tidal  currents  prevail  in  all  of  these  areas.  The  only  other 
significant  concentration  over  shelf  waters  consisted  largely  of  great  albatrosses. 
Black-browed  Albatrosses,  Magellanic  Diving-Petrels,  Southern  Fulmars,  and  prions  in 
Bahia  Grande  on  15  July  1971.  Surface  temperatures  there  were  anamalously  cold  (4.5° 
C),  suggesting  a  strong,  local  upwelling. 

Farther  offshore,  large  flocks  of  terns,  penguins,  and  shearwaters  were  feeding  150 
km  east  of  the  Valdes  Peninsula  on  31  July  1972.  Cooke  and  Mills  (1972)  also  found  sea 
bird  concentrations  there  and  pointed  out  that  the  area  is  rich  in  dissolved  nutrients. 

The  largest  concentrations  were  at  the  edge  of  the  continental  shelf.  On  the  morning 
of  29  July  at  39°22'  S,  55°22'  W,  we  estimated  10.000  Black-browed  Albatrosses,  30 
Royal/Wandering  Albatrosses,  4,000  Cape  Pigeons,  300  Giant  Petrels,  2  Shoemakers,  2 
Southern  Fulmars,  and  1  Sooty  Shearwater,  all  in  association  with  a  pod  of  Pilot  Whales. 
As  we  passed  through  the  flock  we  were  accompanied  by  ranks  of  50  to  100  Black-brows 
sailing  by  in  formation,  and  this  sight  was  repeated  in  all  directions  over  an  area  of 


227 

perhaps  eight  km^.  Many  of  the  birds,  particularly  the  great  albatrosses,  were  feeding  on 
white  wormlike  masses  approximately  20  cm  long,  and  on  dead  reddish  fish  (presumably 
Sehustes  or  Hclicolcnus,  Scorpaenidae).  That  afternoon  we  found  an  even  larger 
concentration,  also  at  the  edge  of  the  shelf,  near  39°40'  S,  55°35'  W.  There,  10,000- 
12,000  Black-browed  Albatrosses,  50  Royal/Wandering  Albatrosses,  6,000  Cape  Pigeons, 
and  500  Giant  Petrels  were  feeding  on  offal  from  a  large  trawler.  In  each  area  sonar 
tracings  revealed  the  presence  of  large  schools  of  fish. 

SEASONAL  DIFFERENCES 

Census  data  indicate  few  pronounced  differences  in  the  distribution  and  abundance 
of  most  species  over  the  Argentine  coastal  shelf  in  winter.  Apparently  wintering  popula- 
tions are  established  by  mid-June  and  remain  largely  stable  through  August.  To  obtain  a 
more  representative  picture  of  average  winter  conditions,  I  pooled  the  data  from  all  three 
transects.  This  procedure  reduces  bias  from  inadequate  sampling  on  individual  transects 
and  minimizes  differences  caused  by  minor  variations  in  routes.  In  Table  3  the  combined 
data  are  compared  with  those  gathered  by  Cooke  and  Mills  in  a  rapid  transect  between 
Buenos  Aires  and  Tierra  del  Fuego  in  summer.  For  convenience  the  data  are  grouped  by 
2°  increments  of  latitude.  The  data  from  each  season  are  not  so  complete  as  to  inspire 
any  great  confidence  as  to  their  general  applicability;  however,  they  are  the  only  available 
quantitative  data  and  can  be  used  to  make  preliminary  comparisons  of  summer  and 
winter  patterns. 

Differences  between  the  summer  and  winter  surveys  are  largely  interpretable  in  terms 
of  the  breeding  biology  of  particular  species.  For  example,  the  high  density  of  Speniscus 
magi'lhmicus  south  of  44°  S  in  summer  is  attributable  to  concentrations  in  the  vicinity  of 
nesting  colonies;  winter  densities  are  lower  because  the  species  disperses  widely  over  the 
northern  shelf  waters.  A  similar  pattern  of  increased  density  near  known  or  presumed 
southern  nesting  grounds  in  summer  followed  by  northward  dispersal  in  winter  is  shown 
by  Pachyptila  ssp.  (presumably  P.  belcheri  from  the  Falkland  Islands),  Eudyptes 
crestatus.  and  Pelecanoides  magellani. 

Albatrosses  and  Giant  Petrels  occupy  the  shelf  waters  year-round,  with  few 
differences  in  distribution  or  abundance.  These  species  have  long  deferred  maturity.  If 
populations  of  great  albatrosses  in  the  area  consist  largely  of  pre-breeding-age 
individuals,  as  seems  to  be  the  case  for  D.  eponiophora,  the  lack  of  large  seasonal 
differences  would  not  be  unexpected.  However,  the  higher  density  of  Macronectcs  in 
winter  may  reflect  a  post-nesting  influx  of  adults.  This  is  suggested  by  the  apparent 
increase  in  adults  in  August  1972  as  compared  with  earlier  censuses.  Large  concentrations 
of  Diomedeu  mclunophris  between  42-48°  S  in  summer  suggest  locally  rich  feeding 
conditions  that  do  not  persist  into  the  winter  months. 

Though  not  as  pronounced,  deferred  maturity  is  also  characteristic  of  smaller  Pro- 
cellariiformes  (Ashmole,  1971:  Table  2),  and  one  would  expect  some  non-breeders  of  most 
species  to  occur  in  the  area  throughout  the  year.  The  absence  of  fulmarine  petrels 
[Fulmarus  glucicdoides.  Daption  capensis)  in  summer  is  probably  attributable  to  their 
breeding  biology:  young  birds  tend  to  search  for  nesting  sites  at  colonies  several  years  in 
advance  of  active  breeding  (G.  E.  Watson,  pers,  comm.). 

Proci'Uariii  acquinoctialis  is  resident  in  the  southern  hemisphere,  nesting  in  the 
austral  summer  (Murphy,  1936:  644;.  Its  apparent  predominance  in  summer  seems 
unusual  and  may  be  due  to  concentrations  of  non-breeders  near  44°  S;  the  situation  may 
be  similar  to  that  shown  by  D.  mclunophris.  Pujfinus  gravis  was  virtually  absent  in 
winter,  having  migrated  to  the  northern  hemisphere;  its  abundance  far  from  any  known 
nesting  grounds  in  summer  presumably  indicates  a  large  population  of  non-breeding 
individuals  (see  also  Watson,  1971;  Tickell  and  Woods,  1972).  Most  Puffinus  griseus 
winter  in  the  northern  hemisphere,  but  large  numbers  occur  over  the  Argentine  shelf  all 
year.  The  limited  data  do  not  suggest  any  important  differences  in  abundance  between 
wintering  and  summering  populations  in  the  area,  but  there  is  an  obvious  shift  northward 
in  winter.  However,  much  greater  abundances  are  expected  during  periods  of  migration. 


228 


TABLE  3.  A  comparison  of  winter  (W)  and  summer  (S)  seabird  densities  over  the  continental  shelf  of  Argen- 
tina. Winter  data  are  pooled  from  three  transects  (see  text);  summer  data  are  from  one  transect  (Cooke  and 
Mills,  1972).  Abundance  indicated  is  number  of  birds  per  hour  of  observation.  Species  seen  on  fewer  than 
five  days  are  omitted  from  the  winter  sample. 


Species 


40-42  42-44 


Latitude  °S 
44-46         46-48         48-50 


50-52         52-54 


Eudyptes  crestatus 


Spheniscus  magellanicus 


Diomedea  exulans/ 
epomophora 


Diomedea  melanophris 


Macro nectes  giganteus 


Fulmarus  glacialoides 


W 
S 

w 
s 


w 
s 

w 
s 

w 
s 

w 
s 


5.4 


0.3 


3.0 


2.8 


0.3 


0.2 


0.5 


0.5 


7.5 


0.4 


41 


0.5 


5.4 

1.7 

0.5 

0.5 

0.8 

— 

— 

19.5 

10.0 

3.0 

0.8 

0.5 

0.6 

0.1 

0.1 

6.0 

0.4 

0.5 

0.7 

1.7 

0.3 

0.8 

— 

5.3 

2.9 

10.8 

9.7 

66 

2.4 

9.2 

32 

13.2 

5.5 

0.8 

3.5 

3.1 

3.6 

2.8 

6.8 

8.0 

3.1 

2.7 

2.2 

3.1 

2.1 

1.6 

1.0 

6.0 


Daption  capensis 
Pachyptila  spp. 
Procellaria  aequinoctialis 
Puff  inus  gravis 
Puffinus  griseus 
Puffinus  puffinus 


Oceanites  oceanicus 
(incl.  petrel  sp.) 


Pelecanoides  magellani 
(and  Pelecanoides  sp.) 


Catharacta  skua 


Stercorarius  parasiticus 
(and  Stercorarius  sp.) 


Larus  dominicanus 


Sterna  hirundinacea 


Hours  of  observation 


W 
S 

W 
S 

W 

s 

w 
s 

w 
s 

w 
s 


w 
s 


w 
s 

w 
s 


w 
s 

w 
s 

w 
s 

w 
s 


2.0 


2.5 


2.1 


2.6 


1.4 


1.6 


1.3  13.2 


5.4 


120 


1.1 


0.5 


2.2 


2.5 

0.8 

2.1 

8.0 

78 

4.4 

— 

— 

— 

— 

16.3 

19.0 

7.5 

0.8 

2.1 

0.7 

6.8 

5.6 

15.1 

60 

4.8 

1.5 

0.8 

0.5 

36 

284 

67 

8.3 

0.8 

0.5 

36 

18.0 

1.7 

2.4 



3.6 

5.6 

51 

0.8 

— 

— 

1.0 

2.5 
0.5 


0.1 
0.4 


0.6 
4.3 


24.8  39.6 


0.5 


0.4 


0.2 


9.2 


0.2 


0.9 


4.4 


0.2 


1.9 


0.9 


0.7  3.5  40 

6.6  3.9  2.4 


4.8 


0.2 


0.8 
27.6 


9.3 


31.0 
6.2 


16.5  17.0  13.5 

2.7  2.3  3.2 


0.8 

0.4 

0.4 


2.5 
1.2 


11.8 
2.5 


0.4 
0.5 


2.0 

5.0 
2.0 


229 


Oceanites  oceanicus  and  Catharacta  skua  were  uncommon  at  both  seasons,  though 
more  widespread  and  northerly  in  winter.  Post-breeding  northward  dispersal  in  Lams 
dominicatius  and  Sterna  hinindinacea  is  largely  responsible  for  their  predominance  in  the 
winter  censuses,  although  the  virtual  absence  of  L.  domitiicanus  in  summer  is  partly 
attributable  to  the  fact  that  Cooke  and  Mills'  route  was  farther  offshore  than  the  normal 
range  of  this  gull. 

Two  migrants  from  the  northern  hemisphere,  Pujfitius  pufjhius  and  Stercorarius 
parasiticus,  occurred  almost  exclusively  in  their  non-breeding  season,  the  austral 
summer;  at  the  latitudes  under  consideration  the  shearwater  is  very  uncommon. 

In  all  three  winter  transects  the  transition  between  the  northern  and  southern  shelf 
avifaunas  occurred  near  47°  S.  Cooke  and  Mills  suggested  that  the  transition  zone  was 
nearer  50°S  in  summer,  but  it  seems  to  occur  near  Golfo  San  Jorge  area  in  that  season  as 
well  (Table  3,  Fig.  5). 

ECOLOGICAL  CONSIDERATIONS 

Despite  their  limitations,  the  quantitative  data  are  useful  in  suggesting  questions  for 
future  research.  For  example,  do  latitudinal  or  seasonal  distributional  patterns  of  sea 
birds  suggest  corresponding  differences  in  the  productivity  of  shelf  waters.  Neither  the 
winter  nor  the  summer  data  show  any  consistent  relationship  between  latitude  and  sea 
bird  abundance  (Fig.  3),  although  in  both  seasons  the  highest  concentrations  were 
recorded  in  the  northern  half  of  the  census  area.  Seasonal  differences  in  abundance  also 
seem  minor,  as  the  summer  and  winter  curves  correspond  fairly  closely  over  most  of  the 
range.  (North  of  40°  S  the  winter  data  were  largely  gathered  beyond  the  continental 
shelf.)  Biomass  is  a  more  useful  index  to  productivity,  for  it  indicates  the  total  mass  of 


300 


200 


3 
O 

X 


100 


50 


36 


Winter 

Summer  


38  40  42  44  46 

OS  Latitude 


48 


50 


52 


Figure  3.  Sea  bird  abundance  along  the  coast  of  Argentina,  plotted  to  nearest  0°30'  of  latitude.  Winter  data 
are  pooled  from  three  transects;  summer  data  are  from  Cooke  and  Mills  (1972). 


230 


organisms  that  is  being  maintained  in  an  area.  To  obtain  this  statistic,  the  density  of  each 
species  in  Table  3  was  multipHed  by  its  average  weight  (Appendix)  and  the  resuhs  were 
summed,  giving  grams/hours/2°  increment  of  latitude.  The  data  hint  at  increased 
biomass  to  the  north  (Fig.  4),  but  they  are  strongly  biased  by  inadequate  sampling, 
particularly  at  the  higher  latitudes.  The  winter  peak  at  50-52°  S  is  especially  suspect, 
being  based  on  only  2-1/2  hours  of  observations  on  a  single  day.  In  summary,  present 
data  on  sea  bird  abundance  and  biomass  do  not  indicate  marked  seasonal  or  latitudinal 
differences  in  the  productivity  of  the  Argentine  coastal  waters.  This  conclusion  is  tentative 
and  requires  additional  study. 

A  more  interesting  question  is  how  seasonal  and  latitudinal  differences  in  species 
composition  may  affect  patterns  of  resource  utilization.  Table  4  presents  a  simplified 
ecological  classification  of  sea  birds  modified  after  Ashmole  (1971),  which  should  be 
consulted  for  details.  In  this  table,  I  have  grouped  the  avifauna  into  nine  categories  based 
on  size  of  bird,  major  feeding  behaviors,  and  food  preferences.  The  Shoemaker  is 
separated  from  the  other  shearwaters  partly  because  of  its  greater  size  and  different 
feeding  behavior,  but  mainly  because  it  is  resident  in  the  southern  hemisphere  and 
therefore  its  ecological  impact  is  expected  to  be  more  constant.  Since  total  biomass  at  any 
latitude  is  variable  (Fig.  4)  and  is  strongly  affected  by  census  errors,  the  data  have  been 
converted  to  a  percentage  basis  for  each  category.  When  these  data  are  presented 
graphically  (Fig.  5)  the  marked  change  in  the  ecological  composition  of  the  sea  bird 
community  at  46-48°  S  is  emphasized. 

In  summer,  north  of  this  area,  virtually  the  entire  biomass  is  made  up  of  large 
species  that  obtain  their  food  from  the  upper  meter  of  the  sea,  mainly  by  surface  seizing 
or  pursuit  plunging  (albatrosses,  large  and  small  shearwaters).  The  remainder  consists 
largely  of  divers  (penguins)  that  feed  on  t'lsh.  Groups  that  feed  at  least  in  part  by  filtering 
small  organisms  (fulmarine  petrels,  prions  and  storm-petrels)  are  absent.  South  of  46-48° 
S  large  surface  feeders  compose  only  50-60  per  cent  of  the  biomass,  and  there  is  a  sharp 
increase  in  the  biomass  of  divers  and  filter  feeders.  The  high  percentage  of  gulls  and 
skuas  at  48-50°  S  is  based  on  a  concentration  of  Parasitic  Jaegers.  Jaegers  typically  derive 
much  of  their  food  by  piracy;  however,  since  likely  prey  species  were  rare  or  absent  they 
may  have  been  feeding  by  surface  seizing  or  scavenging. 


40  2        42  4  44  6         46  8        48  50 

°S  latitude 


Winter    ■ 
Summer  ' 


502 


52  4 


Figure  4.  Seasonal  relationship  between  biomass  (grams/hour)  and  latitude  along  the  coast  of  Argentina, 
plotted  by  2°  increments  of  latitude.  Winter  data  are  pooled  from  three  transects,  summer  data  are  from  Cooke 
and  Mills  (1972). 


231 


TABLE  4.   A  simplified  ecological  classification  of  seabirds  (modified  from  Ashmole,  1971] 


Group 


Species 


Weight 


Major  Food 


Foraging  Behavior 


A. 

Albatrosses  and 
giant  petrels 

Diomedea  exulans 
D.  epomophora 
D.  melanophris 
Macronectes  giganteus 

Larger  than 
3000g 

Fish,  carrion, 
cephalopods 

Surface  seizing, 
scavenging 

B. 

Fulmarine  petrels 

Fulmarus  glacialoides 
Caption  capensis 

350-7  OOg 

Crustaceans, 

cephalopods, 

carrion 

Surface  seizing, 
filtering,  scavenging 

C. 

Gulls  and  skuas 

Larus  dominicanus 
Catharacta  skua 
Stercorarius  parasiticus 

500-1 500g 

Varied 

Scavenging,  surface 
seizing,  piracy 

D. 

Prions  and  storm- 
petrels 

Pachyptila  spp. 
Oceanites  oceanicus 
Garrodia  nereis 

30-1  30g 

Small  fish, 
plankton 

Filtering,  pattering 

E. 

Large  shearwaters 

Procellaria 
aequinoctialis 

1250g 

Cephalopods, 

fish, 

crustaceans 

Surface  seizing, 
pursuit  plunging 

F. 

Smaller  shearwaters 

Puffinus  gravis 
Puffinus  griseus 
Puffinus  puffinus 

400-750g 

Fish, 

cephalopods 

crustaceans 

Surface  seizing, 
pursuit  plunging 

G. 

Terns 

Sterna  hirundinacea 

200g 

Small  fish 

Plunging 

H. 

Penguins 

Spheniscus  magellanicus 
Eudyptes  crestatus 

2500-4900g 

Fish, 

cephalopods, 

crustaceans 

Pursuit  diving 

1. 

Diving-petrels 

Pelecanoides  magellani 
Pelecanoides  urinatrix 

150-160g 

Crustaceans, 
small  fish 

Pursuit  diving 

In  winter  the  transition  zone  persists  at  46-48°  S,  but  biomass  relationships  are  more 
complex.  The  proportion  of  surface  feeders  increases  from  about  75  per  cent  in  the  north 
to  90  per  cent  in  the  south.  In  the  north  this  is  divided  among  albatrosses,  large  and  small 
shearwaters,  and  gulls;  the  remainder  consists  of  diver  (penguins)  and  plungers  (terns).  In 
the  south  albatrosses  alone  make  up  approximately  75  per  cent  of  the  biomass,  the 
remainder  being  contributed  by  smaller  surface  feeders  and  filter  feeders;  the  percentage 
of  divers  is  much  reduced,  and  plungers  are  absent. 

Through  the  year  the  biomass  contributed  by  some  ecological  groups  remains  fairly 
constant.  Surface  feeders  dominate  the  shelf  waters  all  year,  and  piratical  feeders  make 
up  a  fairly  consistent  though  small  proportion  of  the  biomass.  It  is  interesting,  however, 
that  major  seasonal  shifts  in  distribution  shown  by  many  species  are  not  accompanied  by 
a  compensatory  movement  into  the  vacated  area  by  taxa  that  utilize  the  similar  foods.  For 
example,  penguins  vacate  the  southern  shelf  in  winter,  and  fulmarine  petrels,  prions,  and 
diving-petrels  shift  northward.  The  biomass  they  contributed  to  southern  waters  is  not 
replaced  by  other  divers,  small  scavengers,  or  filter  feeders  but  by  large  surface  feeders. 
Also,  terns  congregate  over  the  northern  shelf  in  winter,  an  area  that  contained  no 
plungers  in  summer.  These  shifts  may  indicate  an  increase  in  the  spectrum  of  available 
food  in  winter.  More  likely,  however,  a  wide  variety  of  foods  is  present  all  year  but  cannot 
be  exploited  in  summer  because  this  area  is  too  distant  from  nesting  colonies. 

Marine  bird  populations  beyond  the  continental  shelf. —  The  marine  avifauna 
beyond  the  continental  shelf  differs  importantly  both  in  species  composition  and  in 
relative  abundance  of  species  from  that  nearer  shore.  The  major  differences  noted  in 
winter  have  been  discussed  (p.  226).  Tickell  and  Woods  (1972)  reported  several  species  on 
transects  between  Montevideo,  Uruguay,  and  the  Falkland  Islands  from  late  spring  to 
late  autumn  that  we  did  not  find  over  the  shelf  in  winter.  These  included  Phoebetria 
palpebrata,  Pterodroma  macroptera.  Pt.  lessoni,  Pt.  incerta.  Pt.  mollis,  Halobaena 
caenilea.     Fregetta     tropica,     Fregetta     grallaria,     Stercorarius     pomarinus,     and     S. 


232 


< 

g 

CO 


402 


468  480 

°S    Latitude 


44  6 


48  0 


50-2 


Figure  5.  Biomass  relationships  ot'seabirds  by  feeding  types,  along  the  coast  of  Argentina  in  winter  (upper)  and 
summer  (lower).  Groupings  comprising  less  than  1  per  cent  of  the  biomass  for  any  period  arc  not  plotted. 
A.  Albatrosses  and  giant  petrels.  B.  Fulmarine  petrels.  C.  Gulls  and  skuas.  D.  Prions  and  storm-petrels. 
E.  Large  shear\\aters.  F.  Smaller  shearwaters.  G.   Terns.  H.  Penguins.  1.  Diving-petrels. 


lon^^icaiidus.  Several  other  species  e.g.,  (Procc/laria  cincrcus.  PuJIimis  gravis,  Garrodia 
nereis)  seemed  to  be  tar  commoner  in  deep  waters  than  near  shore.  Almost  certainly,  sea 
bird  density,  latitudinal  patterns  of  abundance  and  distribution,  and  ecological  patterns 
of  resource  utilization  also  differ  significantly  between  these  areas,  but  the  only  semi- 
quantitative data  (Tickell  and  Woods,  1972)  are  insufficient  to  permit  even  preliminary 
comparison  and  analysis. 


233 


MORTALITY 

In  1972  vve  found  the  desiccated  remains  of  Magellanic  Penguins  every  30  m  or  so 
along  the  beaches  of  Golfo  San  Jose  (Jehl  et  al.,  1973);  extensive  mortality  was  also  noted 
at  Punta  Norte  and  elsewhere  on  the  Valdes  Peninsula.  Most  of  the  birds  had  been  dead 
for  a  long  time,  and  although  there  was  no  evidence  that  the  mortality  had  been  caused 
by  a  single  event,  the  majority  of  the  carcasses  were  oiled.  At  sea  it  was  not  uncommon  to 
observe  oiled  albatrosses.  Giant  Petrels,  and  Cape  Pigeons.  I  made  no  quantitative  esti- 
mates, but  the  incidence  of  oiling  was  greatest  off  northern  Argentina,  particularly  in  the 
vicinity  of  the  Rio  de  la  Plata.  This  heavily-trafficked  area  is  close  to  one  of  the  most 
important  feeding  grounds  for  sea  birds  in  the  South  Atlantic  (Murphy,  1936;  Robertson 
and  Kinsky,  1962;  Cooke  and  Mills,  1972).  In  many  miles  of  beachcombing  in  Golfo  San 
Jose,  1  found  the  remains  of  few  pelagic  birds  other  than  penguins,  and  none  that  were 
oiled.  Flying  birds  are  less  likely  than  penguins  to  amass  lethal  doses  of  oil  at  one  sitting, 
but  even  small  amounts  can  break  down  the  insulation  of  the  feather  coat  and  lead  to 
death  far  from  the  area  of  contamination.  Further,  the  pelts  of  Procellariiformes  are  less 
durable  than  the  tough  hides  of  penguins,  and  their  bodies  seem  more  likely  to  be 
devoured  by  Giant  Petrels  and  other  scavengers  before  they  can  drift  ashore.  I  suspect 
that  the  incidence  of  sea  bird  mortality  from  oil  pollution,  even  in  the  remote  reaches  of 
the  South  Atlantic,  is  more  insidious  and  pervasive  than  the  present  documentary 
evidence  indicates  (see  also  Jehl,  1975). 

ACKNOWLEDGMENTS 

The  research  was  made  possible  from  a  grant  (NSF-GV-32739)  from  the  National  Science  Foundation.  The 
assistance  and  cooperation  of  Master  Frank  Liberty,  Master  P.  J.  Lenie.  and  the  crew  of  the  R/V  Hero  is 
sincerely  appreciated.  1  am  indebted  to  Jon  P.  Winter,  W.  C.  Cummings.  J.  F.  Fish,  and  P.  O.  Thompson  for 
assistance  in  the  field,  to  Carl  L.  Hubbs  for  ichthyological  information,  and  to  George  E.  Watson  for  comment- 
ing on  a  draft  of  the  manuscript. 

LITERATURE  CITED 

Alexander,  W.  B.,  et  al. 

1%5.  The  families  and  genera  of  the  petrels  and  their  names.  Ibis  107:  401-405. 
Aragno,  F.  J. 

1%8.   Datos  y  resultados  preliminares  de  las  campafias  pesqueria:  "Pesqucria  1.  II,  III."  Mar  del  Plata, 
Argentina,  Publicaciones  del  projecto  de  dcsarrollo  pesquero  (Series  Informes  Technicos)  Nos.  10/1, 
10  II.  10/111. 
Ashmole,  N.  P. 

1971.  Sea  bird  ecology  and  the  marine  environment,  p.  223-285.  //;.   D.  S.  Farner  and  J.  R.  King  (cds.). 
Avian  Biology.  Vol.  1.  Academic  Press,  N.  Y,  and  London. 

Bourne,  W.  R.  P..  and  J.  Warham 

l%b.  Geographical  variation  in  the  Giant  Petrels  of  the  genus  Macroncctcs.  Ardea  54:  45-67. 
Cooke.  F..  and  E.  L.  Mills 

1972.  Summer  distribution  of  pelagic  birds  off  the  coast  of  Argentina.  Ibis  144:  245-251. 
Cummings.  W.  C,  J.  R.  Fish,  P.  O.  Thompson,  and  J.  R.  Jehl,  Jr. 

1971.    Bioacoustics  of  marine  mammals  off  Argentina.  R.  V.  Hero,  Cruise  71-3.  Antarctic  J.  U.  S.  b(b): 
2hf)-2b8. 
Escalante,  R. 

1970.   Aves  marinas  del  Rio  de  la  Plata.  Montevideo,  Barreiro  y  Ramos,  S.  A. 
Jehl,  J.  R.,  Jr. 

1973a.  The  distribution  of  marine  birds  in  Chilean  waters  in  winter.  Auk  90:  114-135. 

1973b.  Winter  populations  of  marine  birds  along  the  coast  of  Argentina.  Antarctic  J.  U.  S.,  7(2):  32-33. 

1975.   Mortality  of  Magellanic  Penguins  in  Argentina.  Auk  92,  in  press. 
Jehl.  J.  R..  Jr.,  M.  A.  E.  Rumboll.  and  J.  P.  Winter 

1973.  Winter  bird  populations  in  Golfo  San  Jose.  Argentina.  Bull.  Brit.  Ornith.  CI.  93:  56-63. 
King,  W.  B. 

1970.    The  trade  wind  /one  oceanographv  piKit  studv.  Part  7:  Observations  of  sea  birds  March  1964  to  June 
1965.  LI.  S.  Fish  Wildl.  Serv.  Spec.  Sci.  Rept.  Fish.  No.  586. 
Lack,  D. 

1968.   Ecological  adaptations  for  breeding  in  birds.  London,  Methuen  and  Co.,  Ltd, 
Meyer  de  Schauensee,  R. 

1966.   The  species  of  birds  of  South  America.  Narberth.  Pennsylvania.  Livingston  Publ.  Co. 


234 


Murphy,  R-  C. 

1936.  Oceanic  birds  of  South  America,  2  vols.  New  York,  Amer.  Mus.  Nat.  Hist. 
Olrog,  C.  C.  «. 

1958.  Observaciones  sobre  la  avifauna  antarctica  y  de  alta  mar  desde  el  Rio  de  la  Plata  hasta  60°  de  latitud 
sur.  Acta  Zool.  Lilloana,  15:  19-33. 
Robertson.  C.  J.  R.,  and  F.  C.  Kinsky 

1972.  The  dispersal  movements  of  the  Royal  Albatross  (Diomedea  epom'ophora).  Notornis,   19:  289-301. 
Serventy,  D.  L.,  V.  Serventy,  and  J.  Warham 

1971.  The  handbook  of  Australian  Sea-birds.  Sydney,  A.  H.  and  A.  W.  Reed. 
Tickell,  W.  L.  N. 

1968.  The  biology  of  the  great  albatrosses,  Diomedea  exulans  and  Diomedea  epomophora.  p.  1-55.  In, 
O.  L.  Austin,  Jr.  (ed.).  Antarctic  bird  studies.  Antarctic  Res.  Ser.  12.  262  p.  Washington,  D.C., 
American  Geophysical  Union. 

Tickell,  W.  L.  N.,  and  R.  W.  Woods 

1972.  Ornithological  observations  at  sea  in  the  South  Atlantic  Ocean,  1954-64.  Brit.  Antarct.  Surv.  Bull. 
31:  63-84. 

Valdes,  A.  J. 

1969.  Datos  y  resultados  de  las  campafias  pesqueria:  "Pesqueria  V."  Mar  del  Plata,  Argentina,  Publicaci- 
ones  del  projecto  de  desarrollo  pesquero  (Serie  Informes  Technicos),  No.  10/V. 

Villanueva,  S.  D.  (ed.) 

1969-71.   Datos  y  resultados  de  las  campafias  pesqueria:  "Pesqueria  VI,  VII,  VIll,  IX,  X,  XI."  Mar  del 
Plata,  Argentina.  Publicaciones  del  projecto  de  desarrollo  pesquero  (Serie  Informes  Technicos;  Nos. 
10/VI,  10/VII,  10/VIII,  10/IX,  10/X,  lO/XI. 
Watson,  G.  E. 

1971.  Molting  Greater  Shearwaters  (Pufftnus  gravis)  off  Tierra  del  Fuego.  Auk  88:  440-442. 
Watson,  G.  E.,  et  al. 

1971.  Birds  of  the  Antarctic  and  Subantarctic.  Antarctic  Map  Folio  Ser.  Folio  14,  New  York,  Amer. 
Geogr.  Soc. 


San  Diego  Natural  History  Museum,  P.O.  Box  1390,  San  Diego,  California,  92112. 


APPENDIX  I 
Weights  of  seabirds' 

Eudyptes  crestatus,  25(X)  (L).  Spheniscus  magelhnicus.  4900  (L).  Diomedea  exulans/ epomophora.  85(X)  (L, 
SSW,  SD).  Diomedea  melanophris.  3600  (SD).  Macronectes  giganteus,  30(X)  (SSW.  SD).  Fulmants  glacialoides. 
700  (J).  Daption  capensis.  350  (J).  Pac/iyplila  spp.,  130  (J).  Procellaria  aequinoctialis.  1250  (SD).  Pujfinus 
griseus.  750  (J).  Pujfinus  gravis,  650  (E).  Pujfinus  pujfinus,  400  (L).  Oceanites  oceanicus,  30  (L,  J).  Peleca- 
noides  mugelUini.  160  (J).  Pelccunoidcs  urinatrix.  150  (SD,  this  paper).  Catharacta  skua.  1400  (SD).  Stercorar- 
ius  parasiticus.  500  (L).  Lams  dominicanus,  910  (SD).  Sterna  hirundinacea.  200  (SD). 

References:  L  =  Lack,   1968,  appendix  17.  J=Jehl,   1973a.  SSW  =  Serventy,  Serventy.  and  Warham,    1971. 
SD  =  specimens  in  San  Diego  Natural  History  Museum.  E  =  estimate. 


I 


i 


MUS.  C?OMP.  ZOOL- 
LIBRARY 

HARVARD 
UNIVERSITY, 


MEXICAN  SPECIES  OF  THE  GENUS  HETERANDRIA, 
SUBGENUS  PSEUDOXIPHOPHORUS 
(PISCES:  POECILIIDAE) 


ROBERT  RUSH  MILLER 


TRANSACTIONS 


OF  THE  SAN  DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 


VOL.  17,  NO.  17 


28  JUNE  1974 


i 


J 


MEXICAN  SPECIES  OF  THE  GENUS  HETERANDRIA. 
SUBGENUS  PSEUDOXIPHOPHORUS 
(PISCES:  POECILIIDAE) 

ROBERT  RUSH  MILLER 


ABSTRACT. —  The  subgenus  Fscucld.xiphophorus  is  currently  regarded  as  monotypic.  with  a  single  wide- 
spread species,  Hctcnuulria  hiimiciihitu.  inhabiting  the  Atlantic  slope  of  Middle  America.  Actually,  the 
taxon  includes  two  sharply  distinct  species  in  Mexico,  the  more  primitive  being  H.  jonesi;  other  species  of 
Fsciuloxiphoplionis  occur  in  Guatemala.  The  basis  for  recognizing  subgenera  of  Hcterandria  is  presented 
as  well  as  a  detailed  comparison  between  H.  himaculata  and  H.  jonesi.  including  illustrations  of  gonopodia. 
gonopodial  suspensoria,  and  the  whole  fish.  Hetcruudrui  jonesi,  which  occurs  at  elevations  up  to  2,385 
meters,  is  close  to  the  ancestral  stock  of  the  genus. 

RESUMEN. —  El  subgenero  Pseudoxiphophoms.  de  acuerdo  a  la  literatura  corriente.  es  considerado  como 
monotipico.  con  una  sola  especie  de  amplia  distribucion,  Hetenindriu  himaculata.  que  habita  la  vertiente 
Atlantica  de  Mexico  y  America  Central.  Como  se  muestra  aqui,  en  Mexico  hay  dos  especies  claramente 
distintas,  la  mas  primitiva  siendo //.  jonesi:  otras  especies  de  Pseudoxiphophoms  existen  en  Guatemala.  La 
base  para  reconocer  subgeneros  de  Heterandria  se  presenta  asi  como  una  comparacion  detallada  entre  H. 
hinuiculata  y  H.  jonesi.  incluyendo  ilustraciones  de  gonopodios,  suspensores  de  gonopodio,  y  ademas  de  el 
pez  completo.  Heterandria  jonesi.  que  occure  a  alturas  hasta  2.385  metros,  es  cercano  a  el  tronco  ancestral 
de  el  genero. 

Until  the  recent  general  review  of  the  Poeciliidae  by  Rosen  and  Bailey  (1963),  Heter- 
andria and  Pseudoxiphophonts  were  regarded  as  monotypic  genera  represented,  respec- 
tively, by  H.  formosa  Agassiz  in  Florida  and  adjacent  coastal  lowlands,  and  by  P.  bi- 
maculatus  (Heckel)  from  northeastern  Mexico  southward  and  eastward  into  Nicaragua. 
Among  ichthyologists  publishing  on  Pseudoxiphophoms  during  this  century,  only  Regan 
(1904-1913)  consistently  maintained  that  this  subgenus  comprises  two  fully  distinct 
species,  although  Hubbs  (1924-1936)  divided  P.  bimaculatus  into  four  subspecies,  includ- 
ing the  one  that  is  here  restored  to  full  specific  status.  Rosen  and  Bailey  (1963:  131), 
commenting  on  Hubbs'  action,  indicated  that  the  characters  distinguishing  these  forms 
"are  apparently  clinal  and  grade  imperceptibly  from  one  race  to  another."  In  attempting 
to  distinguish  forms  oi  Pseudoxiphophoms,  overemphasis  has  been  placed  on  the  dorsal- 
ray  number  which  varies  widely  in  the  two  species  of  this  group  in  Mexico.  As  usual  in 
poeciliids,  the  detailed  architecture  of  the  gonopodium  proves  to  have  far  more  important 
systematic  value,  although  the  size  and  position  of  the  dorsal  fin  is  also  highly  useful. 
Color  pattern  helps  to  distinguish  the  Mexican  species  (Pseudoxiphophoms)  but  it  is  too 
variable  for  complete  reliance. 

Regan  was  correct  in  concluding  that  P.  Jonesi  (restricted  to  east-central  Mexico)  and 
P.  bimaculatus  are  distinct  species,  even  though  the  male  of  jonesi  was  unknown  to  him. 
As  here  shown  for  the  first  time,  the  gonopodium  of  Heterandria  Jonesi  is  consistently  and 
sharply  distinct  from  that  of  H.  bimacukita.  Although  these  species  have  overlapping 
ranges  they  are  rarely  taken  together,  as  in  central  Veracruz  (see  below).  My  paper  on  the 
zoogeography  of  Middle  American  freshwater  fishes  (Miller,  1966)  did  not  include  the 
extralimital  H.  Jonesi,  but  recognition  of  more  than  one  species  of  Pseudoxiphophoms  in 
Mexico  was  implied  in  the  range  statement  for  Heterandria  bimacukita,  since  the  sub- 
genus Pseudoxiphophoms  ranges  northward  into  southeastern  Tamaulipas. 

This  paper  presents  a  detailed  comparison  between  H.  Jonesi  and  H.  bimacukita  in 
Mexico,  illustrates  their  distinctive  gonopodia  and  suspensoria  as  well  as  the  general  form 
and  appearance  of  each  species,  and  discusses  variation  in  coloration,  body  proportions, 
and  meristic  characters.  The  distribution  of  the  two  species  is  also  treated  and  support  is 
presented  for  the  view  that  H.  Jonesi  is  closest  to  the  common  ancestor  of  the  genus. 

SAN  DIEGO  SOC.  NAT.  HIST..  TRANS  17(17):  235-250,  28  JUNE  1974 


236 


MATERIALS  AND  METHODS 

Specimens  examined  came  from  the  following  museum  collections:  BMNH,  British 
Museum  (Natural  History),  P,  Instituto  Politecnico  Nacional  (Mexico),  UMMZ,  Univer- 
sity of  Michigan  Museum  of  Zoology,  USNM,  United  States  National  Museum.  I  am 
grateful  to  P.H.  Greenwood  for  making  a  syntype  oX  MoUietiisia  jonesii  available,  to  Jose 
Alvarez  for  the  loan  of  topotypes  of  that  species,  to  William  M.  McLane  and  Brandon 
McNair  for  information  regarding  the  sympatric  occurrence  of  H.  bimaculata  and  H. 
jonesi,  to  Royal  D.  Suttkus  for  the  loan  and  exchange  of  specimens,  to  the  staff  of  the 
National  Museum  of  Natural  History  (USNM)  for  facilities  and  working  space,  and  to  the 
John  Simon  Guggenheim  Memorial  Foundation  for  support  as  a  Guggenheim  Fellow 
while  preparing  this  manuscript.  Appreciation  is  also  extended  to  Martha  B.  Lackey, 
former  staff  artist  of  the  Museum  of  Zoology,  for  the  accompanying  illustrations,  except 
Figure  4,  drawn  by  Patricia  J.  Wynne,  current  staff  artist.  I  am  grateful  to  Mexican 
officials  for  permission  to  collect  fishes  in  their  country. 


Table 

1. 

Distingu 

sh 

ng 

characters  of  the 

subgenera 

of  Heterandria. 

Character 

Heterandria 

Pseudoxiph  ophorus 

Gonopodium  (Fig.  2): 

Serrae  on  posterior  margin 
of  ray  4p 

Segments  beyond  distalmost 
serrae  of  ray  4p 

Tip  of  ray  5a 


Distal  part  of  ray  3 


7-9 

5  or  fewer 

Extends  beyond  tip  of  ray  4p 

Widely  separated  from  ray  4 


9-18 

5  or  more 

Extends  to  or  falls  short  of 
of  tip  of  ray  4p 

Closely  adjoining  ray  4 


Gonopodial  suspensorium 
(Fig.  3): 

Ligastyle 


Tip  of  gonapophysis  I 


Reduced  to  an  oval  remnant  below 
10th  vertebra 

Extends  ventrally  about  1/3  way 
from  vertebral  column  to  insertion 
of  pelvic  fin 


An  elongate  bone  lying  below 
1 1th  vertebra 

Extends  ventrally  more  than  1/3 
to  1/2  way  from  vertebral 
column  to  insertion  of  pelvic 
fin 


Reproductive  biology: 
Superfetation 

Egg  size  at  fertilization, 

2 
in  mm 

o 

Brood  interval 
Size 


Strongly  developed 
0.37-0.40 


1 


Averaging  5-6  days  (small  broods) 

Minute;  largest  mature  male  ca. 
14  mm  S.L. 


Absent  as  far  as  known 
2.08-2.56 

35-40  days  (large  broods) 

Moderate;  smallest  mature  male 
(jonesi)  ca.  22  mm  S.L. 


Dorsal  fin  of  female 


About  equal  in  size  to  anal,  its 
origin  behind  anal  origin,  over 
16th  or  17th  vertebra 


Much  larger  than  anal,  its  origin 
usually  farther  forward  (behind 
in  one  species),  over  12th  to 
15th  vertebra 


Dorsal  rays 
Vertebral  number 


6-8 

Sexually  dimorphic;  males  32-34, 
females  30-33  (Table  4) 


9-18 

No  sexual  dimorphism;  total 
variation,  30-34 


As  many  as  6  stages  of  developing  embryos  in  a  single  ovary  (Turner,  1937). 
^From  Scrimshaw,  1946;  based  on  H.  formosa  and  H.  bimaculata  only. 
^At  height  of  reproductive  season  (Turner,  1937). 
"^Over  16th  in  one  form  in  Alta  Verapaz,  Guatemala  (D.E.  Rosen,  pers.  comm.) 


237 


Counts  and  measurements  were  made  as  prescribed  by  Hubbs  and  Lagler  (1958: 
19-26).  Measurements  are  expressed  as  permillages  of  the  standard  length;  they  were 
taken  with  dial  calipers  reading  to  the  nearest  tenth  of  a  millimeter.  One  ratio  (length  of 
depressed  dorsal  into  predorsal  length)  was  stepped  off  with  a  pair  of  dividers  and  esti- 
mated to  the  nearest  tenth.  A  second  ratio  (base  of  dorsal  tin  into  predorsal  length)  was 
measured  with  calipers,  converted  into  permillages,  and  mathematically  calculated — a 
more  accurate  and  objective  means  of  obtaining  the  required  figures.  The  vertebral  count 
includes  the  hypural  plate  as  the  terminal  vertebra;  the  second  vertebra  is  the  first 
rib-bearing  one  in  all  cyprinodontoids. 

CHARACTERS  OF  THE  TWO  SUBGENERA 

When  only  two  species  were  assigned  to  Heterandria,  the  need  for  subgeneric  recogni- 
tion was  minimal.  Now  that  Pseudoxiphophorus  is  polytypic  (probably  containing  three 
or  more  species — Miller,  1966,  and  Rosen  and  Bailey,  MS),  it  is  helpful  to  employ  the 
subgenus  when  discussing  the  Middle  American  forms.  Consequently  I  have  drawn  up  a 
comparison  (Table  1)  which  provides  a  biological  as  well  as  a  structural  basis  for  recog- 
nizing two  subgenera  of  Heterandria:  some  may  feel  the  differences  are  sufficient  for 
generic  recognition.  Characters  that  indicate  a  close  relationship  between  Pseudoxipho- 
phorus and  Heterandria  involve  the  morphology  of  the  reproductive  system  and  the  breed- 
ing behavior  as  well  as  the  osteology  of  the  skull  (as  pointed  out  by  Rosen  and  Bailey, 
1963:  128-129).  Another  interesting  common  feature  discovered  in  the  present  study  is  the 
marked  sexual  dimorphism  in  the  length  of  the  snout:  H.  Jonesi—43  males,  78-96;  47 
females,  92-1 II.  H.  himacuUita—30  males,  82-100;  30  females,  94-111.  (Figures  are 
permillage  of  standard  length;  see  Table  3.)  H.  formosa — 10  males,  58-75;  10  females, 
76-89.  Another  interesting  aspect  of  the  comparison  is  the  sexual  dimorphism  in  vertebral 
number  in  subgenus  Heterandria  only:  males,  32  (8),  2)2)  (26),  34  (3);  females,  30  (1),  31 
(13),  32  (18),  22  (1) — Table  4.  The  tiny  egg  of  the  subgenus  Heterandria  is  correlated  with 
the  high  degree  of  dependence  on  the  mother  for  nourishment  by  the  developing  embryo. 
Such  virtual  elimination  of  yolk  is  paralleled  in  certain  species  of  Poeciliopsis  (e.g., 
Poeciliopsis  elongata  [Giinther]  and  P.  prolijica  Miller— see  Schultz  and  Thibault,  MS), 
in  which  superfetation  is  also  strongly  developed.  Presumably  superfetation  is  not 
developed  in  subgenus  Pseudoxiphophorus  (checked  only  in  H.  himaculata  and  H.  jonesi). 

THE  MEXICAN  SPECIES 

Heterandria  jonesi  (Gunther) 
(Figs.  1-3) 

Mollienisia  joncsii. —  Gunther,  1874:  371  (original  description,  based  on  females  only;  Lago  Alcuhuaca,  Mexico 
=  Lago  de  Aljojuca  —  see  Alvarez,  1950). 
Gamhusia  Joncsii. —  Regan.  1907:  260  (name;  comparisons).  Regan,  1906-08:  94,  97-98,  pi.  12,  fig.  8  (key; 

description;  synonymy;  female  syntype  figured;  distribution). 
Pscudoxiphophnnis  jonesii. —  Regan,  1913:  993  (synonymy;  description;  range). 

Pseudoxiphophorus  himuculatus  jom-sii. —   Hubbs,    1924:  17-18  (characters;   synonymy).   Alvarez,    1950: 

88-91  (redescription  of  topotypes;  correction  of  type  locality  to  Lago  de  Aljojuca,  15  km  NE  of  Ciudad 

Cerdan,  Puebla;  comparison  with  sample  from  Tepeaca,  Puebla,  in  Ri'o  Balsas  basin). 

Pseudoxiphophorus   bimaculatus   (misidentification). —   Woolman,    1894:  55-56    (description;    Rio    Blanco    at 

Orizaba).  Jordan  and  Evermann,  1896:  678  (description,  based  on  Orizaba  specimens).  Meek,  1904:  127 

(in  part;  Orizaba  records  only). 

Heterandria  himaculata  (misidentification). —  Rosen  and  Bailey,   1963:  131   (in  part;  references  io  jonesi 
and  pauciradiatus). 
Pseudoxiphophorus  reticuhitus  (misidentification). —  Jordan  and  Evermann,  1896:  678,  footnote  (description  of 

specimens  from  Ri'o  Blanco  at  Orizaba,  where  only  H.  Jonesi  occurs). 
Pseudoxiphophorus  pauciradiatus. —  Regan,   1904:  256  (original  description,  based  on  8  of  Woolman's  speci- 
mens from  Orizaba).  Regan,   1905:  362-363  (validity  of  species;  comparison  with  bimaculatus).   Regan. 
1907:  260  (listed  as  synonym  oi  Mollienisia  Jonesii). 

Diagnosis. — A  species  of  the  subgenus  Pseudoxiphophorus  (Table  1)  distinguished 
from  H.  himaculata  as  follows  (see  also  Table  5):  Terminal  segment  of  ray  4a  of  gono- 


r  // 


^^^, 


***>■ 


y  ■"*^- 


5^^^  i  1 1  j^r  1%  II  H\'  Vv*  An  r/« 


K'XCv^^:^^^f^. 


Xl<ii0im>xi 


*?"  T     -r       *  iiiiiiiS"'^ 


£12^ 


pjrSS- 


'J6c^:k- - 


Figure  1.  Top  io  botloiii:  Uctcrumhiu  joiicsi.  adult  male,  M  mm,  RaiKho  .Sierra  de  Agiia,  Ori/aba  Valley 
(L'MMZ  1838')4);  Hclcnuidria  jouvsi.  adult  temale,  43.5  mm.  t'rt)ni  same  collection  (topotypes  of  P.  painircnlialtis 
Regan);  Hcicnindria  himaciilata.  adult  male,  35,5  mm.  Nacimiento  de  Cosolapa  (UMMZ  183902);  Hclcrandria 
himuciilata.  adult  female,  50  mm,  from  same  collection. 


239 


podium  short  (not  longer  than  2-3  subdistal  segments  and  often  barely  exceeding  penuhi- 
mate  one),  sHghtly  reciir\ed,  not  reaching  tip  of  enclosing  membrane;  anterior  margin  of 
subdistal  segments  of  ray  4a  smooth;  ray  4p  forming  part  of  curved  tip  of  gonopod  (Fig. 
2).  Base  of  dorsal  tin  enters  predorsal  length  1.6  to  2.4  times  in  males,  and  2.0  to  2.9 
times  in  females;  depressed  dorsal  fin  enters  same  distance  1.2  to  1.6  (rarely  1.1)  times  in 
males,  and  1.4  to  2.0  times  in  females.  Origin  of  dorsal  fin  more  posterior  (Table  3). 
Basicaudal  spot  generally  smaller,  lower,  and  more  anterior,  lying  mostly  on  caudal 
peduncle. 

Type  locality. — This  species  was  described  from  Lago  de  Aljojuca,  a  crater  lake  or 
axalapazco  (Tamayo,  1964:  113)  in  the  endorheic  part  of  the  high  Puebla  Plateau  (Llanos 
de  El  Salado),  15  km  northeast  of  Ciudad  Cerdan,  Puebla,  and  west  of  the  great  volcano 
Pico  de  Orizaba  (5,750  m)  at  an  elevation  of  2,385  m  (Alvarez,  1950,  1972).  Apparently  it 
is  the  only  fish  native  to  this  lake,  although  three  other  similar  lakes  to  the  north  each 

Table  2.   Variation  in  number  of  dorsal  fin  rays  in  two  species  of  Heterandria  from  Mexico. 


Cat.  no.  and/or  authority  and  locality^ 


Number  of  dorsal  rays 


10      11 


12        13     14       15 


16 


17 


No.        Avg. 


H.  jonesi 


P  203,  Alvarez,  1950  (topotypes)^, 

Lago  de  Aljojuca,  Puebla 
P  184,  Tepeaca,  Puebla  (Balsas  basin) 
183986,  Acosac,  Puebla  (Balsas  basin) 
186675,  Tehuacan,  Puebla  (Papaloapan 

basin) 
183894,  Hubbs,  1924,  1926,  Orizaba 

Valley 
162143,  Ri'o  Atoyac,  Veracruz"^ 
183896,  Rfo  Atoyac,  Veracruz 
124304,  Rfo  Necaxa,  Puebla 

(Tecolutia  basin) 
193493,  42  km  WSW  Poza  Rica, 

Veracruz  (Cazones  basin) 
124330,  162141,  Palitla.S.L. 

Potosf  (Panuco  basin) 
183887,  Jaumave,  Tamaulipas 

(Tamesf  basin) 


70 

34 

5 

13 

17 

13 

22 


7   -    - 


38 

149 

14 

48 

43 

1 

96 

29 

— 

-   -   -    24   20 


12 


19   54 


8  24 


104  11.33 

20  11.75 

30  11.43 

30  12.20 

211  12.00 

96  11.42 

125  11.23 

47  12.55 

19  12.32 

76  12.79 

35  13.86 


H.  bimaculata 

Hubbs,  1924,  1926,  Jico  and  Jalapa, 

Veracruz  (Chachalacas  basin) 
USNM  31023,45489,  Mirador, 

Veracruz  (Chachalacas  basin) 
162144,  Rfo  Atoyac,  Veracruz'* 
181309,  Hubbs,  1924,  1926,  Cordoba, 

Veracruz  (subtopotypes) 
Regan,  1905,  Rfo  Tonto,  Veracruz 

(Papaloapan  basin) 
183902,  Cosolapa,  Oaxaca 

(Papaloapan  basin) 
124234,  4  km  E  El  Hule,  Veracruz 

(Papaloapan  basin) 
Regan,  1905,  in  Hubbs,  1924,  Sto. 

Domingo  Petapa,  Oaxaca  (Coatza- 

coalcos  basin) 
178533,  Rfo  Sarabia,  Oaxaca 

(Coatzacoalcos  basin) 


2  30  11  2 

6  23  4 

8  26  4 

1          7  30  13 

-  5  7 

3  12  32  13 
3       21  28  1 


1         3  1 

12      23         16 


45  13.29 

33  13.94 

38  13.89 

51  14.08 

15  14.87 

-        60  13.92 

53  13.51 

1  6  15.33 

51  15.08 


Catalog  numbers  are  those  of  UMMZ  unless  otherwise  stated. 
2|n  28    loaned  from  this  series  I  counted  1 1  (22),  12  (6).   Also  included  is  a  count  of  12  on  a  syntype,  BMNH 

1873.1.13.1  (illustrated  by  Regan,  1906-08:  PI.  12,  Fig.  8). 
-^Sympatric  with  bimaculata  (162144). 
^Sympatric  v\i\th  jonesi  (162143). 


240 


Figure  2.  Gonopodia  of:  A.  Heterundria  jonesi;  B.  Hetenuulrici  bimiiculata;  C.  Heterandria  fonnosa. 

contain  an  atherinid  of  the  genus  Poblana  {-Chirostoma:  see  Bolland  and  Barbour,  MS). 
The  tlsh  was  named  for  its  discoverer,  T.M.  Rymer  Jones. 

Variation. — The  gonopodium  (Figs.  2,  3)  provides  the  major  criterion  for  distinguish- 
ing//.  Jonesi  from  its  relatives.  It  is  therefore  important  to  know  how  much  it  varies.  The 
three  characteristics  given  in  the  diagnosis  include  the  variation  known  for  the  populations 
examined.  Other  features  follow.  Ray  3  terminates  near  the  proximal  end  of  segments  3 
to  6  of  ray  4a.  There  is  sharp  transition  between  the  several  elongate  proximal  segments 
of  ray  4a  and  the  short  subterminal  segments.  There  is  always  a  rather  abrupt  change  in 
height  and  size  between  the  most  proximal  serra-bearing  segment  of  ray  4p  and  the  next 
succeeding  segments  of  this  ray.  These  shorter  segments,  which  precede  the  last  one  of 
ray  4a,  vary  from  2  to  5.  Ray  4p  has  from  12  to  18  strong,  retrorse  serrae.  Ray  5a  ends 
about  2  to  4  segments  from  the  tip  of  ray  4a. 

Ten  measurements  were  made  on  males  and  females  of  four  populations  of  H.  Jonesi 
(Table  3)  representing:  (1)  the  type  locality  of  the  species  (Aljojuca),  at  2,385  m;  (2)  the 
type  locality  off.  pauciradiatus  (Orizaba),  at  1,240  m;  a  locality  (Palitla)  in  the  southern 
part  of  the  Rio  Panuco  basin,  at  about  120  m;  and  the  northernmost  known  population 
(Jaumave),  in  the  headwaters  of  the  Rio  Guayalejo,  at  about  330  m.  These  data  show 
that:  (1)  Aljojuca  and  Orizaba  specimens  have  the  shortest  dorsal-fm  base,  Jaumave  the 
longest,  with  Palitla  intermediate;  (2)  Jaumave  females  have  the  longest  anal  tin,  Orizaba 
and  Aljojuca  the  shortest,  with  Palitla  somewhat  intermediate;  (3)  the  caudal  tin  is  longest 
at  Jaumave,  generally  shortest  at  Orizaba  and  Aljojuca,  and  again  somewhat  intermediate 
at  Palitla,  although  the  measurements  do  not  overlap  those  at  Jaumave;  (4)  body  depth 
varies  greatly,  in  part  because  of  the  reproductive  condition  of  the  female,  as  at  Aljojuca 
(see  below);  (5)  head  length  shows  little  or  no  sexual  dimorphism  at  Aljojuca,  Orizaba,  or 
in  the  Rio  Atoyac  at  Atoyac  (10  males,  269-283,  ave.  273,  10  females,  260-280,  ave.  269), 
but  is  dimorphic  at  Palitla  and  Jaumave  (and  might  be  found  to  be  so  in  populations  at 
lower  elevations  between  Atoyac  and  Palitla);  (6)  snout  length  is  sharply  dimorphic 
between  the  sexes  at  all  four  localities,  as  are  predorsal  length  and  distance  between 
dorsal  origin  and  base  of  caudal  tin;  but  that  (7)  the  distance  from  anal  origin  to  caudal 
base  is  neither  sexually  dimorphic  nor  significantly  different  in  the  four  samples. 

The  number  of  dorsal-fin  rays  is  highest  at  Jaumave,  among  the  lowest  at  Aljojuca. 
and  intermediate  at  Palitla  (Table  2). 

Vertebral  number  is  rather  consistently  32,  varying  from  31  to  33,  in  Puebla  and 
adjoining  parts  of  Veracruz,  but  shows  a  decrease  toward  the  north  (southwest  of  Poza 


241 


Figure  3.   Gonopodial  suspensoria  ot:  A.  HelcnuiJna  Jonrsi;  B.  Hetcrandria  himciciilatu;  C.  Hi'tcnindriu  for- 
iiiosa.  (From  same  specimens  illustrated  in  Fig.  2.) 

Rica,  in  the  Rio  Cazones  basin),  especially  at  Jaumave,  where  the  mode  is  31   and  the 
range  30  to  32  (Table  4). 

Color  pattern  is  rather  consistent  for  the  populations  from  the  Puebla  Plateau 
(Alojojuca,  Tepeaca,  Acosac),  the  Orizaba  Valley,  and  the  Rio  Atoyac.  They  are  moder- 
ately to  strongly  barred,  with  3  to  1 1  rather  narrow  and  usually  short,  vertical  bars  con- 
fined to  the  midside  from  just  behind  the  base  of  the  pectoral  fin  to  just  before  the  basi- 
caudal  spot.  Generally,  the  larger  fish  have  the  most  bars.  These  vary  from  5  to  9  in  the 


242 


Table  3.   Proportional  measurements  of  Heterandria  jonesi  and  H.  bimaculata  (in  permillage  of  standard  length). 


Heterandria  jonesi 

/y.  bimaculata 

Aljojuca 

Orizaba 

Palitia 

Jaumave 

Cordoba 

Cosolapa 

P  203 

183894 

162141 

183887 

108614-^ 
181309 

183902 

Standard  length. 

Range  (mean)  No. 

1 

IVlales 

23.0-36.1 

23.6-36.1 

24.4-37.1 

22.8-27.1 

30.1-48.4 

25.5-51.9 

(27.2)  7 

(30.9)  15 

(30.1)  14 

(25.4)  7 

(38.9)  15 

(39.2)  15 

Females 

31.1-41.6 

29.9-55.5 

30.1-55.0 

26.0-47.1 

32.7-68.4 

43.1-76.2 

(38.2)  4 

(41.5)  15 

(38.9)  13 

(36.7)  15 

(50.0)  15 

(55.9)  15 

Body  depth 

Males 

267-290 

264-295 

264-297 

266-306 

253-290 

249-288 

(282) 

(280) 

(281) 

(293) 

(270) 

(267) 

Females 

255-283 

274-333 

271-296 

296-323 

240-281 

248-281 

(267) 

(299) 

(282) 

(310) 

(261) 

(265) 

Predorsal  length 

Males 

516-534 

500-538 

489-532 

498-522 

453-494 

452-486 

(525) 

(516) 

(519) 

(510) 

(470) 

(470) 

Females 

575-593 

568-597 

553-573 

561-589 

496-536 

516-546 

(587) 

(582) 

(563) 

(576) 

(520) 

(528) 

D.  Origin  to  C.  base 

Males 

490-518 

495-520 

492-531 

515-533 

540-583 

539-571 

(500) 

(507) 

(515) 

(523) 

(563) 

(554) 

Females 

421-430 

427-461 

456-483 

450-476 

477-514 

479-508 

(426) 

(445) 

(466) 

(462) 

(500) 

(493) 

A.  origin  to  C.  base 

Females 

437-453 

413-452 

431-448 

422-450 

431-470 

438-460 

(445) 

(437) 

(439) 

(437) 

(455) 

(449) 

Head  length 

Males 

270-287 

263-280 

263-296 

266-276 

266-287 

250-275 

(279) 

(269) 

(278) 

(270) 

(276) 

(261) 

Females 

270-286 

263-293 

276-308 

272-300 

252-306 

252-282 

(279) 

(277) 

(295) 

(284) 

(281) 

(267) 

Snout  length 

Males 

78-89 

78-87 

86-96 

79-82 

85-100 

82-96 

(83) 

(82) 

(92) 

(81) 

(92) 

(89) 

Females 

93-97 

92-105 

99-111 

92-103 

89-108 

94-107 

(95) 

(98) 

(105) 

(98) 

(102) 

(101) 

C.  peduncle  depth 

Males 

163-177 

158-176 

148-186 

166-193 

165-186 

157-190 

(170) 

(168) 

(169) 

(184) 

(178) 

(178) 

Females 

147-151 

144-161 

160-171 

172-183 

155-178 

153-168 

(150) 

(151) 

(164) 

(176) 

(162) 

(161) 

D.,  basal  length 

Males 

235-255 

225-256 

268-299 

302-325 

326-369 

329-358 

(244) 

(245) 

(284) 

(309) 

(348) 

(342) 

Females 

205-210 

202-226 

242-267 

264-290 

284-310 

273-315 

(207) 

(213) 

(253) 

(274) 

(300) 

(302) 

A.,  depressed  length 

Females 

190-203 

182-211 

203-225 

234-252 

193-239 

199-243 

(196) 

(196) 

(213) 

(244) 

(212) 

(220) 

C,  length  middle  rays 

Males 

238-258 

221-245 

237-275 

279-289 

227-260 

231-275 

(248) 

(232) 

(258) 

(283) 

(239) 

(248) 

Females 

220-227 

206  231 

225-256 

257-284 

203-239 

201-230 

(223) 

(215) 

(246) 

(271) 

(218) 

(216) 

material  examined  from  Aljojuca,  although  large  females  (such  as  the  syntype  figured  by 
Regan,  1906-08:  pi.  12,  fig.  8,  65  mm  S.L.,  examined  by  me)  may  show  no  trace  of  bars. 
At  Acosac,  adults  of  both  sexes  have  from  3  to  8  bars,  although  a  63-mm  female  lacks 


243 


them.  In  the  Orizaba  Valley,  45  fish  (30  males,  15  females)  have  from  4  to  10  bars. 
Vertical  bars  are  most  strongly  developed  in  the  two  samples  from  Rio  Atoyac,  wherein  all 
tish  (including  young  only  11  mm  long)  are  barred,  and  the  number  of  bars  varies  in  53 
adults  from  6  to  11,  usually  8  to  10.  In  the  Rio  Cazones  drainage  (42  km  WSW  of  Poza 
Rica),  the  bars  on  males  are  weakly  developed  (1-7  in  14)  and  are  apparently  lacking  in 
females  and  juveniles.  At  Palitla,  bars  are  also  weakly  developed  in  males  (from  none  to  7) 
and  none  is  evident  in  females  or  juveniles.  The  extreme  variation  is  attained  at  Jaumave, 
where  none  of  the  fish  collected  show  vertical  bars.  The  basicaudal  spot,  also  quite 
uniform  from  the  Puebla  Plateau  to  Rio  Atoyac,  is  rather  small,  generally  oval,  and  lies 
mostly  on  the  base  of  the  caudal  peduncle  not  far  above  the  body  axis  (Fig.  1).  In  the  Rio 
Cazones  collection,  the  spot  is  larger,  higher,  and  almost  as  much  of  it  lies  on  the  caudal 
fin  as  on  the  peduncle,  thus  more  closely  approaching  the  basicaudal  spot  typical  of  H. 
himaculata.  At  Palitla,  the  spot  is  more  like  that  at  Rio  Atoyac  except  that  it  lies  higher 
above  the  body  axis.  At  Jaumave,  the  basicaudal  spot  is  similar  to  that  at  Palitla  but 
tends  to  become  obsolete  in  large  females. 

Biology. — As  suggested  above,  body  depth  in  females  is  strongly  influenced  by 
pregnancy.  In  the  four  mature  females  measured  from  Aljojuca  (Table  3),  collected  21 
May  1949,  there  were  large  mature  eggs  but  no  embryos.  Permillage  values  for  body 
depth  are  from  255  to  283  (avg.  267),  whereas  in  43  females  from  the  three  other  localities 
(with  mean  standard  lengths  not  greatly  different  from  those  of  the  females  from  Aljojuca) 
these  values  are  from  271  to  333  (avg.  282,  299,  310).  Clearly  the  reproductive  season  is 
much  shorter  at  Aljojuca  (2,385  m)  than  it  is  at  the  lower  elevations.  For  example,  in  the 
10  largest  and  fattest  females,  collected  18  March  1968  from  Acosac  (UMMZ  183986,  ca. 
1,830  m),  one  had  advanced  embryos,  one  had  early  embryos,  and  eight  were  packed  with 
large  eggs — demonstrating  that  at  this  lower  elevation  the  reproductive  season  was  well 
under  way,  even  though  the  tlsh  were  taken  earlier  in  the  year.  At  still  lower  elevations 
production  probably  occurs  over  a  long  time  span  as  suggested  by  the  two  collections 
made  during  the  latter  half  of  December  from  Atoyac  (UMMZ  162143)  and  Palitla 
(UMMZ  162141),  each  of  which  contains  individuals  as  small  as  11  mm. 

It  is  a  general  observation  for  poeciliids  (but  not  for  all  viviparous  cyprinodontoids — 
e.g.,  goodeids,  Fitzsimons,  1972:  730)  that  males  have  determinate  growth  and  attain 
maturity  at  widely  different  sizes.  This  is  abundantly  supported  for  H.  jonesi  by  the  fol- 
lowing data  giving  the  frequencies  for  each  standard  length  measurement  (rounded  to 
nearest  whole  number)  followed  by  number  of  specimens  and  mean  value:  Orizaba  Valley 
(UMMZ  183894),  24  (7),  25  (16).  26  (9),  27  (9),  28  (11),  29  (8),  30  (2),  31  (4),  32  (3),  34 
(6),  35  (3),  36  (2),  in  80,  range  23.6-36.1  mm,  mean  28.1  mm;  Rio  Atoyac  at  Atoyac 
(UMMZ  183896).  27  (1),  29  (I),  30  (1),  31  (2),  32  (1),  35  (1),  36  (2),  37  (7),  38  (3),  39  (2), 
41  (1),  42  (1).  45  (1),  46  (1),  25,  37.5  (range  27.5-45.7  mm).  The  Orizaba  collection  is 
from  a  spring-fed,  roadside  ditch,  whereas  the  Atoyac  collection  is  from  a  large  river. 

At  only  one  locality  and  at  only  one  time  were  H.  jonesi  and  H.  himaculata  taken 
together  and  then  the  circumstances  were  unusual  although  the  data  on  number  of  dorsal 
rays  suggest  that  sympatry  may  be  normal  in  this  area.  The  collection  was  made  by  W. 
McLane  and  B.  Schultz  on  23  December  1940  in  the  Rio  Atoyac  (then  in  flood),  6.5  km 
north  of  Hacienda  Potrero  Viejo  (E  of  Cordoba  and  N  of  Hwy  150)  and  contains  96  speci- 
mens of //.  /o//('5/ (1 1-53  mm,  UMMZ  162143),  including  one  transforming  male,  and  38 
of//,  bimaculata  (10-40  mm,  UMMZ  162144),  including  one  mature  male.  The  dorsal 
rays  (Table  2)  show  an  overlap  only  at  13  rays;  the  8  specimens  of  H.  bimaculata  with 
that  number  are  discussed  under  Variation  in  the  account  of  that  species. 

Range. — The  northern  limit  of  this  species  (Fig.  4)  is  the  Rio  Guayalejo  of  south- 
eastern Tamaulipas,  the  major  tributary  of  the  Rio  Tamesi,  where  it  must  be  scarce. 
Darnell  (1962)  failed  to  take  Heterandria  in  the  66  collections  (totaling  over  1 1,000  fishes) 
made  during  1950-53  in  the  Tamesi  basin,  and  that  only  two  collections  are  known  from 
that  drainage:  2  specimens  from  near  the  bridge  just  west  of  Nuevo  Morelos,  Tamaulipas, 
Rosen  and  Gordon,  18  January  1957  (specimens  lost),  and  94  from  Jaumave,  discussed 
herein.  The  species  is  probably  widespread  in  suitable  habitats  throughout  the  Rio 
Panuco  basin  (up  to  elevations  near  2,000  m — e.g.,  Tula,  Hidalgo)  southward  to  the  Rio 


244 


Table  4.   Number  of  vertebrae  in  three  species  of  Heterandria  from  Mexico  and  Florida. 


Species,  cat.  no.   ,  locality 


Number  of  vertebrae 


30 


31 


32 


33 


34 


No. 


Avg. 


H.  jonesi  (Mexico) 


P  203,  L.  Aljojuca,  Puebia  (topotypes) 

P  184,  Tepeaca,  Puebia  (Balsas  basin) 

183986,  Acosac,  Puebia  (Balsas  basin) 

187718,  Orizaba  Valley,  Veracruz 

183896,  Rfo  Atoyac,  Atoyac,  Veracruz 

193493,  WSW  Poza  Rica,  Veracruz  (Cazones  basin) 

162141,  Palitla.S.L.  Potosf  (Panuco  basin) 

183887,  Jaumave,  Tamaulipas  (Tamesf  basin) 


H.  bimaculata  (Mexico) 


1 


USNM  31023,  45489,  Mirador,  Veracruz 

(Chachalacas  basin) 
108614,  181309,  Cordoba,  Veracruz  (subtopotypes) 
183902,  Cosolapa,  Oaxaca  (Papaloapan  basin) 
124234,  near  El  Hule,  Veracruz  (Papaloapan  basin) 
178533,  Ri'o  Sarabia  (Coatzacoalcos  basin) 

H.  formosa  (Florida) 

USNM  133265,  Crows  Bluff,  Deland,  males 

females 
USNM  210703,  Monroe  Station,  Tamiami  Trail, 

males 

females 


— 

12 

2 

2 

8 

— 

— 

13 

— 

— 

19 

6 

4 

24 

1 

11 

6 

1 

4 

22 

— 

24 

1 

— 

1 

13 

— 

- 

13 

17 

- 

12 

7 

8 

8 

— 

3 

20 

1 

2 

12 

7 

9 

— 

_ 

6 

14 

6 

9 

1 

14 

32.14 

10 

31.80 

13 

32.00 

25 

32.24 

29 

31.90 

18 

31.44 

26 

31.85 

30 

30.87 

14 

31.93 

30 

32.73 

20 

32.45 

16 

31.50 

24 

31.92 

15 

32.93 

17 

31.47 

22 

32.82 

16 

31.69 

I 


All  catalogue  numbers  are  UMMZ,  except  as  noted.  Localities  under  H.  jonesi  are  arranged  from  high  to  low 
elevations  (Aljojuca  to  Atoyac)  and  from  south  to  north  (Poza  Rica  to  Jaumave);  under  H.  bimaculata,  they 
are  arranged  from  north  to  south. 

Nautla.  The  species  reappears  in  the  Rio  Atoyac,  southwest  of  Veracruz  (City),  and  is  the 
only  Heterandria  of  the  Orizaba  Valley  and  the  streams,  irrigation  ditches,  and  crater 
lakes  (Aljojuca  only)  of  the  Puebia  Plateau,  where  it  occurs  as  high  as  2,385  m.  In  this 
elevated  region  it  lives  in  tributaries  of  the  Rio  Balsas  and  Rio  Papaloapan  as  well  as  in 
waters  without  exit  to  the  sea.  Its  distribution  south  of  the  latitude  of  Tehuacan  (1,649  m) 
is  unknown,  but  it  has  not  been  found  in  the  lowlands  of  the  Papaloapan  basin  or  any- 
where in  the  Rio  Coatzacoalcos  system,  the  next  major  drainage  to  the  south. 

Zoogeography. — The  occurrence  oi  Heterandria  jonesi  \n  a  high-elevation  crater  lake! 
(Aljojuca)  in  the  endorheic,  semidesert  basin  (area  ca.  8,000  km^)  occupied  by  the  Llanos 
de  Puebia  or  San  Juan  (Tamayo,  1964:   113,  Fig.  4)  calls  for  explanation.  The  remnant 
native  fish  fauna  (Chirostoma,    "Poblana",  and  Heterandria)  in  this  area  indicates  that 
the  interior  basin  they  now  occupy  was  connected  with  at  least  three  different  major 
drainages  during  late  Cenozoic  time.  Barbour  (1973)  has  presented  convincing  biological] 
evidence  to  tie  the  area  in  with  the  Rio  Lerma  basin  via  the  Valle  de  Mexico.  The  occur- 
rence of  the  cyprinid    "Aztecula  vittata  (Girard)"  in  Pleistocene  lake  deposits  and  anj 
adjacent  Recent  tributary  of  the  Balsas  basin  just  south  of  the  city  of  Puebia  (Miller  and 
Uyeno,  MS)  demonstrates  a  former  connection  between  the  Valle  de  Mexico,  Llanos  de, 
San  Juan,  and  the  Rio  Balsas  drainage.  Finally,  the  occurrence  of  the  otherwise  exclu- 
sively Atlantic  (in  Mexico)  subgenus  Pseudoxiphophoms  in  both  Lago  de  Aljojuca  and  in 
Rio  Balsas  tributaries  proclaims  a  former  connection  between  the  Llanos  de  San  Juan,  the] 
Atlantic  slope,  and  the  Balsas  system. 

Why  docs  not  Heterandria  jonesi  also  occur  in  the  three  other  fish-inhabited  crater  j 
lakes  and  in  Laguna  de  El  Carmen  (the  large  marsh-like,  shallow  lake  just  west  of  the 
crater  lakes)?  Probably  because  those  lakes  contain  species  of  atherinids  that  either  prey! 
on  the  newborn  young  of  the  poeciliid  or  outcompete  Heterandria  in  other  ways.  Lago  de 
Aljojuca  contains  only  Heterandria  which  elsewhere  in  this  plateau  region  occurs  only  in 


245 


spring-ted  ditches  and  ponds  tliat  contain  no  other  tlshes. 

Alvarez  (1972)  has  presented  the  following  hypothesis  to  explain  the  distribution  of 
the  tlshes  of  the  Llanos.  Prior  to  the  Pleistocene  this  plateau  region  contained  an  enor- 
mous shallow  lake  that  was  connected  to  the  Valle  de  Mexico  by  way  of  Apizaco  and 
Apani  (Barbour,  1973:  Fig.  5).  The  crater  lakes  in  question  were  formed  much  later  by 
volcanic  explosion,  and  their  cavities  were  tilled  with  water  because  they  all  lay  below  the 
highest  level  of  the  Plio-Pleistocene  lake.  Because  of  their  different  altitudes,  the  crater 
lakes  were  isolated  from  each  other  at  varying  times  as  the  level  of  the  hypothetical  lake 
fell  below  2.440  meters. 

Lago  de  Aljojuca  is  the  highest  of  these  tlsh-supporting  crater  lakes  and  some  partic- 
ular aspect  of  the  history  of  its  colonization,  as  yet  undetermined,  must  account  for  the 
fact  that  it  alone  contains  Hetcnnidria.  Very  likely  other  species  of  tlshes  became  extinct 
in  the  area  following  virtual  desiccation  of  the  original  lake. 

Specimens  examined  (All  in  Mexico). —  Hidalgo:  M74-48.  in  UMMZ,  ditch  near  Tula;  Puebla:  BMNH 
1873.1.13.1  (syntvpe),  Lago  de  Aljojuca;  P  184  (10),  Tepeaca;  P  203  (20  topotypes),  Lago  de  Aljojuca;  UMMZ 
124304  (47),  Ki'o  Necaxa;  UMMZ  183986  (50).  trib.  Rio  Balsas  at  Acosac;  UMMZ  186675  (57).  trib.  Rio  Papa- 
loapan,  Teluiacan;  UMMZ  193493  (19),  trib.  Rio  San  Marcos.  42  km  WSW  of  Poza  Rica.  San  Luis  Potosi: 
UMMZ  124330  (48).  162141  (45).  Palitla;  Tamaulipas:  UMMZ  183887  (94),  Jaumave.  Veracruz:  UMMZ  162143 
(96),  Rfo  Atovac,  6.5  km  N  Potrero  Viejo;  UMMZ  183894  (500).  Orizaba  Valley;  UMMZ  183896  (869).  Rio 
Atoyac  at  Atoyac;  UMMZ  187718  (47b),  Orizaba  Valley. 

Heterundria  bimaciilata  (Meckel) 
(Figs.  1-3) 

Xiphoplionis  himuculatus. —  Meckel.  1848:  297-299,  pi.  9,  figs.  1-2  (original  description;  a  clear  brook  ot  the 

Orizaba  Mountains,  Mexico). 

Poecilioiile.s  himaculutus. —  Steindachner,  1863:  176  (original  description;  Teapa,  Tabasco,  Mexico —  see 

Rosen  and  Bailey,  1963). 

Pseudoxiphophonts  himaculutus. —  Garnian.  1895:  81-82.  pi.  3.  fig.  6,  pi.  8,  fig.  9  (in  part;  synonymy; 
description).  Meek,  1902:  98  (brief  description;  maximum  adult  size;  notes  on  eggs,  embryos,  time  of 
birth).  Meek.  1904:  127-128  (in  part;  synonymy,  excluding  P.  pauciradiatus  Regan;  description;  range). 
Regan,  1904:  256  (comparison  with  P.  pauciradiatus;  P.  reticulatus  Troschel  in  synonymy).  Regan,  1913: 
993-994,  fig.  170C  (synonymy;  description;  gonopodium  figured).  Scrimshaw,  1946  (size  of  ova  and 
ovisac).  Rosen  and  Gordon,  1953:  26,  Fig.  32C  (gonopodium).  Rosen  and  Mendelson.  1960:  fig.  4M 
(hypothetical  correlation  between  sensory  canals  of  head  and  feeding  habits).  Rosen  and  Tucker.  1961: 
fig.  2  (secondary  sex  characters  and  sexual  behavior). 

Heierandria  himaculata. —  Rosen  and  Bailey.  1963:  131.  figs.  49B,  51 D,  55B  (in  part;  synonymy,  excluding 
references  to  jonesi  and  pauciradiatus;  range;  skeleton  and  gonopodial  suspensorium  of  male  figured). 
Miller.  1966:790  (range). 

Gambusia  himaculata. —  Regan.  1906-08:  98,  pi.  14,  fig.  4  (synonymy;  description;  range,  excluding 
Orizaba). 

Gamhusia  (Pseudoxiphophorus)  himaculata. —  Regan,  1907:  260  (listed  in  comparison  with  G.  annectens). 

Pseudoxi/tho/Jtiorus    himaculatus    himaculatus. —    Hubbs,     1924:  18    (synonymy;    distribution;    dorsal-ray 
counts). 
Pseudoxiphophorus  himaculatus  taeniatus. —  Regan.  1905:  363  (original  description;  San  Domingo  de  Guzman. 

Oaxaca.  Mexico;  this  locality,  now  called  Petapa.  is  on  a  tributary  of  the  Rio  Coatzacoalcos  SW  of  Mati'as 

Romero). 
Pseudoxiphophorus  himaculatus  peninsulae. —  Hubbs.  1936:  230-232,  pi.  8,  fig.  1  (original  description;  vicinity 

or  Progreso.  Yucatan.  Mexico). 
Pseudoxiphophorus  reticulatus. —  Troschel  in  von  Muller,  1865:  638-639  (original  description;  Mexico). 

Diagnosis. — A  species  of  the  subgenus  Pseudoxiphophorus  (Table  1)  distinguished 
from  H.  Jonesi  as  follows  (see  also  Table  5):  Terminal  segment  of  ray  4a  of  gonopodium 
greatly  elongate  (as  long  as  4-8  subdistal  segments),  its  tip  strongly  hooked  forward  (J- 
shaped),  reaching  tip  of  enclosing  membrance;  anterior  margin  of  4  to  6  subdistal  seg- 
ments of  ray  4a  with  keel-like  prominences;  ray  4p  not  entering  into  curved  tip  of  gonopod 
(Fig.  2).  Base  of  dorsal  tin  enters  predorsal  length  1.2  to  1.5  times  in  males,  and  1.6  to 
1.9  times  in  females.  Origin  of  dorsal  tin  more  anterior  (Table  3).  Basicaudal  spot  gen- 
erally larger,  higher,  and  more  posterior,  lying  mostly  on  caudal  tin. 

Type  Locality. — Confusion  has  resulted  from  the  common  misinterpretation  of 
Meckel's  type  locality  as  "Orizaba".  In  the  same  paper  in  which  H.  himaculata  is 
described,  Meckel  (1848)  also  described  Xiphophoms  helleri  and  Poeciliopsis  gracilis  (see 


246 


♦ 


Kigurc  4.  Distribution  ot  two  species  o\  Hctvnindriu  in  Mexico.  For  //.  hinuiculata  (which  ranges  eastward  and 
southward  into  Nicaragua)  only  the  records  from  the  Rfo  Coat/.acoalcos  basin  northward  are  included.  Stations 
are  Uom  UMMZ  records  and  Meek's  (1404:  128)  localities  (except  Sanborn,  not  found). 


Table  5.   Comparison  between  two  species  of  Heterandria  inhabiting  Mexico. 


247 


Character 


H.  jonesi' 


H.  bimaculata^ 


Gonopodium  (Fig.  2): 

Terminal  segment  of  ray  4a 


Anterior  margin  of  subdistal 
segments  of  ray  4a 

Ray  4p 


Ray  5a 


Gonopodial  suspensorium 
(Fig.  3) 


Dorsal  origin  to  caudal  base 

Predorsal  length 
Base  of  dorsal  fin 

Predorsal  length 

Depressed  dorsal-fin  length 

Basicaudal  spot  (Fig.  1) 


Cross-hatching  on  sides 


Short,  slightly  curved  forward,  not 
reaching  tip  of  enclosing  mem- 
brane; not  longer  than  2-3  sub- 
distal  segments,  often  only 
exceeding  penultimate  one 

Evenly  smooth  on  all 


Extending  far  beyond  tip  of  ray  3 
to  form  part  of  curved  tip  of 
gonopod 

Distal  part  curves  evenly  toward 
tip  of  gonopod;  ray  4p  is  only 
slightly  elevated  in  this  region 

Angle  of  gonactinosts  about  45° 
from  vertical.  Ligastyle  as  long 
as  or  longer  than  basal  stem  of 
gonapophysis  III 

Shorter.    In  permillage  of  S.L., 
males  490-533;  females  421-483 
(Table  3) 

Males  1.6-2.4;  females  2.0-2.9 


Greatly  elongate,  tip  strongly 
hooked  forward  (J-shaped), 
extending  to  end  of  enclosing 
membrane;  as  long  as  4-8  sub- 
distal  segments 

With  keel-like  prominences  on  4-6 
segments 

Extending  just  beyond  tip  of  ray  3, 
not  entering  into  curved  tip  of 
gonopod 

Distal  part  descends  abruptly  to  ray 
4p  which  is  strongly  elevated  in 
this  region 

Angle  of  gonactinosts  about  33° 
from  vertical.    Ligastyle  shorter 
than  basal  stem  of  gonapophysis 
III 

Longer.    In  permillage  of  S.L., 
males  539-575;  females  479-508 
(Table  3) 

Males  1.2-1.5;  females  1.6-1.9 


Males  1.1-1.6^;  females  1.4-2.0  Males  0.9-1.1 ;  females  1.2-1.4 


Generally  smaller,  its  position 
lower  and  more  anterior 
(weakest  in  females  from 
Jaumave) 

Well  developed  and  generally 
extending  ventrally  around 
caudal  peduncle 


Generally  larger,  its  position 
usually  higher  and  more 
posterior 


Well  developed  above  body  axis 
but  fading  ventrally;  none  on 
venter  of  caudal  peduncle 


Ratios  and  proportions  based  on  specimens  from  5  populations  (including  topotypes  of  M.  jonesil  and  P. 

pauciradiatus)  as  follows:     UMMZ  162141   (Palitia),  183887  (Jaumave),  183894  (Orizaba  Valley),  183896 

(Atoyac),  and  P  203  (Aljojuca). 
o 
Ratios  and  proportions  based  on  specimens  from  2  populations  (3  collections,  including  subtopotypes  of 

X.  bimaculatus)  as  follows;    UMMZ  108614  and  181309  (Cordoba),  and  183902  (Cosolapa). 
o 
This  ratio  was  determined  mathematically  by  dividing  the  measurement  of  the  base  of  the  dorsal  fin  into 

that  of  the  predorsal  length  for  each  fish. 

The  ratio  of  1.1  \r\  H.  jonesi  occurred  in  only  1  male  from  Jaumave;  otherwise  the  range  was  1 .2-1 .6. 

Rosen  and  Bailey,  1963:  131-133)  and  stated  that  all  three  species  live  together  "in  einem 
klaren  Bache  des  Gebirges  Orizaba".  Although  X.  helleri  inhabits  streams  of  the  Orizaba 
Valley,  neither  H.  hinuiculata  nor  F.  gracilis  live  there  (the  only  other  known  fish  is  H. 
Jonesi).  Therefore,  apparently  none  of  Meckel's  species  came  from  any  tributary  of  or 
stream  in  the  trough-like  Orizaba  Valley,  which  lies  at  an  elevation  of  about  1,240  m. 
Menzel  and  Darnell  (1973:  232),  in  discussing  the  type  locality  of  Poccilia  mexicana 
Steindachner,  also  said  to  be  from  Orizaba,  concluded  that  the  types  came  from  a  much 
lower  elevation  in  either  the  Ri'o  Jamapa  or  Ri'o  Blanco  drainages.  Most  likely  X.  helleri 
and  H.  himaculaiu  (if  not  P.  gracilis)  came  from  the  vicinity  of  Cordoba  at  an  elevation  of 
about  870  m.  Woolman  (1894:  65)  described  the  rapids  and  barrier  falls  in  the  Rio 
Blanco,  the  drainage  system  of  the  Orizaba  Valley,  which  prevent  the  ascent  of  fishes  into 


248 


this  valley  from  lower  elevations  to  the  east  of  Orizaba. 

Variation. — The  salient  characters  distinctive  of  the  gonopodium  of  this  species  have 
been  given  in  the  Diagnosis  and  also  appear  in  Table  5.  Additional  traits  follow.  Ray  3 
extends  to  the  base  of  the  penultimate  segment  or  to  that  of  the  terminal  one  (the  J- 
shaped  hook)  of  ray  4a.  Distal  to  the  elevated  tlange  or  keel  of  ray  3  are  from  6  to  9  small 
segments.  Ray  4a  bears  from  2  to  4  squarish  segments  between  the  terminal  one  and  the 
last  keeled  segment,  and  from  6  to  9  segments  distal  to  the  terminal  serra  on  ray  4p. 
There  are  6  to  12  segments  distal  to  the  last  serra  of  ray  4p,  and  this  ray  bears  10  to  14 
strong,  retrorse  serrae.  Ray  5a  ends  several  segments  of  ray  4a  short  of  the  base  of  the 
J-shaped  hook. 

Ten  measurements  (Table  3)  were  made  on  30  males  and  30  females  sampled  from 
two  places  in  Veracruz  and  Oaxaca — Cordoba  (approximate  type  locality  of  the  species) 
and  Cosolapa.  approximately  50  airline  km  SE  of  Cordoba.  These  data  show  very  close 
agreement  except  in  head  length,  which  is  longer  at  Cosolapa.  Sexual  dimorphism  is 
strongly  marked  in  dorsal  origin  (as  shown  by  predorsal  length,  and  dorsal  origin  to 
caudal  base)  and  basal  length  of  dorsal  tin;  it  is  less  striking  in  caudal  peduncle  depth 
and  length  of  middle  caudal  rays. 

The  number  of  dorsal  tin  rays,  lowest  in  the  highlands  (Jico-Jalapa)  near  the  northern 
limit  of  the  range,  appears  to  show  an  increase  southward  and  toward  lower  elevations. 
The  extreme  range  for  this  species  is  from  11  to  18  (11  in  one  specimen  from  Honduras, 
UMMZ  173305,  and  18  in  three  from  Belize,  formerly  British  Honduras — specimens 
taken  by  David  W.  Greentleld  at  Sta.  G70-139). 

Vertebral  number  shows  modes  of  32  or  2)2>  in  samples  from  tlve  populations  in 
Mexico  (Table  4). 

Color  pattern  is  generally  more  consistent  in  bimaculata  than  in  Joiiesi.  Cross- 
hatching  is  well  developed  on  the  upper  and  mid-sides  but  begins  to  pale  ventrally  and 
fades  out  entirely  over  the  ventral  surface  of  the  caudal  peduncle.  Vertical  bars  are  rare 
although  Hubbs  (1936:  231)  stated  that  H.  b.  pcninsulac  from  Yucatan  has  2  to  4  such 
bars,  "like  narrow  parr  marks",  behind  the  shoulder  spot.  Among  35  young  to  juvenile 
individuals  from  the  Rio  Atoyac  (UMMZ  162144)  are  8  with  2  to  5  faint  bars  anteriorly;  7 
of  these  have  13  dorsal  rays,  the  number  that  overlaps  that  of  H.  Jonesi  at  this  same 
locality.  However,  in  no  other  features  do  these  specimens  resemble  /o/a'.s/;  measurements 
of  the  basal  length  of  the  dorsal  tin  and  predorsal  length  yielded  calculated  ratios  of  less 
than  1.0  in  all  eight  specimens  (see  Table  5).  Bars  thus  appear  to  be  only  rarely  developed 
in  H.  bimaculata.  The  basicudal  spot  is  somewhat  variable  in  size  and  position.  Typically 
it  is  large,  roundish,  more  or  less  equal  to  the  diameter  of  the  eye,  set  higher  than  in 
/V>//<'.s/ and  mostly  on  the  caudal  tin.  However,  in  a  collection  from  the  Rio  Coatzacoalcos 
basin  (Ri'o  Sarabia,  Oaxaca,  UMMZ  178533),  the  spot  lies  almost  entirely  on  the  base  of 
the  caudal  tin,  generally  only  slightly  above  the  body  axis,  and  varies  from  round  to  tri- 
angular with  the  apex  of  the  triangle  (often  drawn  out)  directed  posteriorly.  This  popula- 
tion (corresponding  to  H.  b.  taeniata  of  Regan)  also  shows  a  well-developed  midlateral 
stripe  that  is  disrupted  in  young  specimens. 

Biology. — Meek  (1902:  98)  reported  that  this  species  probably  gives  birth  "near  the 
first  to  the  middle  of  June"  (at  Jalapa,  Veracruz,  1,427  m).  Possibly  this  is  true  but  if  so, 
successful  fertilization  and  development  take  place  later  in  the  year  than  it  does  in  H. 
Jonesi.  A  collection  (UMMZ  108614)  made  on  22  March  from  Cordoba  (872  m)  contains 
individuals  as  small  as  14  mm  standard  length,  indicating  that  brood  production  had 
been  under  way  for  some  time.  As  already  indicated  (see  Biology,  H.  Jonesi),  both  species 
had  produced  young  in  the  Ri'o  Atoyac  (about  600  m)  by  late  December. 

Mature  males  of//,  biiuaculata.  like  those  of//,  jonesi,  vary  greatly  in.  size:  25  (1), 
27  (1),  29  (2),  30  (1),  31  (1),  il  (3),  i:>  (6),  34  (2).  35  (3),  37  (1),  38^2),  39  (2),  40  (2),  43 
(1),  44  (3),  45  (5),  46  (1),  47  (2),  48  (3),  49  (3),  50  (1),  51  (1),  52  (1),  48,  avg.  39.4  mm 
(UMMZ  183902,  Cosolapa).  The  largest  of  20  immature  males  in  this  collection  was  51, 
the  smallest  40,  and  16  were  41  or  more  mm  long  (all  these  males  had  the  gonopodium 
elongated  but  undifferentiated  at  the  tip).  The  74  mature  females  in  this  collection, 
varying  from  43  to  76  mm  long,  averaged  58.1  mm. 


249 


Synipatry  between  H.  hiniaciilatu  and  //.  joncsi  has  already  been  discussed  (see 
Biology,  H.  ioiwsi). 

Raii}^c. —  Ihe  precise  northern  limit  of//,  himaculata  on  the  Atlantic  coastal  plain  is 
uncertain,  but  it  evidently  does  not  extend  to  the  Rio  Nautla  basin,  which  as  far  as  known 
contains  only  //.  Joncsi.  The  northernmost  collection  known  to  me  is  from  the  Rio 
Misantla  (M74-6,  in  UMMZ;  Fig.  4),  an  independent  tributary  to  the  Gulf  of  Mexico 
lying  just  southeast  of  the  Rio  Nautla,  Veracruz;  this  stream  is  north  of  Jalapa,  which  lies 
near  the  northernmost  inland  limit  of  H.  himaculata.  In  Mexico  the  species  occurs  at 
elevations  from  near  sea  level  to  at  least  1,430  m  (Jalapa). 

Spccinn-ns  examined  (All  in  Mexico). —  Oaxaca:  UMMZ  17853.1  (52),  Rio  Sarabia  on  Trans-Isthmian  Hwy; 
UMMZ  18.1902  (757),  Cosolapa.  Veracruz:  U.SNM  ,1102.1  (5)  and  45489  (27),  Mirador;  UMMZ  108614  (207), 
Ki'o  Chico,  Cordoba;  UMMZ  124234  (h9).  4  km  E  ot  Papaloapan  (  =  E1  Hiilc);  UMMZ  l(i2144  (.18),  Rio  Atovac. 
(1.5  km  N  ot  Potrero  Viejo;  UMMZ  181.109  (35),  Cordoba:  UMMZ  184512  (94),  32.5  km  N  ol  Jose  Cardel.' 

PHYLOGENY 

In  considering  the  relationships  of  phyletic  lines  within  the  subgenus 
P.sciulo.xiphop/ionis.  it  is  clear  from  Table  5  and  Figures  2  and  3  that  //.  Joncsi  is  less 
specialized  than  //.  himaculata.  The  gonopodium,  especially,  is  of  simpler  construction  in 
joncsi.  Although  several  species  of  this  subgenus  are  yet  to  be  described  from  Guatemala. 
I  have  examined  all  of  them  and  conclude  that  none  is  more  primitive  than  //.  Joncsi.  In 
body  form  and  proportions,  position  and  size  of  the  dorsal  tin,  head  shape,  length  of 
mandible,  and  detailed  architecture  of  the  gonopodium,  each  of  the  Guatemalan  species 
shows  some  features  that  indicate  a  less  generalized  condition  than  is  found  in  H.  j'oncsi. 
The  species  represented  by  UMMZ  193893  (Alta  Verapaz,  Guatemala)  is  perhaps  as 
close  to  Joncsi  as  is  any  of  the  Guatemalan  species,  but  it  shows  certain  modifications 
about  the  distal  end  of  the  gonopodium  (e.g.,  thickening  of  ray  3.  increased  number  of 
small  segments  in  ray  4a)  that,  to  me.  mark  it  as  more  specialized. 


LITERATURE  CITED 

Alvarez.  Jose 

1950.   Contribucion  al  conocimiento  de  los  peces  de  la  region  de  los  Llanos,  estado  de  Puebla  (Mexico).  An. 
Esc.  Nac.  Cien.  Biol.,  b  (1-4):  81-107.  tigs.  1-3. 

1972.  Algiinos  ejemplos  de  especiacion  en  peces  niexicanos.  Acta  Polit..  Mex.,  13(b0):  81-89.  tigs.  1-6. 
Barbour.  CD. 

1973.  A  biogeographical  history  of  Chirosioiiui  (Pisces:  Atherinidae):  a  species  tlock  from  the  Mexican 
Plateau.  Copeia.  1973(3):' 533-556,  tigs.  1-18. 

Boiland.  R.F.,  and  CD.  Barbour 

MS.      Svstematic  status  ot  the  genus  Fohhuui  Dc  Buen  (Pisces:  Atherinidae). 
Darnell,  R. 

1962.   Fishes  of  the  Rio  Tamesi  and  related  coastal  lagoons  in  east-central  Mexico.  Publ.  Inst.  Mar.  Sci.,  8: 
299-365,  tigs.  1-2. 
Fitzsinions.  J.M. 

1972.   A  revision  of  two  genera  of  goodeid  tlshes  (Cyprinodontitormes,  Osteichthyes)   troni   the  Mexican 
Plateau.  Copeia,  1972(4):  728-756,  tigs.   1-10. 
Carman,  S. 

1895.    Ihc  cyprinodonts.  Mem.  Mus.  Comp.  Zool..  Harvard  Coll..   19:   1-P9,  pis.   1-12. 
Gunther,  A. 

1874.    Descriptions  of  new  species  of  tlshes  in  the  British   Museum.    Ann.    Mag.    Nat.    Hist.,   ser.   4,    14: 
.168-3^1. 
Meckel,  J. 

1848.   Fine  ncue  Gattung  von  Poecilien  mit  rochenartigen  Amklammcruiigs-Organc.   .Sitzber.    K.   Akad. 
Wiss.  Wien,  Math^Natwiss.  CI..  1:  289-.l()3.  pis.  8-9. 
Hulibs,  CL. 

1924.    .Studies  of  the  tlshes  of  the  order  Cyprinodontes.    1-I\'.   Misc.   Publ.    Mus.  Zool.   Univ.   Mich..    13: 

1-31,  pis.   1-4. 
1926.   Studies  of  the  tlshes  of  the  order  Cyprinodt)ntcs.  \  1.  Material  for  a  revision  of  the  American  genera 

and  species.  Misc.  Publ.  Mus.  Zool.  Univ.  Mich..  16:   1-86.  pis.   1-4. 
1936.   Fishes  of  ihc  'i  ucatan  Peninsula.  Cam.  Inst.  Wash.  Publ.  45":  15"-28",  tig.  1.  pK.  1-15. 


250 


Hubbs.  C.L.  and  K.F.  Lagler 

1958.   Fishes  of  the  Great  Lakes  region.  Bull.  Cranbrook  Inst.  Sci.,  26:  i-xiii,   1-213,  figs.   1-251.  col.  pis. 
1-44. 
Jordan.  D.S.,  and  B.W.  Evermann 

18%.   The  fishes  ot  North  and  Middle  America.  Bull.  U.S.  Nat.  Mus.,  47(1):  I-LX.  1-1240. 
Meek.  S.E.  ' 

1902.   A  contribution  to  the  ichthyology  of  Mexico.  Field  Col.  Mus.,  Publ.  65,  Zool.  Scr.  3(6):  63-128,  pis.     j 

14-31. 
1904.  The  fresh-water  fishes  of  Mexico  north  of  the  Isthmus  of  Tehuantepec.  Field  Col.  Mus.,  Publ.  93, 
Zool.  Ser.  5:  i-lxiii,  1-252,  figs.  1-72,  1  map,  pis.  1-17. 
Menzel,  B.W.,  and  R.M.  Darnell 

1973.   Systematks  of  Poecilia  mexicana  (Pisces:  Poeciliidae)  in  northern  Mexico.  Copeia,  1973(2):  225-237. 
figs.  1-3. 
Miller.  R.R. 

1966.  Geographical  distribution  of  Central  American  freshwater  fishes.  Copeia,  1966(4):  773-802,  figs.  1-5. 
Regan.  C.T. 

1904.  Descriptions  of  new  or  little-known  fishes  from  Mexico  and  British  Honduras.  Ann.  Mag.  Nat.  Hist., 
ser.  7.  13:  255-259. 

1905.  A  collection  of  fishes  made  by  Dr.  H.  Gadow  in  southern  Mexico.  Ann.  Mag.  Nat.  Hist.,  ser.  7,  16: 
361-363. 

1906-08.  Pisces.  In:  Biologia  Centrali-Americana,  8:  1-203.  7  figs.,  pis.  1-26. 

1907.   Descriptions  of  six  new  freshwater  fishes  from  Mexico  and  Central  America.  Ann.  Mag.  Nat.  Hist., 

Ser.  7.  19:  258-260. 
1913.   A  revision  of  the  cyprinodont  fishes  of  the  subfamily  Poeciliinae.  Proc.  Zool.  Soc.  London,   1913: 

977-1018,  figs.  168-173,  pis.  99-101. 
Rosen.  D.E..  and  R.M.  Bailey 

1963.  The  poeciiiid  fishes  (Cyprinodontiformes).  their  structure,  zoogeography,  and  systematics.  Bull.  Am. 
Mus.  Nat.  Hist..  126:  1-176.  figs.  1-61,  maps  1-19,  pis.  1-2. 

Rosen,  D.E.,  and  M.  Gordon 

1953.   Functional  anatomy  and  evolution  of  male  genitalia  in  poeciiiid  fishes.  Zoologica,  38(1):  1-47,  figs. 
1-47.  pis.  1-4. 
Rosen,  D.E..  and  J.R.  Mendelson 

1960.  The  sensory  canals  of  the  head  in  poeciiiid  fishes  (Cyprinodontiformes).  with  reference  to  dentitional 
types.  Copeia,  1960(3):  203-210,  figs.  1-4. 

Rosen,  D.E.,  and  A.  Tucker 

1961.  Evolution  of  secondary  sexual  characters  and  sexual  behavior  patterns  in  a  family  of  viviparous  fishes 
(Cyprinodontiformes:  Poeciliidae).  Copeia,  1961(2):  201-212,  figs.  1-3. 

Schultz.  R.J. ."and  R.  Thibault 

MS.     Reproductive  strategies  among  viviparous  fishes.  Amer.  Nat. 
Scrimshaw,  N.S. 

1946.   Egg  size  in  poeciiiid  fishes.  Copeia.  1946(1):  20-23. 
Steindachner.  F. 

1863.   Beitrage  zur  Kcnntniss  der  Scianoiden  Brasiiiens  und  der  Cyprinodonten  Mejicos.  Sitzber.  K.  Akad. 
Wiss.,  Wien,  Math.-Natwiss..  48(1):  162-185.  pis.  1-4. 
Tamayo.  J.L. 

1964.  The  hydrography  of  Middle  America,  pp.  84-121.  figs.  1-6.  //;:  Handbook  of  Middle  American  Indi- 
ans, vol.  1.  R.  Wauchope  and  R.C.  West,  eds.  Univ.  Texas  Press,  Austin. 

Turner.  C.L. 

1937.   Reproductive  cycles  and  superfetation  in  poeciiiid  fishes.  Biol.  Bull.,  72(2):  145-164. 
von  Muller.  J.W. 

1865.   Reisen  in  den  Vereinigten  Staaten,  Canada  und  Mexico.  Vol.  3.  Leipsig.  pp.  i-xii,  1-643. 
Wooiman,  A.J. 

1894.   Report  on  a  collection  of  fishes  from  the  rivers  of  central  and  northern  Mexico.   Bull.   U.S.   Fish 
Comm..  14(1895):  55-66.  pi.  2. 


4 


I 


Museum  of  Zoology,  The  University  of  Michigan.  Ann  Arbor.  Michigan  48104  U.S.A. 


SAN  (^(sn 


LITHOSTRATIGRAPHIC  VARIATIONS 

IN  THE  POWAY  GROUP 

NEAR  SAN  DIEGO,  CALIFORNIA 


GARY  L.  PETERSON  AND  MICHAEL  P.  KENNEDY 


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TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 


VOL.  17,  NO.  18 


6  DECEMBER  1974 


;■  i^'*  "*"- 


LITHOSTRATIGRAPHIC  VARIATIONS  IN  THE  POWAY  GROUP 
NEAR  SAN  DIEGO,  CALIFORNIA 

GARY  L.  PETERSON  AND  MICHAEL  P.  KENNEDY 


ABSTRACT. — The  Eocene  Poway  Group  consists  of  two  mutually  intertongued  bodies  of  rock,  or 
iithosomes.  One  consists  predominantly  of  coarse  conglomerate  of  fluvial  origin  and  is  located  principally 
in  the  eastern  San  Diego  area.  The  other  consists  of  sandstone  and  siltstone  and  lies  predominantly  west 
of  the  conglomerate  lithosome.  Tongues  of  the  conglomerate  lithosome  extend  toward  the  west  and 
include  the  Stadium  Conglomerate  and  the  herein  named  Pomerado  Conglomerate  (Upper  Eocene). 
Tongues  of  the  sandstone  lithosome  extend  toward  the  east  and  pinch  out  within  the  conglomerate. 
Lower  sandstone  tongues  constitute  the  Mission  Valley  Formation,  and  an  upper  tongue  is  herein  named 
the  Miramar  Sandstone  Member  of  the  Pomerado  Conglomerate  (Upper  Eocene).  The  sandstone 
lithosome  is  partly  of  nearshore-marinc  and  partly  of  nonmarine  origin,  whereas  the  conglomerate  is  of 
tluviatile  origin.  The  conglomerate  was  deposited  as  a  delta  at  the  site  where  a  large  Eocene  river 
emerged  onto  a  low -lying  coastal  plain. 

One  of  the  most  distinctive  and  widespread  stratal  units  in  the  San  Diego  area  is 
referred  to  in  the  older  literature  as  the  "Poway  Conglomerate"  (Ellis  and  Lee.  1919; 
Hanna.  1926;  Bellemin  and  Merriam,  1958;  and  many  others).  More  recently,  Kennedy 
and  Moore  (1971)  recognized  that  this  Eocene  stratal  unit  is  composed  not  only  of 
conglomerate,  but  also  contains  at  least  one  widespread  mappable  sandstone  unit.  They 
revised  the  nomenclature  accordingly  and  raised  the  rank  of  the  "Poway  Conglomerate" 
to  the  Poway  Group.  Within  the  Poway  Group,  Kennedy  and  Moore  (1971)  recognized  a 
lower  rock  unit  designated  the  Stadium  Conglomerate,  a  middle  unit  dominated  by 
fine-grained  sandstone  designated  the  Mission  Valley  Formation,  and  a  third  unnamed 
conglomerate,  herein  designated  the  Pomerado  Conglomerate. 

A  geologic  map  of  part  of  San  Diego,  and  adjacent  areas,  now  completed  at  a  scale 
of  1:24,000  (Kennedy  and  Peterson,  1974).  shows  the  distribution  of  these  formations  and 
their  relationships  to  one  another.  A  small  portion  of  that  map  is  included  here  as  Fig- 
ure 1.  The  purpose  of  this  paper  is  to  name,  briefly  describe,  and  establish  type  sections 
for  new  rock  units,  as  well  as  to  describe  the  vertical  and  lateral  variations  in  lithologic 
character  within  the  Poway  Group. 

The  lateral  distribution  of  rock  types  within  the  Poway  Group  is  best  illustrated  and 
explained  by  utilizing  the  lithosome  concept  of  Wheeler  and  Mallory  (1956).  rather  than 
more  traditional  lithostratigraphic  units  (formations  and  members).  Briefly,  a  lithosome 
is  a  rock  body  of  uniform  character  that  intertongues  with  one  or  more  other  rock  bodies 
of  uniform  but  differing  character.  The  individual  tongues  of  the  lithosome  are  referred  to 
by  Wheeler  and  Mallory  as  lithostromes,  and  are  here  what  we  have  mapped  as 
formations  and  members  (Fig.  1). 

The  lithostratigraphic  variation  in  the  Poway  Group  is  most  pronounced  in  an 
east-west  direction  (Fig.  2).  The  Poway  Group  is  subdivided  into  two  mutually 
intertongued  Iithosomes,  each  representing  a  different  depositional  environment.  One 
lithosome  is  dominated  by  conglomerate  and  is  herein  informally  referred  to  as  the 
conglomerate  lithosome.  The  other  is  dominated  by  soft,  friable  sandstone  and  is 
hereafter  referred  to  as  the  sandstone  lithosome. 

The  basically  simple  lithosomal  relations  that  we  illustrate  in  Figure  2  are  in  part 
interpretational,  since  the  Poway  Group  is  only  partially  presened.  The  principal 
complicating  factor  is  the  presence  of  the  Lindavista  Terrace,  a  Pleistocene  wave-cut 
platform  capped  by  a  thin  veneer  of  reddish-brown  sandstone  and  conglomerate  (the 
Lindavista  Formation).  Because  of  this  later  erosional  episode,  the  Poway  Group  has  a 
large  notch  removed,  and  the  variations  within  this  missing  portion  of  the  group  are  no 
longer  evident.  This  erosional  notch  affects  the  western  part  of  the  Poway  Group,  or  that 

SAN  DIEGO  SOC.  NAT.  HIST.,  TRANS.  17(18):  251-258,  6  DECEMBER  1974. 


252 


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Figure  1.  Geologic  map  of  the  Miramar  Reservoir  area  showing  locations  of  type  sections  of  the  Late  Eocene 
Pomerado  Conglomerate  and  Miramar  Sandstone  Member. 

portion  which  we  interpret  to  be  dominated  by  the  sandstone  lithosome.  We  have 
reconstructed  within  this  missing  portion  what  we  consider  to  be  the  most  reasonable 
lithostratigraphic  variation  to  account  for  the  relations  observed  and  mapped  within  the 
preserved  portion  of  the  section. 

The  lithosomes,  together  with  their  lithostromes,  which  are  mappable  rock  units  of 
member  and  formational  rank,  are  briefly  described  below.  A  discussion  of  the  regional 
genetic  significance  follows  in  the  last  section. 

CONGLOMERATE  LITHOSOME 


The  conglomerate  lithosome  is  composed  of  one  of  the  most  distinctive  rock  types  in 
the  San  Diego  area.  The  clasts  range  in  size  from  pebbles  to  small  boulders,  and  locally 
are  up  to  nearly  a  meter  in  diameter.  Clasts  over  30  cm  in  diameter  are  rare.  The  clasts 
are  subrounded  to  rounded  and  are  set  in  a  medium-  to  coarse-grained  sandstone  matrix. 


I 


253 


In  general  field  appearance  the  conglomerate  is  so  distinctive  that  with  only  some  minor 
exceptions  involving  reworking,  it  can  be  readily  distinguished  from  the  older  and 
younger  conglomerates  of  the  area  (Peterson,  1970).  Thin  beds  and  lenses  of 
cross-stratit'ied  sandstone  lithologically  similar  to  the  conglomerate  matrix  occur 
throughout  the  section  and  increase  in  persistence  toward  the  west.  The  conglomerate  is 
chiefly  nonmarine,  and  we  interpret  the  interstratified  sandstone  to  be  partly  marginal 
marine  but  predominantly  of  fluvial  origin. 

The  conglomerate  is  characterized  by  the  Poway  suite  of  clasts  (Bellemin  and 
Merriam,  1958;  DeLisle  et  al.,  1965;  Woodford  et  al.,  1968;  Peterson,  1971),  an  exotic 
assemblage  consisting  predominantly  of  rhyolitic  to  dacitic  volcanic  and  volcaniclastic 
rocks  with  a  smaller  but  significant  proportion  of  quartzite.  This  assemblage  of  clasts  first 
appears  in  the  stratigraphic  succession  at  San  Diego  in  the  Eocene  (Peterson  and 
Nordstrom,  1970),  is  extensively  reworked  into  post-Eocene  rock  units,  and  in  most  places 
is  the  dominant  clast  suite  found  in  the  modern  stream  and  beach  gravels.  The  Poway 
clasts  are  exceedingly  durable,  having  travelled  an  exceptionally  long  distance  to  the  site 
of  deposition.  Their  probable  area  of  origin  seems  to  be  on  the  Sonoran  side  of  the  Gulf 
of  California  (Bellemin  and  Merriam,  1958;  DeLisle  et  al.,  1965;  Woodford  et  al.,  1968; 
Minch,  1970,  1972). 

The  character  of  the  conglomerate  lithosome  is  surprisingly  uniform  throughout  the 
San  Diego  area.  It  varies  little  either  geographically  or  stratigraphically.  Thus  the 
Stadium  Conglomerate  is  lithologically  nearly  identical  to  the  Pomerado  Conglomerate. 
These  conglomerates  can  be  differentiated  only  because  they  are  separated  by  the  Mission 
Valley  Formation. 

Stadium  Conglomerate. — The  lower  conglomerate  lithostrome  was  designated  the 
Stadium  Conglomerate,  with  a  type  section  near  San  Diego  Stadium  in  Mission  Valley. 
This  rock  unit  is  very  widespread  and  has  been  recognized  throughout  the  San  Diego 
region  (Kennedy  and  Moore,  1971;  Peterson,  1971).  Its  thickness  is  highly  variable  and 
ranges  from  a  few  meters  to  perhaps  75  m.  In  general  it  is  thickest  and  most  typically 
developed  in  the  central  San  Diego  area  and  becomes  progressively  thinner  to  the  north 
and  west.  The  Stadium  Conglomerate  is  overlain  by  the  finer-grained  Mission  Valley 
Formation.  The  contact  between  the  two  units  is  gradational,  and  locally  the  two  units  are 
intertongued. 

Pomerado  Conglomerate. — The  upper  conglomerate  lithostrome  is  here  named  the 
Pomerado  Conglomerate.  A  well-exposed  section,  here  designated  the  type  section,  is 
located  along  the  roadcuts  of  Pomerado  Road  and  Sycamore  Canyon  access  road  between 
San  Diego  and  Poway  (location  P  in  Figure  1). 

At  the  type  section,  the  Pomerado  Conglomerate  gradationally  overlies  the  Mission 
Valley  Formation,  a  unit  consisting  of  gray  to  light  brown  sandstone  containing  a  small 
amount  of  whitish  caliche,  scattered  pebbles,  and  small  cobbles  of  rhyolitic  rock.  The 
basal  Pomerado  contact  is  placed  at  the  base  of  a  7  m  massive  conglomerate  of  typical 
Poway  type.  Overlying  the  conglomerate  is  a  7  m  thick  medium-grained,  soft,  friable 
sandstone  resembling  the  underlying  Mission  Valley  Formation  but  interpreted  here  as  a 
lens  of  sandstone  within  the  Pomerado  Conglomerate. 

Overlying  the  sandstone  lens  is  a  14  m  thick  massive  cobble  conglomerate  with  the 
typical  Poway  suite  of  clasts,  many  of  which  are  fractured  in  situ.  This  conglomerate 
grades  upward  into  a  sandstone  containing  small  scattered  pebbles  and  a  thin  bed  of 
pebble  conglomerate.  Thickness  of  the  sandstone  is  7  m.  It  is  overlain  by  5  m  of  cobble 
conglomerate  with  some  clasts  up  to  30  cm  diameter.  Overlying  this  conglomerate  is  a 
2  m  thick  sandstone  lens  which  is  in  turn  overlain  by  a  1  m  massive  cobble  conglomerate 
bed.  Overlying  the  conglomerate  is  a  3  m  bed  of  soft,  friable,  medium-grained  sandstone 
containing  some  Poway-type  pebbles.  The  highest  exposed  unit  of  the  Pomerado 
Conglomerate  is  a  10  m  thick  bed  of  cobble  to  boulder  conglomerate.  Some  of  the 
boulders  are  up  to  30  cm  in  diamter  and  many  are  fractured  in  situ.  Some  thin,  mostly 
discontinuous  beds  and  lenses  of  sandstone  are  present  in  this  otherwise  massive 
conglomerate  bed. 

The  top  of  the  type  section  of  the  Pomerado  Conglomerate  ends  at  the  crest  of  the 


254 


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Figure  2.  Generalized  diagram  of  relationship  between  sandstone  lithosome  (Mission  Valley  Formation  and 
Miramar  Sandstone  Member  of  Pomerado  Conglomerate)  and  conglomerate  lithosome  (Stadium  Conglomerate 
and  Pomerado  Conglomerate). 

I 

hill.  In  this  area  no  further  units  overlie  the  Pomerado.  Farther  to  the  south,  the  Pliocene 

San  Diego  Formation  unconformably  overlies  the  Pomerado  Conglomerate. 

The  Pomerado  Conglomerate  is  gradational  with  the  underlying  Mission  Valley 
Formation  throughout  the  map  area  (Fig.  1)  and  much  of  the  San  Diego  region.  To  the 
east,  however,  the  Mission  Valley  Formation  becomes  thinner  and  finally  pinches  out 
between  the  Pomerado  Conglomerate  and  Stadium  Conglomerate. 

No  fossils  have  been  found  in  the  Pomerado  Conglomerate;  it  is  here  considered  to  be 
Late  Eocene  in  age,  as  it  is  gradational  with  and  overlies  the  Mission  Valley  Formation, 
which  is  Late  Eocene  (Kennedy,  1973). 

The  Pomerado  Conglomerate  is  not  nearly  as  widespread  as  the  Stadium 
Conglomerate,  but  the  difference  in  distribution  is  at  least  in  part  a  matter  of 
preservation.  The  Stadium  Conglomerate  extends  beneath  the  Lindavista  wave-cut 
platform  and  is  present  in  much  of  the  western  part  of  the  San  Diego  area.  The  Pomerado 
Conglomerate,  for  the  most  part,  terminates  at  the  old  sea  cliff  associated  with  the 
Lindavista  Terrace  (Fig.  2). 

Within  part  of  the  Pomerado  Conglomerate  is  a  moderately  widespread  sandstone 
and  siltstone  unit  here  designated  the  Miramar  Sandstone  Member.  We  interpret  this  as 
a  tongue  of  the  sandstone  lithosome  (Fig.  2)  described  below. 

SANDSTONE  LITHOSOME 

The  sandstone  lithosome  is  composed  primarily  of  soft  white,  gray,  yellow,  and  light 
brown  friable  sandstone  with  interbedded  soft  gray  to  green-gray  siltstone.  In  general, 
this  rock  type  crops  out  much  more  poorly  and  is  not  nearly  as  obvious  as  the 
conglomerate.  In  addition,  the  sandstone  and  siltstone  rock  units  tend  to  be  covered  with 
slopewash  derived  from  the  overlying  conglomerate.  However,  exposures  are  present  at 
road  cuts  and  because  of  a  distinctive  topographic  expression  their  distribution  can  be 
mapped  with  some  degree  of  certainty. 

In  addition  to  the  dominant  lithology,  the  sandstone  lithosome  contains  random  thin 
beds  and  lenses  of  conglomerate.  The  conglomerate  is  similar  in  all  respects  to  that  of  the 
conglomerate  lithosome  and  is  considered  to  represent  minor  tongues  of  that  unit.  The 
conglomerate  beds  and  lenses  locally  constitute  up  to  about  20  per  cent  of  the  sandstone 
lithosome,  which  is  also  equivalent  to  the  maximum  per  cent  of  sandstone  beds  and  lenses 
locally  present  in  the  conglomerate  lithosome. 

A  marginal  marine  and  nonmarine  environment  of  deposition  for  the  sandstone 
lithosome  is  based  on  the  presence  of  fossil  mammals,  tlsh,  lagoonal  oysters,  and 
nearshore-marine  moUusks  (Kennedy,  1973). 

Mission  Valley  Formation. — The  Mission  Valley  Formation  is  a  rock  unit  named  by 
Kennedy  and  Moore  (1971),  with  a  type  section  along  the  south  side  of  Mission  Valley 
near  State  Highway  163  (old  U.S.  395).  From  that  locality  the  Mission  Valley  Formation 
extends  over  a  wide  area  that  includes  parts  of  the  La  Mesa,  La  Jolla,  Del  Mar,  and 


255 


Poway  IVi  minute  quadrange.  The  Mission  Valley  Formation  is  thickest  in  its 
westernmost  exposures  near  Mission  Valley.  To  the  east,  within  the  upper  Mission  Gorge 
area,  the  lower  part  of  the  formation  intertongues  with  the  upper  part  of  the  Stadium 
Conglomerate  (Fig.  2).  An  upper  tongue  of  the  Mission  Valley  Formation  can  be  seen  in 
upper  Murphy  Canyon  but  pinches  out  rapidly  toward  the  eastern  boundary  of  the  Poway 
and  La  Mesa  quadrangles.  Beyond  this  line,  the  Pomerado  and  Stadium  Conglomerates 
are  in  contact.  A  Late  Eocene  age  has  been  assigned  to  the  Mission  Valley  Formation 
based  on  the  presence  of  Tejon  mollusks  in  the  nearshore-marine  part  of  the  section  and 
Uinta  C  mammals  in  the  nonmarine  part  (Kennedy,  1973). 

Miramar  Sandstone  Member  of  Pomerado  Conglomerate. — The  uppermost  tongue 
of  the  sandstone  lithosome  is  found  entirely  within  the  Pomerado  Conglomerate  in  the 
general  vicinity  of  Miramar  Reservoir.  Lithologically,  it  is  identical  to  the  Mission  Valley 
Formation,  and  its  outcropping  characteristics  are  similar.  However,  because  this  unit  is 
wholly  within  the  Pomerado  Conglomerate,  and  because  it  is  nowhere  in  contact  with  the 
Mission  Valley  Formation,  we  are  here  designating  it  the  Miramar  Sandstone  Member  of 
the  Pomerado  Conglomerate.  We  interpret  the  Miramar  Member  as  well  as  the  Mission 
Valley  Formation  to  be  tongues  of  the  sandstone  lithosome,  but  this  interpretation 
depends  on  evidence  within  that  part  of  the  Poway  Group  that  has  been  erosionally 
removed  by  the  cutting  of  the  Lindavista  Terrace  (Fig.  2). 

The  type  section  for  the  Miramar  Sandstone  Member  is  here  designated  to  be  along 
the  fire  road  extending  along  the  ridge  at  the  northern  margin  of  Miramar  Reservoir  (see 
Fig.  1).  At  the  type  section,  the  Pomerado  overlies  the  Mission  Valley  Formation  which 
consists  of  soft,  friable,  red  to  brown  weathering  sandstone.  The  lower  part  of  the 
Pomerado  consists  of  17  m  of  pebble  to  cobble  conglomerate  composed  of  the  Poway  suite 
of  clasts. 

Overlying  the  lower  conglomerate  is  20  m  of  soft  medium-  to  coarse-grained,  gray  to 
gray-brown,  red-brown  weathering  sandstone  here  designated  the  Miramar  Sandstone 
Member.  The  sandstone  is  best  exposed  in  gullies  at  the  edge  of  the  fire  road.  Locally  it  is 
pebbly,  containing  the  Poway  suite  of  clasts,  and  locally  it  is  fractured  with  the  fractures 
filled  with  caliche.  In  all  respects,  the  Miramar  Member  strongly  resembles  the  Mission 
Valley  Formation. 

The  Miramar  Member  is  overlain  by  32  m  of  massive  cobble  conglomerate,  the  upper 
unit  of  the  Pomerado  Conglomerate  in  this  section.  The  conglomerate  is  dominated  by 
cobbles  and  small  boulders  with  some  of  the  clasts  ranging  up  to  30  cm  in  diameter. 

From  the  type  section,  the  Miramar  Member  can  be  traced  around  the  hills 
surrounding  Miramar  Reservoir.  Where  traced  to  the  east  it  pinches  out  within  the 
Pomerado  Conglomerate  (Figs.  1,  2). 

No  fossils  were  found  within  the  Miramar  Member,  although  the  general  lithologic 
similarity  to  the  Mission  Valley  Formation  suggests  a  similar  environment  of  deposition. 
The  fact  that  it  is  within  the  Pomerado  Conglomerate,  which  is  gradational  with  the 
underlying  Mission  Valley  Formation,  suggests  an  age  of  late  Eocene. 

REGIONAL  IMPLICATIONS 

In  the  Late  Eocene  in  the  San  Diego  area  a  large  river  valley  emerged  along  the 
Pacific  Coast.  This  ancient  valley,  called  the  Ballena  Channel  by  Minch  (1970,  1972),  is 
traceable  eastward  almost  to  the  Elsinore  Fault.  It  enters  the  San  Diego  area  in  the 
vicinity  of  San  Vicente  Reservoir.  There  are  narrowly  distributed  channel  deposit,  (the 
"Ballena  Gravel")  rapidly  fans  out  and  grades  westwardly  into  the  Late  Eocene  stratal 
units  of  the  San  Diego  area  (Kennedy  and  Moore,  1971;  Peterson,  1971). 

Apparently  the  Ballena  Channel  carried  much  of  the  coarse  sediment,  especially  that 
of  the  coarse  conglomerate  characterized  by  "Poway"  clasts,  which  is  now  so  abundant  in 
the  Eocene  succession  of  the  San  Diego  area.  The  Ballena  River  entered  the  San  Diego 
embayment  from  the  east,  apparently  dropping  much  of  its  coarsest  load  in  the  area  of 
lowering  gradient  as  it  entered  the  coastal  plain  of  San  Diego.  The  conglomerate 
lithosome  represents  predominantly  fiuvial  deposits.   These  conglomerate  beds   grade 


256 


laterally  and  intertongue  westward  (also  to  an  extend  northward  and  southward)  with  the 
sandstone  lithosome.  # 

The  sandstone  lithosome  is  at  least  partially  marine.  Thus  a  significant  portion  of  the 
tine  detritus  may  have  been  derived  via  longshore  transport  or  roughly  at  right  angles  to 
the  westward  paleoslope  indicated  by  the  Ballena  Channel.  In  addition  ,  much  of  the  fine 
detritus  appears  to  be  supplied  by  local  minor  drainage  channels  (Peterson,  1971). 

Generally  continuous  submergence  was  necessary  to  preserve  the  rocks  of  the  Poway 
Group.  The  large  scale  intertonguing  of  the  two  distinctive  lithosomes  can  be  interpreted 
in  several  ways.  First,  the  eastward  extensions  of  the  partially  marine  sandstone-siltstone 
lithosome  indicate  eastward  transgression  of  the  strand  line.  The  maximum  transgression 
would  be  represented  by  the  Mission  Valley  Formation.  The  regressive  phases  would  be 
represented  by  the  tluvial  conglomerate  lithosome,  with  the  maximum  regressions 
represented  by  the  Stadium  and  Pomerado  Conglomerates.  The  transgressive-regressive 
fluctuations  could  have  been  caused  by  interacting  eustatic  sea-level  changes  and  local 
tectonic  subsidence.  Such  fluctuations  tit  well  with  earlier  transgressive-regressive 
episodes  represented  within  the  underlying  La  Jolla  Group  (Kennedy  and  Moore.  1971). 

A  second  possible  interpretation  is  that  the  size  and  position  of  the  conglomerate 
lithosome  may  be  due  to  variations  in  the  amount  of  coarse  fluvial  detritus  being 
transported  into  the  San  Diego  area.  That  is,  in  times  of  voluminous  supply,  such  as 
during  the  deposition  of  the  Stadium  and  Pomerado  conglomerates,  a  conglomeratic 
fan-delta  could  have  built  westward  at  the  expense  of  the  marine  environment.  During 
times  of  less  fluvial  sediment  supply  during  the  subsidence,  the  marine  environment 
might  again  have  encroached  eastward. 

A  third  possibility  is  that  the  intertonguing  lithosomes  indicate  a  combination  both 
of  changes  in  rate  of  submergence  and  of  changes  in  rate  of  sediment  influx.  We  consider 
this  possiblity  the  most  plausible. 


LITERATURE  CITED 


Bellemin,  G.J.,  and  R.H.  Merriam 

1958.   Petrology  and  origin  of  the  Poway  Conglomerate.  San  Diego  County.  California.  Geol.  Soc.  Amer. 
Bull.  69:  199-220. 
DeLisle.  M.,  J.R.  Morgan,  J.  Heldenbrand,  and  G.  Gastil 

1965.   Lead-alpha  ages  and  possible  sources  of  metavolcanic  rock  clasts  in  the  Poway  Conglomerate,  south- 
west California.  Geol.  Soc.  Amer.  Bull.  76:  1069-1074. 
Ellis.  A.J.,  and  C.H.  Lee 

1919.  Geology  and  ground  waters  of  the  western  part  of  San  Diego  County,  California.  U.S.  Geol.  Survey 
Water-Supply  Paper  446:  1-321. 
Hanna.  M.A. 

1926.  Geology  of  the  La  Jolla  quadrangle,  California.  Univ.  Cal.  Publ.  Geol.  Sci.  16:  247-398. 
Kennedy.  M.P. 

1973.  Stratigraphy  of  the  San  Diego  embayment,  California.  Ph.D.  Thesis.  Univ.  California  (Riverside). 
Kennedy.  M.P.,  and  G.W.  Moore 

1971.  Stratigraphic  relations  of  Upper  Cretaceous  and  Eocene  formations.  San  Diego  coastal  area,  Cali- 
fornia. Amer.  Assoc.  Petroleum  Geol.  Bull.  55:  709-722. 

Kennedy.  M.P.,  and  G.L.  Peterson 

1974.  Geology  of  the  La  Mesa,  Poway.  and  southwest  quarter  of  the  Escondido  quadrangles,  eastern  San 
Diego  metropolitan  area,  California.  California  Div.  Mines  and  Geol.  Bull.  200B. 

Minch,  J. A. 

1970.  Early  Tertiary  paleogeography  of  a  portion  of  the  northern  Peninsular  Ranges.  In  Pacific  slope 
geology  of  northern  Baja  California  and  adjacent  Alta  California.  Amer.  Assoc.  Petroleum  Geol. 
(Pacific  Sec.)  Fall  Field  Trip  Guidebook:  4-9. 

1972.  The  Late  Mesozoic-Early  Tertiary  framework  of  continental  sedimentation,  northern  Peninsular 
Ranges,  Baja  California,  Mexico.  Ph.D.  Thesis.  Univ.  California  (Riverside). 

Peterson,  G.L. 

1970.  Distinctions  between  Cretaceous  and  Eocene  conglomerates  in  the  San  Diego  area,  southwestern 
California.  In  Pacific  slope  geology  of  northern  Baja  California  and  adjacent  Alta  California.  Amer. 
Assoc.  Petroleum  Geol.  (Pacific  Sec.)  Fall  Field  Trip  Guidebook:  90-98. 

1971.  Stratigraphy  of  the  Poway  area,  southwestern  California.  San  Diego  Soc.  Nat.  Hist.  Trans.  16: 
225-236. 


257 


Peterson.  G.L..  and  C.E.  Nordstrom 

1970.  Sub-La  Jolla  unconformity  in  vicinity  of  San  Diego,  California.  Amer.  Assoc.  Petroleum  Geol.  Bull 
54:  265-274. 
Wheeler,  H.E.,  and  V.S.  Mallory 

1956.   Factors  in  lithostratigraphy.  Amer.  Assoc.  Petroleum  Geo!.  Bull.  40:  2711-2723. 
Woodford,  A.O..  E.E.  Welday,  and  R.H.  Merriam 

1968.   Siliceous  tuff  clasts  in  the  upper  Paleogene  of  southern  California.  Geol.   Soc.   Amer.    Bull.    79: 
1461-1486. 


Department  of  Geological  Sciences.  San  Diego  State  University,  San  Diego.  California. 
California  Division  of  Mines  and  Geology.  Geological  Research  Division,  Scripps 
Institution  of  Oceanography.  La  Jolla,  California. 


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MU«.  COM,.,  roou 

LIBRARY     ^ 

MB  1  9 1973 


THE  AUTECOLOGY  OF 
XANTUSIA  HENSHAWI  HENSHAWI 
(SAURIA:  XANTUSIIDAE) 


HARVARO 


JULIAN  C.  LEE 


TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 


VOL.  17,  NO.  19 


22  APRIL  1975 


THE  AUTECOLOGY  OF 
XANTUSIA  HENSHA  WI  HENSHA  WI 
(SAURIA:  XANTUSIIDAE) 

JULIAN  C.  LEE 


ABSTRACT. — Xantusia  henshawi  henshawi  is  a  secretive,  crevice-dwelling  lizard  confined  to  southern 
California  and  adjacent  Baja  California,  Mexico.  Field  studies  during  portions  of  four  years  at  two  sites  in 
western  San  Diego  Co.  and  studies  of  musuem  specimens  revealed  that  the  distribution  of  this  saxicolous 
species  is  correlated  with  the  presence  of  granitic  rocks.  The  availability  of  suitable  rock  crevices  is  an 
important  factor  limiting  both  population  size  and  geographic  distribution.  Brush  fires  may  generate 
additional  crevices  by  accelerating  the  exfoliation  of  boulders. 

This  species  is  slow-growing,  late-maturing,  long-lived,  and  has  the  lowest  reproductive  potential  yet 
reported  for  any  lizard.  Males  probably  first  breed  at  about  2.5  years;  females  breed  first  at  3.5  years,  and 
produce  one  brood  (mean  size  1.46)  per  season.  Head  and  tail  length  exhibit  allometric  growth  relative  to 
snout-vent  length;  linear  growth  is  determinant.  The  frequency  of  caudal  autotomy  is  high  among  adults 
and  probably  results  from  intra-specific  fighting  rather  than  predation.  The  mortality  rate  from  all  causes  is 
low. 

Xantusia  h.  /zens/iauj  occupies  a  thermally  buffered  microhabitat,  and  in  summer  maintains  body  tempera- 
tures within  approximately  the  same  limits  day  and  night.  This  species  is  eurythermic,  showing  no  distinct 
temperature  preference.  Body  temperature  is  highly  dependent  on  substrate  temperature. 

The  life  history  of  this  lizard  is  similar  to  that  of  Xantusia  vigilis. 

Of  the  approximately  3,000  living  species  of  lizards,  the  life  histories  of  perhaps  no 
more  than  50  have  been  studied  thoroughly  (Fitch,  1970).  Because  of  its  limited  and 
highly  disjunct  distribution,  the  family  Xantusiidae  is  of  particular  interest.  Yet  the  only 
general  life  history  for  any  xantusiid,  is  Miller's  (1951)  study  of  Xantusia  vigilis.  The 
present  study,  designed  to  fill  partially  this  gap  in  our  knowledge  of  lizard  ecologies, 
presents  an  analysis  of  the  life  history  of  Xantusia  henshawi. 

Possibly  because  of  its  limited  distribution,  secretive  habits,  and  specialized 
microhabitat,  little  information  has  been  published  on  this  species  since  its  discovery  in 
1893.  Authors  who  have  discussed  aspects  of  the  biology  of  X  henshawi —  often  in  casual 
or  anecdotal  fashion  —  include:  Atsatt  (1925),  Brattstrom  (1951,  1952,  1965),  Grinnell 
and  Camp  (1917),  Klauber  (1926,  1931.  1939),  Lee  (1974),  Mautz  and  Case  (1974),  Scott 
(1971),  Shaw  (1949),  and  Stephens  (1921). 

Description. — Xantusia  henshawi  is  a  small  lizard.  Adult  males  average  56  mm  SVL 
and  weigh  about  2.9  g.  Adult  non-gravid  females  average  62  mm  and  3.3  g.  These  lizards 
are  dorso-ventrally  compressed  (Fig.  1),  a  feature  associated  with  their  crevice  dwelling 
habits.  The  limbs  are  well  developed,  pentadactyl,  and  the  digits  bear  small,  strongly 
recurved  claws.  The  head  is  covered  with  enlarged,  symmetrical  shields;  the  dorsal  and 
lateral  body  surfaces  and  throat  are  covered  with  granular  scales;  the  venter  bears  14 
longitudinal  rows  of  enlarged  rectangular  plates;  and  the  tail  is  covered  with  whorls  of 
smooth,  rectangular  scales.  Femoral  pores  are  present  in  both  sexes.  As  in  all  xantusiids, 
eyelids  are  lacking,  the  eye  being  covered  by  a  transparent  spectacle.  The  pupil  is 
vertically  elliptical.  This  species  exhibits  a  daily  rhythmic  color  change  (Atsatt,  1939): 
during  the  day,  the  animal  is  dark  gray  or  black  with  a  fine  yellowish  reticulum;  at  night 
the  yellowish  network  expands,  and  the  animal  becomes  grayish  with  dark  spots. 
I  Distribution. — Prior  to  1970,  X.   henshawi  was  known  only  from  "Rocky  areas  on 

both  sides  of  the  mountains  from  northern  Riverside  Co.,  California,  to  the  San  Pedro 
Martir  Mountains,  Lower  California"  (Stejneger  and  Barbour,  1943;  see  also  maps  in 
Stebbins,  1954,  1966).  Webb  (1970)  described  a  disjunct  population  which  he  named 
Xantusia  henshawi  bolsonae,  from  a  single  locality  in  eastern  Durango,  Mexico,  1280  km 


SAN  niFr.o  <;nr  nat  hi<;t    tran^   17  (iq^  ?';q-->7s  7^  aprii  iq7^ 


260 


Figure  1.   Adult  male  Xantusia  henshawi  henshawi.  56  mm  snout-vent  length. 

southeast  of  the  nearest  population  of  the  nominate  race.  Figure  2  presents  the 
distribution  of  X.  h.  henshawi  based  upon  published  localities,  museum  records,  and 
data  from  this  study. 

Webb  (1970)  summarized  variation  in  X.  h.  henshawi  based  upon  an  examination  of 
108  specimens  from  throughout  its  geographic  range.  Comparison  with  Webb's  data 
indicated  that  the  Mt.  Woodson  and  Lee  Valley  populations  sampled  in  this  study  are 
referable  to  X  h.  henshawi  (Table  1). 

Habitat. — X.  henshawi  is  saxicolous  and  is  rarely  found  far  from  crevices,  especially 
those  formed  by  the  exfoliation  of  granitic  boulders  (Fig.  3).  Such  rocks  are  a  requirement 
for  this  species,  and  their  absence  is  probably  a  factor  limiting  both  distribution  and 
population  density. 

Chaparral  is  the  principal  plant  community  in  the  range  of  the  nominate  race, 
although  ecotonal  chaparral-coastal  sage  scrub  situations  are  inhabited  in  western  San 
Diego  Co.,  as  is  the  chaparral-creosote  bush  scrub  ecotone  in  eastern  San  Diego  Co., 
southwestern  Imperial  Co.,  and  portions  of  Riverside  Co. 

Summers  are  generally  hot  and  dry  throughout  the  range  of  this  subspecies. 
However,  the  rock-crevice  microhabitat  occupied  hy  Xantusia  henshawi  henshawi  protects 


116 

— r- 


Riverside 


•     San  Diego 


1  Imperial 

1 


I' 


Baja   California 


-3f 


Figure  2.  Distribution  of  Xantusia  henshawi  henshawi.  Open  arrow  indicates  study  site  1,  solid  arrow  indicates 
study  site  2. 


261 


TABLE  1 .  Comparison  of  taxonomic  characters  in  20  adult  Xantusia  henshawi henshawi  from  Mount  Wood- 
son, San  Diego  County,  California,  20  adults  from  Lee  Valley,  San  Diego  County,  and  108  specimens  of  the 
subspecies  from  throughout  its  range  (Webb,  1970).  The  first  figure  is  the  mean;  the  figures  in  parentheses, 
the  range. 

Character  Webb,  1970  Mount  Woodson  Lee  Valley 

Number  of  infralabials  5.1  (4-7)  5.8  (5-6)  6.4  (5-7) 

Number  of  supralabials  6.2(5-8)  6.7(6-9)  7.0(6-8) 

Head  width/body  length  0.18(0.16-0.28)        0.17(0.16-0.18)        0.17(0.16-0.19) 

Number  of  dorsal  granules  at  midbody  62.8(56-71)  61.3(55-66)  63.4(57-69) 

Number  of  transverse  rows  of  ventral  scales  32.7(30-36)  34.1(32-36)  33.0(31-34) 

Number  of  enlarged  scales  on  gular  fold  10.3(7-14)  11.2(9-13)  11.8(7-14) 

Number  of  temporal  scales  5.6  (4-8)  5.6  (5-7)  5.6  (4-8) 

Number  of  femoral  pores  10.7(7-16)  11.0(10-13)  11.3(10-14) 

them  from  temperature  extremes.  This  thermal  buffering  effect  is  illustrated  in  Figure  4. 
The  meager  rainfall  (often  less  than  36  cm  per  year)  is  mostly  restricted  to  the  cooler  fall, 
winter,  and  spring  months  (Fig.  5). 

The  possible  ecological  relationships  of  vertebrates  known  to  coexist  with  X.    h. 
henshawi  are  indicated  in  Table  2. 


MATERIALS  AND  METHODS 

Field  work  at  two  sites  was  conducted  intermittently  in  1968,  1970,  and  1971,  and 
systematically  from  April  through  October,  1972.  Site  1,  in  Lee  Valley,  San  Diego  Co., 
California  (Fig.  2)  is  described  elsewhere  (Lee,  1974).  Site  2  is  located  4.8  km  north  and 
4.8  km  east  of  Poway,  San  Diego  Co.,  California  (Fig.  2).  The  site  includes  portions  of 
Warren  Canyon  and  the  south  and  southwest  slopes  of  Mount  Woodson.  Elevation  ranges 
from  430  m  at  the  bottom  of  Warren  Canyon  to  890  m  at  the  top  of  Mount  Woodson.  The 
steep  flanks  of  Warren  Canyon  and  tlie  slopes  of  Mount  Woodson  are  strewn  with 
exfoliating  granitic  boulders  and  covered  with  chaparral  (Fig.  6).  Portions  of  the  area 
were  burned  in  1967. 

In  this  investigation  735  living  lizards  and  171  preserved  specimens  from  the  San 
Diego  Society  of  Natural  History  (SDSNH)  were  examined.  Locality  data  were  obtained 
from  collections  in  the  San  Diego  Society  of  Natural  History,  Museum  of  Vertebrate 
Zoology,  Los  Angeles  County  Museum  of  Natural  History,  and  the  California  Academy  of 
Sciences. 

Population  structure. — From  late  April  to  mid  October,  1972,  samples  were  taken  at 
approximately  monthly  intervals  from  contiguous  areas  at  site  2.  Specimens  were  captured 
by  removing  granitic  flakes  with  a  crowbar  (see  Klauber,  1926).  Weights  and 
measurements  were  taken  in  the  laboratory  within  24  hours  of  capture.  Weights  were 
taken  on  a  Mettler  balance  and  read  to  the  nearest  0.01  g.  Snout-vent,  tail,  and  axilla- 
groin  lengths  were  measured  with  a  plastic  millimeter  rule  and  read  to  the  nearest  mm. 
Head  length  (anterior  margin  of  auditory  meatus  to  tip  of  rostrum)  and  head  width 
(greatest  width  anterior  to  auditory  meatus)  were  measured  with  vernier  calipers  to  the 
nearest  0.1  mm.  Caudal  autotomy  was  noted  for  each  lizard,  as  was  the  number  of 
femoral  pores  and  the  sex  of  each  adult.  All  lizards  were  released  unharmed  in  the 
approximate  area  of  capture.  The  same  data,  with  the  exception  of  weight,  were  obtained 
from  specimens  in  the  San  Diego  Society  of  Natural  History. 

Reproduction. — Testes  of  preserved  specimens  were  measured  to  the  nearest  0.01 
mm  with  an  ocular  micrometer.  Testicular  volume  was  calculated  using  the  formula  for 
the  volume  of  an  ellipsoid.  Copulatory  activity  in  recently  captured  lizards  was  recorded, 
and  gravid  females  were  held  in  captivity  until  parturition  to  provide  information  on 
brood  size,  characteristics  of  the  newborn,  and  timing  of  parturition. 


262 


TABLE  2.   Vertebrate  associates  of  Xantusia  henshawi  henshawi. 


Group 


Potential  Competitors 
For  Food 


Potential  Predators 


Relationship  Unknown 


Amphibians 
Lizards 


Snakes 


Birds 


Mammals 


Hyla  regilla 
Bufo  boreas 

Sceloporus  occidentalis 
Sceloporus  orcutti 
Uta  stansburiana 
Urosaurus  microscutatus 
Phrynosoma  coronatum 
Cnemidophorus  tigris 
C.  hyperythrus 
Coleonyx  variegatus 
Phyllodactylus  xanti 


Numerous  insectivorous 
passerines 


Sceloporus  orcutti 


Lichanura  trivergata 
Lampropeltis  getulus 
Masticophis  lateralis 
Salvadora  hexalepis 
Hypsiglena  torquata 
Trimorphodon  vandenburghi 
Crotalus  ruber 
Crotalus  mitchelli 

Faico  sparverius 
Buteo  jamaicensis 
Tyto  alba 
Otus  asio 
Bubo  virginianus 
Geococcyx  californianus 
Corvus  corax 
Aphelocoma  coerulescens 

Neotoma  sp. 
Canis  latrans 


Pituophis  melanoleucas 


Numerous  granivorous 
passerines 


Dipodomys  agilis 
Perognathus  sp. 
Peromyscus  sp. 
Sylvilagus  sp. 
Myotis  subulatus 
Odocoileus  hemionus 


Radiographic  examination. — Following  the  technique  described  by  Etheridge  (1962), 
15  preserved  specimens  were  x-rayed  to  determine  the  presence  or  absence  of  epiphysial- 
diaphysial  fusion  and  to  verify  caudal  autotomy  in  certain  specimens. 

Thermal  biology. — From  14  June  through  19  October,  1972,  lizards  were  captured  at 
night  at  site  1  and  the  following  data  were  recorded:  date  and  time  of  capture,  sex, 
cloacal  temperature,  air  temperature  (1  cm  above  substrate),  and  substrate  temperature. 
Temperatures  were  taken  with  a  Schultheis  rapid  equilibrium  thermometer  and  read  to 
the  nearest  0.1  C.  Cloacal  temperatures  were  read  within  10  seconds  of  capture,  and  were 
taken  only  from  lizards  which  were  abroad  on  boulders. 


RESULTS 

Sexual  dimorphism. — Adult  females  average  6  mm  longer  in  SVL  than  adult  males 
(62  mm  vs.  56  mm;  Fig.  7),  and  weigh  more  (3.3  g  vs.  2.9  g),  but  at  any  given  length, 
males  and  non-gravid  females  weigh  the  same.  I  found  no  sexual  dichromatism  or 
intersexual  differences  in  relative  head  length,  head  width,  axilla-groin  length,  or  tail 
length  of  adults. 

As  in  many  species  of  lizards,  X.  h.  henshawi  has  a  row  of  femoral  pores  along  the 
postero-ventral  margin  of  the  thighs.  In  males  they  are  large  and  produce  an  obvious 
secretion;  in  females  they  are  small  and  inconspicuous  (Fig.  8).  Dissection  of  preserved 
specimens  confirms  that  these  secondary  sex  characters  permit  accurate  sexing  of  adult 


263 


Figure  3.  Crevice  formed  by  the  exfoliation  of  a  granitic  boulder.  Photographed  at  site  1. 

lizards.  I  interpreted  the  presence  of  a  waxy  plug  within  the  pore  as  evidence  of  active 
secretion.  In  preserved  lizards  and  in  living  material,  a  secretory  plug  was  first  evident  in 
males  at  a  SVL  of  42  mm  and  43  mm  respectively.  The  number  of  pores  ranges  from  six 
to  14  per  thigh;  often  the  number  on  one  thigh  exceeds  that  on  the  other  by  two  or  three. 
I  found  no  significant  intersexual  difference  in  mean  number  of  femoral  pores  on  the 
right  thigh  (males  =  11.0,  females  =  11.2,  N  =  100  and  97  respectively,  t  =  1.36,  P> 
0.2). 


0^ 

a; 

a 


^ 


00     02      04 


08 


10       12       14 
Time  of  Day 


16 


-t- 
18 


20      22      24 


Figure  4.  Twenty-four  hour  temperature  cycles  on  a  granitic  boulder.  Solid  circles  indicate  temperature  on 
outer  surface  of  exfoliating  flake;  open  circles  indicate  temperature  at  edge  of  crevice;  triangles  indicate 
temperature  25  cm  inside  crevice.  Recorded  22  September  1972  at  site  1. 


264 


35 


30  O 


25 

0) 

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4-< 

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cu 

L. 

0) 

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JFMAMJJASOND 

Months 

Figure  5.  Ten  year  mean  monthly  maximum  (solid  circles)  and  minimum  (open  circles)  temperatures  and  mean 
precipitation  for  the  period  1957-1966  from  Ramona,  San  Diego  Co.,  California,  13.5  km  from  study  site  2. 
Data  from  U.S.  Weather  Bureau,  Ramona-Spalding  station. 

Population  structure. — No  significant  deviation  from  a  1  :  1  sex  ratio  exists  in  adults 
from  site  2  (X^  =  2.81,  P>  0.05),  or  in  specimens  of  first  year  (21  males,  23  females), 
second  year  (12  males,  11  females),  or  adult  lizards  (X^  =  2.34,  P>0.10)  in  the  SDSNH 
collected  throughout  San  Diego  Co.  Figure  7  presents  frequency  distributions  of  SVL  for 
six  successive  monthly  samples  from  site  2.  For  lizards  which  produce  only  one  brood  per 
season  and  in  which  parturition  occurs  over  a  short  period,  a  SVL  frequency  distribution 


Figure  6.  Study  site  2,  Mount  Woodson,  San  Diego  Co.,  California. 


265 


w 

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September 


30  35  40  45 


55  60  65  70 


Snout-vent    Length  (mm) 

Figure  7.  Population  structure  of  Xantusia  henshawi  henshawi.  For  lizards  44  mm  or  above,  open  bars  are 
females,  shaded  bars  are  males.  Lizards  below  44  mm  were  not  sexed.  Black  bars  indicate  lizards  in  the  first 
year  of  life.  October  sample  is  inverted  to  facilitate  comparison  of  age  classes. 

will  be  polymodal,  with  the  modes  representing  size  classes,  which  in  turn  represent  age 
classes.  Thus  in  the  October  sample,  lizards  29  to  33  mm  SVL  are  deemed  newborn  (see 
reprodution,  below),  those  37  to  40  mm  are  one  year  old,  those  43  to  49  mm  two  years 
old,  and  those  52  mm  and  above  are  three  years  or  older.  Designation  of  age  class 
boundaries  is  sometimes  arbitrary,  especially  for  older  lizards  where  growth  rates  are 
slower,  causing  size  classes  to  overlap.  Thus  in  the  July  sample  I  consider  as  first  year 
lizards  those  with  a  SVL  of  40  mm  or  less.  To  the  extent  that  age  classes  can  be  inferred 
from  size  classes,  Figure  7  indicates  that  in  each  month  lizards  three  years  or  older 
comprised  over  50  per  cent  of  the  sample.  Over  the  entire  six  months,  lizards  in  the  first 
year  of  life  comprised  18.2  per  cent,  second  year  lizards  9.3  per  cent,  and  lizards  in  the 
third  year  or  older,  72.5  per  cent  of  the  total  sample. 


266 


Figure  8.  Femoral  pores  of  adult  male  and   female  Xantusia  henshawi  henshawi.  Pores  are  darkened  for 
emphasis. 

Growth. — A  statistical  approximation  of  growth  is  obtained  by  plotting  the  mean 
SVL  for  each  size  class  against  time.  Results  (Fig.  9,  based  upon  data  in  Fig.  7)  are  only 
as  accurate  as  the  age  class  designations,  which  are  somewhat  arbitrary.  At  birth  mean 
SVL  is  31  mm  and  mean  weight  is  0.44  g;  at  one  year,  39  mm  and  0.89  g;  at  two  years,  47 
mm  and  1.51  g;  and  by  June  of  the  third  year,  when  lizards  are  approximately  32  months 
old,  mean  SVL  is  51  mm  and  mean  weight  is  1.96  g. 

Growth  in  X.  h.  henshawi,  as  in  most  vertebrates,  is  allometric  (Figs.  10,  11). 
Relative  to  SVL,  small  lizards  have  short  tails  and  long  heads.  A  possible  deviation  from 
simple  allometry  is  suggested  in  Figure  11.  Lizards  above  60  mm  apparently  have 
relatively  shorter  tails  than  lizards  of  intermediate  sizes;  possibly  I  failed  to  detect  caudal 
autotomy  in  some  of  the  larger  lizards. 

Contrary  to  the  situation  among  turtles  and  crocodilians,  some  lizards  exhibit 
determinate  growth  which  results  from  fusion  of  the  primary  centers  of  ossification 
(diaphyses)  with  the  secondary  centers  (epiphyses)  of  the  endochondral  bones  (Haines, 
1969).  Such  epiphysial-diaphysial  union  is  illustrated  for  X.  h.  henshawi  in  Figure  12.  Of 
15  lizards  examined  by  radiography,  the  smallest  male  showing  such  union  was  52  mm 
SVL;  the  smallest  female,  58  mm.  These  are  close  to  the  sizes  at  which  I  estimate 
reproductive  maturity  is  attained.  The  approximately  normal  size  distribution  of  adults 
from  site  2  (Fig.  7)  and  the  lack  of  extraordinarily  large  individuals  also  suggests 
determinant  growth  for  this  species. 

Mortality. — Available  life  table  data  for  X.  h.  henshawi  are  inadequate  for 
calculation  of  mortality  rates,  but  an  approximation  of  mortality  can  be  inferred  from  the 
relative  abundance  of  age  classes.  Newborn  lizards  in  the  October  sample  from  site  2 
comprise  28  per  cent  of  that  month's  sample.  If  the  population  is  at  equilibrium,  annual 
mortality  approximates  28  per  cent.  The  abundance,  in  each  sample,  of  lizards  in  the 
third  year  or  older  indicates  that  large  numbers  of  lizards  survive  the  first  two  years  — 
convincing  evidence  that  mortality  is  low. 

In  October,  1970,  in  the  vicinity  of  Jamul,  San  Diego  Co.,  lizards  were  collected  in 
areas  devastated  two  days  previously  by  the  Laguna  Fire.  Lizards  were  frequently  removed 
from  beneath  flakes  that  had  been  blackened  by  fire,  yet  no  dead  or  injured  lizards 
were  found,  nor  did  lizards  seem  less  abundant  than  in  contiguous  unburned  situations. 
Samples  from  a  burned  area  and  an  adjacent  unburned  area,  collected  approximately 
three  months  after  the  fire,  showed  little  length-specific  weight  difference  (Fig.    13), 


267 


First  Year 


Second  Year 


Third  Year 


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50- 


45- 


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16 


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0  N  D  J    F 


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J    J    A  S  0  N   D  J    F 


J    JASONDJ    FMAMJ 


Time  (months) 


Figure  9.  Growth  rate  in  Xantusia  henshawi  henshawi  during  the  first  three  years  of  life.  For  each  sample, 
vertical  line  indicates  range,  horizontal  line  indicates  mean.  Rectangles  encompass  95%  confidence  limits  for 
the  parametric  means.  Number  above  vertical  line  indicates  sample  size. 

indicating  that  lizards  from  burned  and  unburned  areas  are  equally  well  nourished. 

No  incidents  of  predation  on  X.  h.  henshawi  were  observed  during  this  study,  but 
potential  predators  (Table  2)  were  encountered  frequently.  Murray  (1955)  recorded  a 
Sceloporus  orcutti  with  the  hind  portion  of  a  X.  h.  henshawi  protruding  from  its  mouth. 
Petrosaunis  mearnsi  eats  X.  h.  henshawi  in  captivity  (Cozens,  pers.  comm.);  Petrosaurns 
is  widely  sympatric  with  X.  h.  henshawi,  and  occupies  a  similar  microhabitat.  In  this 
study,  Hypsiglena  torquata  was  twice  oberved  at  night  resting  about  one  and  a  half 
meters  above  ground  between  the  leaves  of  a  shrub  and  the  side  of  a  boulder.  In  both 
cases  the  snakes  were  within  a  meter  of  individual  X.  h.  henshawi.  In  captivity  this 
species  readily  devours  X.  h.  henshawi,  as  do  Masticophis  lateralis,  Trimorphodon 
vandenburghi,  Salvadora  hexalepis,  and  Lampropeltis  getulus,  all  of  which  are  sympatric 
with  X.  h.  henshawi. 

Potential  diurnal  avian  predators  include  Falco  sparverius,  Buteo  jamaicensis, 
Geococcyx  califomianus,  Aphelocoma  coerulescens,  and  Corvus  corax.  Due  to  its 
secretive  habits,  X.  h.  henshawi  probably  rarely  falls  prey  to  these  birds.  Nocturnal  birds 
such  as  Bubo  virginianus,  Tyto  alba,  and  Otus  asio  may  occasionally  capture  these 
lizards,  but  Xantusia  are  probably  below  the  usual  size  range  of  prey  items  for  the  former 
two  species. 

Many  rodents  are  omnivorous  (Landry,  1970),  and  I  consider  Neotoma  a  potential 
predator,  particularly  because  of  its  size,  its  propensity  for  rocky  situations,  and  its 
nocturnality  which  undoubtedly  brings  it  into  contact  with  X.  h.  henshawi. 

Caudal  autotomy. — Like  many  lizards,  X.  h.  henshawi  can  lose  and  subsequently 
regenerate  portions  of  the  tail.  It  is  frequently  assumed  that  regenerated  tails  can  be 
distinguished  on  the  basis  of  size,  color,  and  scutellation.  Yet,  Zweifel  and  Lowe  (1966) 
were  sometimes  unable  to  distinguish  regenerated  tails  in  Xantusia  vigilis  without  the  aid 
of  radiographs.  Of  15  X.  h.  henshawi  examined  in  this  study  by  radiography,  however, 


268 


regeneration  was  invariably  associated  with  one  or  more  of  the  above  criteria.  Nonetheless, 
some  autotomy  may  have  gone  undetected.  Thus  data  concerning  the  frequency  of 
autotomy  are  minimal  estimates.  Of  257  sexable  lizards  from  site  2,  159  (61.4%)  have 
experienced  caudal  autotomy  at  least  once,  with  larger  (presumably  older)  lizards  showing 
the  highest  frequency  (Fig.  14).  The  incidence  of  autotomy  was  the  same  in  males  (74  of 
122,  60.0%)  and  females  (85  of  135,  62.5%). 

Reproduction. — Of  75  preserved  males  in  the  SDSNH,  the  smallest  exhibiting 
testicular  enlargement  were  47  mm  SVL.  Because  slight  shrinkage  occurs  in  preservative, 
I  infer  that  sexual  maturity  in  males  is  attained  at  about  50  mm  SVL,  when  lizards  are  in 
their  third  year.  Testicular  volume  varies  considerably  through  the  year  (Table  3). 
Testicular  volume  is  low  during  winter,  maximizes  in  late  spring  and  early  summer,  then 
apparently  declines  in  late  summer  and  fall. 

TABLE  3.     Seasonal  variation  in  testicular  volume  in  Xantusia 
henshawi  henshawi.    N  indicates  number  of  testes  examined. 


Month 

N 

X 

SD 

January 

13 

3.83 

2.17 

March 

5 

7.53 

2.64 

April 

10 

17.01 

10.49 

May 

3 

30.30 

7.14 

June 

5 

19.49 

7.61 

July 

3 

23.18 

5.13 

November 

2 

11.20 

0.01 

The  smallest  gravid  female  I  encountered  was  56  mm  SVL,  suggesting  that  sexual 
maturity  is  attained  at  about  that  length.  Moreover,  females  above  54  mm  SVL  undergo 
pronounced  seasonal  weight  change,  whereas  those  below  54  mm  do  not  (Fig.  15).  Above 
54  mm,  April-May  females  are  significantly  heavier  than  January  females,  and  June 
females  are  significantly  heavier  than  the  April-May  sample.  By  October  weights  have 
regressed  to  slightly  below  January  levels.  This  weight  change  is  probably  the  result  of  fat 
deposition  necessary  to  carry  gravid  females  through  the  hot  dry  period  of  gestation. 
Males  do  not  exhibit  this  seasonal  weight  change.  I  therefore  infer  that  females  are 
sexually  mature  at  about  54-56  mm  SVL,  a  length  attained  late  in  the  third,  or  early  in 
the  fourth  year  of  life. 

I  observed  attempted  copulation  among  recently  captured  lizards  on  18  June,  6  July, 
and  7  July.  The  attempts  were  similar  to  the  general  saurian  pattern,  and  presumably 
reflected  the  timing  of  copulatory  activity  in  the  population  from  which  they  came.  Thus 
breeding  extends  from  at  least  mid  June  through  early  July. 

Like  other  xantusiids,  X.  h.  henshawi  is  viviparous.  Parturition  usually  occurs  from 
mid  September  through  mid  October.  Parturition  dates  for  28  broods  born  to  recently 
captured  females  spanned  the  period  17  September  to  19  October,  but  a  newborn  lizard 
collected  at  site  2  on  3  September  shows  that  parturition  occasionally  occurs  earlier. 
Assuming  that  fertilization  occurs  shortly  after  copulation,  gestation  requires  about  90 
days. 

Population  structure,  timing  of  seasonal  weight  increase,  and  timing  of  copulation 
and  parturition  indicate  that  females  at  site  2  produce  one  brood  per  year.  Because  most 
sexually  mature  females  exhibit  a  pronounced  weight  increase  in  spring,  I  assume  that 
females  are  able  to  breed  each  year. 

Of  28  broods,  13  contained  two  offspring  and  15  contained  one  (X  =  1.46).  For 
broods  consisting  of  two  offspring,  a  strong  positive  correlation  exists  between  the  SVL  of 
the  female  and  the  combined  weight  of  her  offspring  (r  =  0.915,  P<  0.01). 

Thermal  biology. — Sixty-eight  cloacal  temperatures  ranging  from  18.1  to  31.8  C  were 
obtained  from  lizards  abroad  at  night  at  site  2.  Mean  temperatures  for  males  and  females 
do  not  differ  significantly  (25.6  and  25.2  respectively).  Cloacal  temperatures  are  strongly 


269 


26-30    31-35    36-40    41  45   46-50    51-55   56-60    61-65   66-70 

Snout-vent  Length    (mm) 
Figure  10.  AUometric  growth  in  the  head  of  Xantusia  henshawi  henshawi.  Symbols  as  in  Figure  9. 

correlated  with  both  substrate  and  air  temperatures  (Figs.  16  and  17  respectively),  but  the 
higher  coefficient  of  determination  indicates  that  lizards  are  more  closely  coupled  to 
substrate  temperatures  than  to  air  temperatures. 


DISCUSSION  AND  CONCLUSIONS 


Sexual  dimorphism. — Like  many  species  of  lizards,  X.  h.  henshawi  exhibits  sexual 
size  dimorphism,  with  females  larger  than  males.  Because  larger  female  X.  h.  henshawi 
produce  heavier,  presumably  more"  fit  offspring,  this  may  be  an  adaptation  to 
accommodate  relatively  large  embryos. 

Among  lizards,  sexual  dichromatism  is  widespread  and  often  assumed  to  have  a 
social  function.  Dichromatism  therefore  implies  color  vision,  and  the  absence  of  color 
vision  should  preclude  the  evolution  of  social  sexual  dichromatism.  Xantusia  vigilis  and 
Klauberina  riversiana  possess  retinas  adapted  to  conditions  of  low  light  intensity  (Walls, 
1942),  conditions  under  which  visual  information  in  terms  of  color  would  be  difficult  to 
perceive,  and  color  vision  would  be  unlikely  to  evolve.  Both  are  sexually  monochromatic. 
Similarly,  the  secretive  crepuscular-nocturnal  habits  of  X.  h.  henshawi  suggest  the 
absence  of  color  vision  and  probably  account  for  the  monochromatism  of  this  species. 

Males  of  many  species  of  lizards,  including  X.  h.  henshawi,  possess  enlarged  femoral 
pores,  whereas  those  of  the  females  are  rudimentary.  Pores  and  their  secretions  may 
somehow  be  associated  with  sexual  activity  (Atland,  1941;  Bellairs,  1970;  Bostic,  1964; 
Cole,  1966),  but  in  X.  h.  henshawi  femoral  pore  secretion  is  first  evident  in  males  at  a 
SVL  of  about  44  mm  and  thus  preceeds  the  acquisition  of  sexual  maturity. 

Population  structure. — Many  investigators  have  found  unbalanced  sex  ratios  among 
reptiles  (Fitch,  1961).  Among  lizards  such  deviations  usually  favor  females  and  are 
interpreted  as  the  result  of  differential  predation  acting  against  the  more  conspicuous 
territorial  males.  The  expected  1:1  sex  ratio  in  X.  h.  henshawi  for  all  age  classes  indicates 
that  no  such  differential  mortality  is  operating  on  this  species.  This  is  reasonable,  for  in 
this  secretive  monochromatic  species,  males  are  no  more  conspicuous  than  females. 

A  remarkable  feature  of  the  age  class  structure  of  the  X.  h.  henshawi  population  is 
the  high  proportion  of  adults,  implying  a  low  rate  of  turnover.  In  this  X.  h.  henshawi  is 
similar  to  Xantusia  vigilis  in  which  51  per  cent  of  a  winter  population  consisted  of  mature 


270 


26-30     31-35   36-40    41-45  46-50    51-55    56-60    61-65    66-70 

Snout-vent    Length   (mm) 

Figure  1 1 .  Allometric  growth  in  the  tail  of  Xantusia  henshawi  henshawi.  Symbols  as  in  Figure  9. 

individuals  (Zweifel  and  Lowe,  1966).  Such  age  class  structure  is  not  unique  to  Xantusia, 
but  it  is  in  sharp  contrast  with  the  situation  in  many  iguanids  where  high  reproductive 
potential  and  high  mortality  combine  to  produce  populations  in  which  rates  of  turnover 
are  high  and  immature  lizards  are  abundant  relative  to  adults.  The  iguanid  lizard  Uta 
stansburiana  in  west  Texas  is  an  extreme  example  in  which  life  expectancy  is  little  more 
than  a  year,  and  annual  turnover  approaches  100  per  cent  (Tinkle,  1965). 

For  X.  h.  henshawi,  a  species  of  narrow  ecological  tolerance,  limited  geographical 
distribution,  existing  in  an  area  of  low  and  unpredictable  rainfall,  a  high  proportion  of 
long-lived  breeding  adults  is  adaptive  in  that  it  permits  the  population  to  survive  several 
successive  seasons  of  reproductive  failure.  The  probability  of  such  failure  may  be  high,  for 
the    correlation    between    reproductive    failure    and    inadequate    mositure    has    been 


B 


Figure  12.  Epiphyseal-diaphyseal  fusion  in  the  humerus  oi Xantusia  henshawi  henshawi.  A.  46  mm  SVL  B.  49 
mm  SVL  C.  62  mm  SVL. 


271 


^_ 

30 

O) 

4-1 

r. 

U) 

0 

^ 

2.0 

o 
o 
o  o 

o  O  o 


o 

8 


o 


o 


•  t 


o 


H 1 1 h 


H 1 1 h 


H 1 h 


45 


50  5  5  60 

Snout-vent  Length  (mm) 


H — I — h- 

65 


Figure  13.  Comparison  of  the  relationship  between  weight  and  snout-vent  length  in  samples  of  Xantusia 
henshawi  henshawi  from  burned  (solid  circles)  and  unbumed  (open  circles)  areas. 

documented  for  some  lizards  (Mayhew,  1965;  Nagy,  1973),  including  X  vigilis  (Zweifel 
and  Lowe,  1966). 

Growth. — For  its  size,  X.  h.  henshawi  is  an  unusually  slow  growing,  late  maturing 
species;  males  first  breed  at  about  two  and  a  half  years,  females  at  three  and  a  half  years. 
In  this  respect  X.  h.  henshawi  is  identical  to  X  vigilis  (Miller,  1951). 

Like  most  vertebrates,  X.  h.  henshawi  has  a  relatively  larger  head  at  birth  than  at 
adulthood.  This  presumably  results  from  the  concentration  of  sense  organs  and  nerve 
tissue  in  that  area.  Once  ossified,  the  numerous  complex,  interlocking  cranial  elements 
preclude  very  much  expansion  in  the  head  region.  The  biological  significance  of 
allometric  growth  in  tail  length  of  X.  h.  henshawi  is  obscure.  It  might  reflect  an 
ontogenetic  change  in  the  importance  of  the  tail  as  an  organ  for  fat  storage,  or  changing 
demands  on  the  tail  as  an  organ  of  balance  during  locomotion.  Fitch  (1954)  found  similar 
allometric  growth  in  the  tail  of  the  skink,  Eumeces  fasciatus. 

Mortality. — Available  evidence  indicates  that  in  X.  h.  henshawi  mortality  from  all 
causes  is  low,  as  it  must  be  for  a  species  with  low  reproductive  potential.  This  is  so  despite 
the  fact  that  coexisting  with  X.  h.  henshawi  are  numerous  potential  ,predators.  These 
lizards  are  probably  most  vulnerable  when  abroad  at  night,  but  their  light  nocturnal  color 
phase  closely  approximates  their  granitic  background,  rendering  them  difficult  to  detect 
visually.  Hypsiglena  torquata  and  Trimorphodon  vandenburghi,  both  nocturnal,  perhaps 
occasionally  prey  on  X.  h.  henshawi,  but  since  neither  species  is  numerous  they  may  not 
make  serious  inroads  in  the  lizard  population.  Nonetheless,  the  precise  background 
matching  coloration  indicates  that  selection  in  the  form  of  visually  oriented  predation  has 
been  a  significant  factor  in  the  ecology  of  this  species. 

Brush  fires  might  be  a  source  of  mortality  in  X.  h.  henshawi,  as  suggested  by 
Klauber  (1939).  His  conclusion  that  X.  h.  henshawi,  in  contrast  to  Uta  and  Sceloporus, 
was  little  affected  by  fires  agrees  with  my  observations  that  these  lizards  not  only  survive 
fires,  but  remain  well  nourished  after  living  several  months  in  a  burned  area. 

Most  chaparral  fires  in  southern  California  occur  during  late  summer  and  early  fall 
during  periods  of  high  temperature,  a  time  when  X.  h.  henshawi  seeks  refuge  deep  within 
rock  crevices  from  unfavorable  temperatures.  So  situated,  they  are  protected  from  fires 


272 


5   10  15  20  25  30  35  40  45  50  55  60  65  70  75  80 


Per  Cent  With  Regeneration 

Figure  14.  Frequency  of  caudal  autotomy  in  Xantusia  henshawi  henshawi  by  size  class.  Numbers  inside  bars 
indicate  sample  size. 

which,  although  generating  high  temperatures,  are  of  short  duration. 

Because  the  presence  of  granitic  exfoliations  is  important  for  the  existance  of  this 
species,  fires,  rather  than  being  detrimental,  may  be  an  asset;  the  rapid  expansion  of 
boulders  caused  by  the  heat  of  brush  fires  accelerates  exfoliation,  thereby  generating 
more  habitat. 

I  have  no  evidence  that  food  shortage  contributes  to  mortality  in  post-partum  lizards, 
although  it  might  lower  reprodutive  success.  Of  735  lizards  collected  in  this  investigation, 
only  two  were  obviously  malnourished;  this  could  have  been  the  result  of  pathology 
unrelated  to  the  availability  of  food. 

Of  those  species  which  are  potential  competitors  for  food  (Table  2),  most  are  either 
temporally  or  spatially  separated  from  X.  h.  henshawi.  Coleonyx,  Phyllodactylus, 
Urosaurus,  Uta,  Sceloponis,  and  Petrosaurus  are  likely  to  share  the  same  boulder-crevice 
microhabitat,  but  only  the  activity  of  the  first  two  overlaps  both  temporally  and  spatially 
with  that  of  X  h.  henshawi,  which  tend  to  be  active  within  crevices  during  late  afternoon 
and  early  evening  and  quiescent  while  abroad  on  boulders  at  night  (Lee,  1974).  In  the 
area  of  sympatry  between  X.  h.  henshawi  and  Coleonyx,  the  latter  is  nowhere  numerous, 
and  Phyllodactylus  overlaps  X.  h.  henshawi  geographically  only  on  the  desert  slopes  of 
mountains.  Thus,  although  potential  competitors  for  food  are  numerous,  actual 
interspecific  competition  —  if  in  fact  food  is  limiting  —  is  rare. 

Caudal  autotomy. — A  high  frequency  of  caudal  autotomy  suggests  heavy  predation 
pressure,  but  such  a  conclusion  is  inconsistent  with  the  ecology  of  X.  h.  henshawi,  for 
actual  predator  species  are  few,  and  individuals  never  abundant.  Observations  by 
Heimlich  and  Heimlich  (1947),  Lowe  (1948),  and  Brattstrom  (1952)  provide  an  alternative 
explanation.  Both  Heimlich  and  Heimlich,  and  Brattstrom  found  tails  of  conspecifics  in 
the  stomachs  of  Xantusia  vigilis;  Lowe  reported  agonistic  interactions  among  captive  X. 
vigilis  involving  biting;  and  Zweifel  and  Lowe  (1966)  concluded  that  intraspecific  fighting 
accounted  for  most  of  the  autotomy  observed  by  them  during  their  nine  year  study  of  X. 
vigilis  in  the  Mojave  Desert.  My  observations  on  captive  X.  h.  henshawi  agree  closely  with 
those  of  Lowe  (1948)  for  X.  vigilis.  Among  captive  X.  h.  henshawi,  fights  were  common 
and  involved  twitching  of  the  tail  and  biting,  with  the  bites  often  directed  toward  the  base 
of  the  tail.  Several  lizards  exhibiting  recent  autotomy  undoubtedly  lost  their  tails  in  this 
manner.  The  high  incidence  of  caudal  autotomy  in  X.  h.  henshawi  probably  is  more  a 
reflection  of  intraspecific  fighting  than  a  high  rate  of  attempted  predation. 


273 


6.0 


5.5 


5.0 


4.5 


4.0 1 
r       3.5 

i 

3.0  + 


2.5 


2.0 


1.5 


•  April-May 

o  June 

A  October 

▲  January 


^-^-^  /^A  A    A  A  ^ 


^A 


— I — I— I — till 1 — I 1 — I — I 1 1 — I — I — I — I — I 1— I — I — I—" — I — 

44   46  48   50   52   54   56   58   60  62   64  66   68 


Snout-vent  Length  (mm) 


Figure  15.  Relationship  between  weight  and  snout-vent  length  for  female  Xarttusia  henshawi  henshawi  collected 
at  different  times  of  the  year.  Below  54  mm  95%  confidence  limits  for  all  regressions  broadly  overlap.  Above  61 
mm  confidence  limits  for  January  entirely  overlap  October  and  slightly  overlap  April-May. 

Reproduction. — Mean  brood  size  for  X  h.  henshawi  is  lower  than  that  reported  for 

Xantusia  vigilis  by  Zweifel  and  Lowe  (1966)  (L46  and  L87  respectively).  Because  female 

X.  h.  henshawi  probably  require  three  years  to  reach  reproductive  maturity,  produce  only 

a  single  brood  per  season,  and  average  L46  offspring  per  brood,  this  species  has  the 

lowest  reproductive  potential  (in  the  sense  of  Ballinger,  1973)  reported  for  any  lizard. 

However,  the  difference  in  brood  size  between  X.  h.  henshawi  and  X.  vigilis  may  not  be 

significant,  because  Zweifel  and  Lowe  (1966)  demonstrated  a  strong  positive  correlation 

between  brood  size  and  amount  of  winter  precipitation.  If  a  similar  relationship  exists  for 

X.  h.  henshawi.  my  estimate  of  brood  size,  based  upon  lizards  collected  in  1972,  is  a 

minimal  one.  Precipitation  for  the  winter  of  1971-72  in  San  Diego  Co.  was  well  below 

normal. 

The  low  reproductive  potential  of  X.  h.  henshawi  may  be  contrasted  with  that  of 

Sceloporus  olivaceous,  a  species  with  perhaps  the  highest  known  reproductive  potential  of 


274 


(r 

I- 
< 

IT 
UJ 

a. 

UJ 


< 

< 
O 


34 

32 

V* 

30 

Y=  920X+2.22 

^ 

• 

r'=.966 

.  *'r 

28 

•••/  • 

*w 

26 

• 

•  • 

24 

••^^ 

•         .*. 

22 

•.V 

20 

.  y 

18 

— 1 — 1 — 1 — 1 — 1 — 1 — 1 — 

— 1 1 1 1 1 H 

— 1 \ 1 1 

18 


20       22 


24       26        28        30 


32 


34 


< 

< 
o 


34 
32 

30 
28 
26 
24 

22 
20 
18 


Y=  .737X+8  47 
r'  =  .679 


—I — I — I — I — I — I — I — t — 1 — 
16         18        20        22        24 


26 


— ( — t — » — \ — I — 
28        30        32 


ROCK   SURFACE    TEMPERATURE 


AIR    TEMPERATURE     C 


Figure    16.  Regression   of  cloacal    temperature    of  Figure    17.  Regression   of  cloacal    temperature    of 

Xantusia  henshawi  henshawi  against  rock  surface  XaMfus/a  AeniAawj  AensAam  against  air  temperature, 

temperature. 

any  lizard.  In  that  species,  yearling  females  may  produce  four  clutches  of  eggs  per  season 
with  a  mean  clutch  size  of  11.3  eggs.  Clutch  size  increases  to  18.4  in  two-year-olds,  and 
24.5  in  lizards  three  years  of  age  (Blair,  1960).  Theoretically,  a  female  Sceloporus 
olivaceous  surviving  through  the  third  breeding  season  could  produce  about  217  offspring 
in  the  time  it  would  take  X.  h.  henshawi  to  produce  one  or  two.  Mortality,  of  course,  is 
very  different  in  these  two  species.  An  average  of  75  per  cent  of  Sceloporus  olivaceous 
eggs  fail  to  hatch,  and  mean  life  expectancy  at  hatching  is  about  three  months  (Blair, 
1960).  In  the  viviparous  X.  h.  henshawi,  developing  embryos  are  protected,  reducing 
mortality  during  development,  and  post-partum  mortality  is  low,  allowing  females  a 
protracted  breeding  life. 

On  the  basis  of  reproductive  strategy  and  attendant  life  history  characteristics, 
Tinkle  (1969)  and  Tinkle  et  al.  (1970)  have  identified  two  categories  of  lizards.  One 
contains  small  species  that  are  early-maturing,  short-lived,  and  highly  fecund  (e.g., 
Sceloporus  olivaceous);  the  other  contains  larger  species  that  are  late-maturing,  have  long 
life  expectancy,  and  produce  few  offspring  per  season.  Members  of  the  former  category 
are  relatively  r-selected  (Pianka,  1970);  they  allocate  large  amounts  of  energy  for 
reproduction,  produce  many  offspring,  but  apportion  little  energy  per  individual 
offspring.  Members  of  the  latter  category  are  relatively  A'-selected,  channeling  energy  into 
production  of  a  few  highly  fit  offspring.  Except  for  size,  X.  h.  henshawi  typifies  the  latter 
category  and  is  a  highly  AT-selected  species. 

Thermal  biology. — In  previous  studies  dealing  with  the  thermal  biology  of  X.  h. 
henshawi  (Brattstrom  1965;  Mautz  and  Case,  1974;  Scott,  1971)  cloacal  temperatures 
obtained  from  lizards  in  the  field  were  apparently  taken  only  during  daylight  hours,  and 
from  lizards  that  had  been  removed  from  crevices.  In  the  present  study,  cloacal 
temperatures  obtained  at  night  from  lizards  abroad  on  boulders  indicate  that  X.  h. 
henshawi  is  highly  dependent  on  rock  surface  temperatures;  thus,  Brattstrom's  (1965) 
designation  of  this  species  as  a  thigmotherm  is  appropriate. 

Cowles  and  Bogert  (1944)  define  the  normal  activity  range  of  temperatures  in  reptiles 
as  ".  .  .  the  thermal  range  extending  from  the  resumption  of  ordinary  routine  (after  the 
animal  has  ceased  basking,  in  the  case  of  diurnal  forms)  and  terminates  at  a  point  just 
below  the  level  at  which  high  temperatures  drive  the  animal  to  shelter."  Elsewhere  (Lee, 
1974)  I  show  that  maximum  activity  in  this  species  occurs  before  lizards  issue  forth  from 
their  crevices  at  night.  Therefore,  the  range  of  temperatures  recorded  for  these  lizards 
abroad  at  night  may  not  reflect  the  normal  activity  range,  but  rather  the  range  over  which 
a  relatively  quiescent  portion  of  the  activity  cycle  occurs.  Thus,  my  data  cannot  be  directly 


275 


compared  with  published  data  on  normal  activity  ranges  of  lizards.  It  is  of  interest, 
however,  to  compare  them  with  the  range  of  temperatures  tolerated  voluntarily  during  the 
day  as  determined  by  Scott  (1971).  The  close  correspondence  between  Scott's  data  (18.6 
to  33.0  C,  X  =  26.0)  and  mine  (18.1  to  31.8  C,  X  =  25.3)  indicates  that  in  general, 
during  the  warmer  part  of  the  year,  lizards  maintain  body  temperatures  within  about  the 
same  limits  both  day  and  night.  This  species  is  clearly  eurythermic,  tolerating  a  wide 
range  of  body  temperatures  and  with  no  clearly  defined  preferred  temperature. 

The  biology  of  Xantusia  henshawi  compared  with  that  of  other  xantusiids. — The 
Xantusiidae  are  a  small,  apparently  ancient  family  of  unknown  origin  (Bezy,  1972). 
Savage  (1963)  recognized  four  living  genera:  Cricosaura,  monotypic  and  known  only  from 
Cabo  Cruz,  Oriente  Province,  Cuba;  Klauberina,  also  monotypic  and  restricted  to  the 
Channel  Islands  off  the  coast  of  southern  California;  Lepidophyma,  with  perhaps  12 
species,  confined  to  southern  Mexico  and  Central  America;  and  Xantusia,  with  two 
species,  restricted  to  northeastern  Mexico  and  the  southwestern  United  States. 

Ecologically,  the  family  is  relatively  homogeneous.  Xantusiids  tend  to  be  small, 
secretive,  and  terrestrial.  So  far  as  known,  all  are  viviparous  and  most  have  low 
reproductive  potential  (Goldberg  and  Bezy,  1974).  As  a  group  they  are  relatively 
AT-selected.  Xantusia  henshawi  henshawi  and  Xantusia  vigilis  are  especially  similar  in 
reproductive  biology,  population  structure,  growth  (Miller,  1951),  and  thermal  biology 
(Kour  and  Hutchinson,  1970). 

Xantusiids  exhibit  highly  disjunct,  frequently  relictual  distributions,  and  often  have 
specialized  microhabitat  requirements  (rock  crevices,  fallen  yuccas,  decomposing  logs  in 
humid  tropical  forests).  Bezy  (1972)  views  this  as  a  response  to  increasing  aridity  during 
the  Tertiary.  Perhaps  an  additional  factor  has  been  competition  with  other  lizards.  If 
xantusiids  are  competitively  inferior,  the  few  existing  species  are  those  that  have  survived 
the  Tertiary  radiations  of  other  lizard  groups  by  avoiding  competition,  either  by  increased 
specialization  (e.g.,  Xantusia  henshawi  henshawi)  or  by  fortuitous  establishment  on  (or 
restriction  to)  islands  (e.g.,  Klauberina). 


ACKNOWLEDGEMENTS 

Roger  Carpenter,  Richard  Etheridge,  and  Paul  Nichols  critically  reviewed  portions  of  the  manuscript. 
Michael  U.  Evans  took  the  photographs  which  appear  as  Figures  3  and  6.  Janet  Lee  provided  financial  support 
for  this  study.  This  report  is  a  portion  of  a  thesis  submitted  to  the  faculty  of  San  Diego  State  University  in 
partial  fulfillment  of  the  requirements  for  the  degree  of  Master  of  Science. 


LITERATURE  CITED 

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Atsatt,  S.R. 

1925.  Observations  on  \\\\ng  Xantusia  henshawi.  Copeia  1925:71-72. 
Atsatt,  S.R. 

1939.  Color  changes  controlled  by  temperature  and  light  in  the  lizards  of  the  desert  regions  of  southern 
California.  Univ.  California  (Los  Angeles)  Pub.  Biol.  Sci.  1:237-276. 
Ballinger.  R.B. 

1973.  Comparative  demography  of  two  viviparous  iguanid  lizards   (Sceloporus  jarrovi  and  Sceloporus 
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Bellairs,  A. 

1970.  The  life  of  reptiles.  2  vols.  Universe  Books,  New  York.  590  p. 
Bezy,  R.L. 

1972.  Karyotypic  variation  and  evolution  of  the  lizards  of  the  family  Xantusiidae.  Los  Angeles  Co.  Mus. 
Nat.  Hist.,  Contrib.  Sci.  227:1-29. 
Blair,  F.W. 

1960.  The  rusty  lizard.  University  of  Texas  Press,  Austin.  185  p. 


276 


Bostic.  D.L. 

1964.  The  ecology  and  behavior  oi  Cnemidophorus  hyperythrus  beldingi  (Sauria:  Teiidae).  Master's  thesis, 
San  Diego  State  University.  112  p.  * 

Brattstrom.  B.H. 

1951.  The  number  of  young  of  Xantusia.  Herpetologica  7:143-144. 

Brattstrom,  B.H. 

1952.  The  food  of  the  night  lizards,  genus  Xantusia.  Copeia  1952:168-172. 
Brattstrom,  B.H. 

1965.  Body  temperatures  of  reptiles.  Am.  Midi.  Nat.  73:376-422. 
Cole,  C. 

1966.  Femoral  glands  in  lizards:  a  review.  Herpetologica  22:199-206. 
Cowles,  R.B.  and  CM.  Bogert. 

1944.  A  preliminary  survey  of  the  thermal  requirements  of  desert  reptiles.  Am.  Mus.  Nat.  Hist.,  Bull. 
83:265-296. 
Etheridge,  R. 

1962.  Skeletal  variation  in  the  iguanid  lizard  Sator  grandaevus.  Copeia  1%2:613-619. 
Fitch,  H.S. 

1954.  Life  history  and  ecology  of  the  five-lined  skink,  Eumeces  fasciatus.  Univ.  Kansas  Publ.  Mus.  Nat. 
Hist.  8:1-156. 

Fitch,  H.S. 

1961.  Longevity  and  age-size  group  in  some  commons  snakes.  In:  Vertebrate  speciation,  a  University  of 
Texas  symposium.  University  of  Texas  Press,  Austin. 
Fitch,  H.S. 

1970.  Reproductive  cycles  in  lizards  and  snakes.  Univ.  Kansas  Mus.  Nat.  Hist.,  Misc.  Publ.  52:1-247. 
Goldberg,  S.R.  and  R.L.  Bezy. 

1974.  Reproduction  in  the  island  night  lizard,  Xantusia  riversiana.  Herpetologica  30:350-360. 
Grinnell,  J.  and  C.L.  Camp. 

1917.  A  distributional  list  of  the  amphibians  and  reptiles  of  California.   Univ.  California  Publ.  Zool. 
17:127-208. 
Haines,  R.W. 

1969.  Epiphyses  and  sesamoids.  In:  Cans,   C,   Bellairs,   A.,   and  T.S.   Parsons  (eds.).   Biology  of  the 
Reptilia.  vol.  1.  Academic  Press,  New  York.  373  p. 

Heimlich,  E.M.  and  M.G.  Heimlich. 

1947.  A  case  of  cannibalism  in  captive  Xantusia  vigilis.  Herpetologica  3:149-150. 
Klauber,  L.M. 

1926.   Field  notes  on  Xantusia  henshawi.  Copeia  1926:115-117. 
Klauber,  L.M. 

1931.  A  new  species  of  Xantusia  from  Arizona,  with  a  synopsis  of  the  genus.  San  Diego  Soc.  Nat.  Hist., 
Trans.  7:1-16. 
Klauber,  L.M. 

1939.   Studies  of  reptile  life  in  the  arid  southwest.  Zool.  Soc.  San  Diego,  Bull.  15:1-23. 
Kour,  E.L.  and  V.H.  Hutchison. 

1970.  Critical  thermal  tolerances  and  cooling  rates  of  lizards  from  diverse  habitats.  Copeia  1970:219-229. 
Landry,  S.O. 

1970.  The  rodentia  as  omnivores.  Quart.  Rev.  Biol.  45:351-372. 
Lee,  J.C. 

1974.  The  diel  activity  cycle  of  the  lizard,  Xantusia  henshawi.  Copeia  1974:934-940. 
Lowe,  C.H. 

1948.  Territorial  behavior  in  Xantusia  vigilis.  Herpetologica  4:221-222. 
Mautz,  J.  and  T.J.  Case. 

1974.  A  diurnal  activity  cycle  in  the  granite  night  lizard,  Xantusia  henshawi.  Copeia  1974:243-251. 
Mayhew,  W.W. 

1965.  Comparative  reproduction  in  three  species  of  the  genus  Uma.  In:  W.W.  Milstead  (ed.).  Lizard 
ecology,  a  symposium.  University  of  Missouri  Press,  Columbia.  300  p. 
Miller,  M. 

1951.  Some  aspects  of  the  life  history  of  the  yucca  night  lizard,  Xantusia  vigilis.  Copeia  1951:114-120. 
Murray,  K.F. 

1955.  Herpetological  collections  from  Baja  California.  Herpetologica  11:33-48. 
Nagy,  K.A. 

1973.  Behavior,  diet,  and  reproduction  in  a  desert  lizard,  Sauromalus  obesus.  Copeia  1973:93-102. 
Pianka,  E.R. 

1970.  On  rand  A"  selection.  Am.  Nat.  104:592-597. 
Savage,  J.M. 

1963.  Studies  on  the  lizard  family  Xantusiidae.  IV.  The  genera.  Los  Angeles  Co.  Mus.  Nat.  Hist.,  Contrib. 
Sci.  71:1-38. 


277 


Scott,  R. 

1971.  Thermal  biology  of  the  granite  night  lizard.  Master's  thesis,  San  Diego  State  University. 
Shaw,  C.E. 

1949.  Notes  on  broods  of  two  xantudiids.  Herpetologica  5:23-26. 
Stebbins,  R.C. 

1954.  Amphibians  and  reptiles  of  western  North  America.  McGraw-Hill  Book  Co.  Inc.,  New  York.  536  p. 
Stebbins,  R.C. 

1966.  A  field  guide  to  western  reptiles  and  amphibians.  Houghton  Mifflin  Co.,  Boston.  279  p. 
Stejneger,  L.  and  T.  Barbour. 

1943.  Checklist  of  North  American  amphibians  and  reptiles.  5th  ed.  Mus.  Comp.  Zool.,  Bull.  93(1):  1-260. 
Stephens,  F. 

1921.  An  annotated  list  of  the  amphibians  and  reptiles  of  San  Diego  County,  California.  San  Diego  Soc. 
Nat.  Hist.,  Trans.  3:57-69. 
Tinkle,  D. 

1965.  Home  range,  density,  dynamics,  and  structure  of  a  Texas  population  of  the  lizard  Uta  stansburiana. 
In:  W.W.  Milstead  (ed.)  Lizard  ecology,  a  symposium.  University  of  Missouri  Press,  Columbia.  300  p. 

Tinkle,  D. 

1969.  The  concept  of  reproductive  effort  and  its  relation  to  the  evolution  of  life  histories  of  lizards.  Am. 
Nat.  103:501-516. 

Tinkle,  D.,  Wilber,  H.M.  and  S.G.  Tilley. 

1970.  Evolutionary  strategies  in  lizard  reproduction.  Evolution  24:55-74. 
Walls,  G. 

1942.  The  vertebrate  eye  and  its  adaptive  radiation.  Cranbrook  Inst.  Sci.  785  p. 
Webb,  R. 

1970.  Another  new  night  lizard  {Xantusia)  from  Durango,  Mexico.  Los  Angeles  Co.  Mus.  Nat.  Hist., 
Contrib.  Sci.  194:1-10. 
Zweifel,  R.  and  C.  Lowe. 

1966.  The  ecology  of  a  population  oiXantusia  vigilis,  the  desert  night  lizard.  Am.  Mus.  Novitates  2247:1-57. 


San  Diego  State  University,  San  Diego,  California.  Present  address: 
Museum  of  Natural  History  and  Department  of  Systematics  and  Ecology, 
University  of  Kansas,  Lawrence,  Kansas  66045. 


S/W   Gqc^^ 


AU6  1  9 1970 

HARVAWO 


TRAN 


OF  THE 

SOCIETY   OF  p 
NATURAL  hist' 


IONS 


VOL.  17,  NO.  20       16  MAY  1975 


A  CAT' 


p.  2 


p.i 

r 

p.  i 


p.  i 


A  CATALOGUE  OF  MURICACEAN  GENERIC  TAXA 


ERRATA 

p.  279,  last  line  -  should  read  —  monotypy;  ^  -extinct  genus 
p,  281,  right  column,  immediately  after  line  52  insert  — 

CYMIA  Morch,  1860:  97,  98 
Type  sp.  (M.)'  Cuma  sulcata 
Swainson,  1340,  A  Treatise  on 
Malacology,  p.  87,  fig.  4 
(=  Buccinum  tectum  Wood,  1828) 
p,  283,  right  column,  line  34  -  should  read  —  LYROTYPHIS. 
p.  234.  left  column,  line  35  —  delete  whole  line 

left  column,  line  44  —  acute  accent  over  "e"  in  Linne. 
right  column,  line  59  —  "Jousseaume"  instead  of  ''Jousseaum". 
p.  286,  left  column,  line  43  —  should  read  ~  lieth.  pi.  436,  fig.  1, 

Liste,  p.  8. 
right  column,  line  1  —  should  read  —  figs,  la,  lb,  Liste,  p.  1. 
right  column,  line  10  ~  "non-biononinal"  should  be  -  "non-binominal'' 
p.  288,  right  column,  line  53  —  "Collected"  should  begin  with  a  lower 

case  letter, 
p.  289,  right  column,  line  59  —  "recent"  should  begin  with  an  upper  case 

letter. 


SAN  DIEGO  SOC.  NAT.  HIST.,  TRANS.  17(20):  279-292,  16  MAY  1975. 


A  CATALOGUE  OF  MURICACEAN  GENERIC  TAXA 


GEORGE  E.  RADWIN  AND  ANTHONY  D'ATTILIO 


ABSTRACT. — A  compilation  of  genus-level  taxa  and  their  type-species  is  provided,  including  the 
mode  of  type  designation  and  references  to  the  original  descriptions  of  the  genera  and  the  type-species. 

Despite  long-standing  and  recently  intensified  interest  in  muricacean  mollusks,  no 
compilation  of  generic  taxa  is  currently  available.  The  most  complete  listing  (Wenz,  1941) 
lacks  the  depth  needed  by  taxonomists  and  other  workers.  Wenz  listed  only  223  taxa  and 
he  treats  half  of  these  as  synonyms,  for  which  he  does  not  supply  type-species;  where  he 
does  note  type-species  he  rarely  cites  the  mode  of  type  designation  or  gives  references  to 
these  type-species.  Yokes  (1971)  presented  a  more  complete  listing,  at  the  species  level, 
for  several  subfamilies  of  the  Muricidae,  and  Keen  (1944)  catalogued  the  Typhinae  at  the 
specific  and  generic  levels. 

In  preparing  a  guide  to  the  Muricidae,  we  compiled  nomenclatural  data  on  genera  in 
that  and  other  muricacean  families.  They  are  catalogued  below.  In  the  first  section  we  list 
nominal  generic  and  subgeneric  muricacean  taxa,  most  cited  from  primary  sources, 
arranged  alphabetically,  including  type-species  designations  and  original  references.  The 
list  is  essentially  complete  through  1974.  In  the  second  section  we  present  a  bibliography 
of  works  in  which  muricacean  generic  taxa  have  been  introduced.  We  have  supplied 
complete  citations  to  generic  references,  abbreviated  references  of  type-species  and 
subsequent  type  designations.  As  this  catalogue  is  intended  to  serve  as  a  reference  work, 
rather  than  as  a  vehicle  for  our  taxonomic  opinions,  we  have  avoided  all  but  the  most 
essential  comments  on  synonymy.  The  following  abbreviations  are  used:  O,  D. — type 
species  by  original  designation;  S.  D. — type  species  by  subsequent  designation;  M. — type 
species  by  monotypy;  T. — type  species  by  tautonymy;  S.  M. — type  species  by  subsequent 
monotypy; — extinct  genus. 


SAN  DIEGO  SOC.  NAT.  HIST.,  TRANS.  17(20):  279-292,  16  MAY  1975. 


280 


GENERIC  AND  SUBGENERIC  TAXA  OF  THE  MURICACEA 


AARONIA  A.  H.  Verrill,  1950:  4 

Type  sp.  (O.  D.):  Murex  (Aaronia)  strausi  Verrill, 
1950,  Min.  Conch.  Club  S.  California  103:  4. 

ACANTHINA  Fischer  de  Waldheim,  1807:  174 
Type  sp.  (S.  D.,  Gray,  1847b):  Buccinum  mono- 
cerus   Chemnitz,    1788    (=    Buccinum    monodon 
Pallas,   1774).  Neues  Systematisches  Conchylien- 
Cabinet  10:  197,  pi.  154,  figs.  1469,  1470). 

tACANTHlNELLA  Shuto,  1969:  109 

Type  sp.  (O.  D.):  Acantina  (sic.)  _/ava«a  Martin, 
1899,  Samml.  Geol.  Mus.  Leiden,  N.  F.  1:  109. 

ACANTHINUCELLA  Cooke,  1918:  8 

Type  sp.  (O.  D.):  Acanthina  punctulata  (Sowerby, 
1835),  Proc.  Zool.  Soc.  London  3:  50. 

tACANTHOLABIA  Olsson  &  Harbison.  1953:  252 
Type  sp.  (O.  D.):  Acantholabia  floridana  Olsson 
&  Harbison,  1953,  Acad.  Nat.  Sci.  Phila.  Monogr. 
8:  251.  pi.  33,  fig.  10. 

ACANTHOTROPHON  Hertlein  &  Strong,  1951:  86 
Type  sp.  (O.  D.):  Trophon  (Acanthotrophon) 
sorensoni  Hertlein  &  Strong.  1951,  pt.  X.  Zoo- 
logica  36(2):  86,  p.  2.  fig.  1. 

ACTINOTROPHON  Dall,  1902:  534 

Type  sp.  (M.):  Trophon  (Boreotrophon)  actino- 
phorus  Dall,  1889.  Mus.  Comp.  Zool.  Harvard 
18:  206. 

ACUPURPURA  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Murex  tenuispina  Lam.,  1822, 
(=  M.  pecten  Lightfoot,  1786)  Hist.  Nat.  Anim. 
s.  Vert.  7:  158. 

ADAMSIA  Dunker,  1857:  357  (Not  Forbes,  1840) 
Type  sp.  (O.  D.):  Adamsia  typica  Dunker,   1857 
(=  Purpura  tritonifonnis  Blainville,  1832),  Proc. 
Zool.  Soc.  London  25:  357. 

AFRITROPHON  Tomlin,  1947:  271 

Type  sp.  (O.  D.):  Trophon  kowieensis  Sowerby. 
1901,  Proc.  Malac.  Soc.  London  4:  213.  pi.  22. 
fig.  16. 

AGNEWIA  Tenison-Woods,  1878:  29 

New  name  for  Adamsia  Dunker,  1857.  not  Forbes 
1840. 

tALDRICHIA  K.  Palmer.  1937:  262 

Type  sp.  (O.  D.):  Murex  cancellaroides  Meyer  & 
Aldrich,  1886  (=  Muricopsis  aldrichi  Cossmann, 
1903,  new  name  for  M.  cancellaroides  Meyer  & 
Aldrich,  1886,  not  Grateloup,  1833),  J.  Cinn.  Soc. 
Nat.  Hist.  9:  44,  pi.  2  fig.  15. 

ALIPURPURA  P.  Fischer,  1884:  641 

Type  sp.  (O.  D.):  Murex  acanthopterus  Lamarck, 
1822,  Hist.  Nat.  Anim.  s.  Vert.  7:  165. 

ANATROPHON  Iredale.  1929a:  186 

Type  sp.  (O.  D.):  Trophon  sarmentosa  Hedley  & 
May,  1908,  Rec.  Australian  Mus.  7:  121. 

ANTIMUREX  Cossmann,  1903:  12 

New  name  for  Crassilabrum  Jousseaume,  1880. 
not  Megerle  von  Muhifeid  (Ms)  in  Scudder,  1882. 
(This  unnecessary  replacement  name  was  intro- 
duced by  Cossmann  on  the  assumption  that 
Megerle's  manuscript  genus  Crassilabrum  pre- 
occupied Crassilabrum  Jousseaume.  In  actuality 
Megerle's  (Ms)  name  was  not  validated  until 
Scudder  (1882),  thus  making  it  junior  to  Jous- 
seaume's  taxon.) 

APIXYSTUS  Iredale,  1929a:  185 

Type  sp.  (O.  D.):  Trophon  stimuleus  Hedley. 
1907,  Rec.  Australian  Mus.  6:  293,  pi.  55,  fig.  19. 


ARADOMUREX  Coen,  1929:  1281 

Type  sp.  (O.  D.):  Murex  sophiae  Aradas  &  Benoit. 

1870,  Conch.  Viv.  Mar.  Sicil.  p.  270-271,  pi.  5, 

fig.  7. 
ARANEA  Perry,  1810:  225  (not /lra«ea  Linne,  1758) 

Type  sp.  (M.):  Aranea  gracilis  Perry,  1810,  The 

Arcana,  p.  225,  pi.  47. 
ASPELLA  Morch.  1877:  24 

Type  sp.  (M.):  Ranella  anceps  Lamarck,    1822, 

Hist.  Nat.  Anim.  s.  Vert.  7:  154. 
ATTILIOSA  Emerson.  1968a:  380 

Type  sp.  (O.  D.):  Coralliophila  incompta  Berry, 

1960,  Leaflets  in  Malacology  1(19):  119-120. 
AUSTROTROPHON  Dall,  1902:  534.  548 

Type  sp.  (S.  D.  Grant  &  Gale.   1931):   Trophon 

cerrosensis  Dall.  1891,  Proc.  U.  S.  Natl.  Mus.  14: 

181,  pi.  5,  figs.  5.  7. 
AXYMENE  Finlay.  1927:  426 

Type  sp.  (O.  D.):  Trophon  (A.)  turbator  Finlay. 

1927,  Trans.  Proc.  New  Zealand  Inst.  57:  426,  pi. 

23.  figs.  127.  128. 
AZUMAMORULA  Emerson.  1968b:  380 

New  name  for  Morulina  Dall,   1923.  not  Borner. 

1906. 
BABELOMUREX  Coen.  1922:  68 

Type  sp.  (O.  D.):  Fusus  babelis  Requien,   1848, 

Cat.  des  Coquilles  de  Tile  de  Corse,  pt.  49  p.  76, 

sp.  549. 
BASSIA  Jousseaume,  1880:  335  (not  Quoy  & 
Gaimard,  1830) 

Type  sp.  (O.  D.):  Murex  stainforthi  Reeve,  1842, 

Proc.  Zool.  Soc.  London  9:  104. 
BASSIELLA  Wenz,  1941:  1089 

New  name  for  Bassia  Jousseaume.  1880.  not  Quoy 

&  Gaimard.  1830. 
BATHYMUREX  Clench  &  Perez  Farfante.  1945:  41 

Type  sp.  (O.  D.):  Bathymurex  atlantis  Clench  & 

Perez  Farfante.   1945.  Johnsonia  17:  41,  pi.  21, 

figs.  3-5. 
BEDEVA  Iredale.  1924:  183 

Type  sp.  (O.  D.):  Trophon  hanleyi  Angas.   1867 

(=    Trophon  paivae  Crosse.    1864).   Proc.   Zool. 

Soc.  London  31:  110,  pi.  13,  fig.  1. 
BEDEVINA  Habe,  1946:  198 

Type  sp.  (O.  D.):  Trophon  birilefft  Lischke.  1871, 

Malak.  Bi.  18:  39. 
BENTHOXYSTUS  Iredale.  1929a:  185 

Type  sp.  (O.  D.):  Trophon  columnarius  Hedley  & 

May,  1908.  Rec.  Australian  Mus.  7:  121,  pi.  24, 

fig.  22. 
BIZETIELLA  Radwin  &  D'Attilio.  1972:  341 

Type  sp.  (O.  D.):  Tritonalia  carmen  Lowe,  1935. 

Trans.  San  Diego  Soc.  Nat.  Hist.  8(6):  20.  pi.  2 

fig.  6. 
BOLINUS  Pusch.  1837:  134 

Type  sp.  (O.  D.):  Murex  brandaris  Linne,   1758, 

Syst.  Nat.,  Ed.  10,  p.  747,  no.  446. 
BOREOTROPHON  P.  Fischer.  1884:  640 

Type  sp.  (M.):  Murex  clathrata  Linne,  1767,  Syst. 

Nat.,  Ed.  12,  p.  1223,  no.  563. 
BRONTA  Pusch,  1837:  130 

New    name    for    Brontes    Montfort,     1810,     not 

Fabricius,  1801. 
BRONTES    Montfort,    1810:    623    (not    Fabricius, 
1801) 

Type  sp.  (O.  D.):  Brontes  haustellum  Montfort, 


281 


1810.  (—  Murex  haustellum  Linne,  1758)  Conch. 
Syst.  2:  623.  pi.  622. 

BRONTESIA  Reichenbach.  1828:  91 

New  name  for  Brontes  Montfort,  1810,  not 
Fabricius,  1801. 

CALCITRAPESSA  Berry,  1959:  13 

Type  sp.  (O.  D.):  Murex  leeamis  Dall,  1890,  Proc. 
U.  S.  Nat.  Mus.  12:  329,  pi.  7,  fig.  1. 

CALOTROPHON  Hertlein  and  Strong,  1951:  87 
Type  sp.  (M.):  Calotrophon  bristolae  Hertlein  & 
Strong.   1951   (=    Tritonalia  turrita   Dall,    1919), 
Zoologica  36(2):  87,  pi.  2.  fig.  2. 

CANRENA  Link,  1807:  126 

Type  sp.  (O.  D.):  Murex  neritoideus  Gmelin, 
1791,  Syst.  Nat.  Ed.  13,  p.  3537,  no.  43. 

CARIBIELLA  Perrilliat,  1972:  82 

Type  sp.  (O.  D.):  Murex  intermedius  C.  B. 
Adams,  1850,  Contrib.  to  Conch.  1(4):  60. 

CENTRIFUGA  Grant  and  Gale,  1931:  706-707 
Type  sp.  (O.  D.):  Murex  centrifuga  Hinds,  1844, 
Mollusca,  pi.  1,  p.  8,  pi.  3,  figs.  7,  8. 

CENTRONOTUS  Swainson,  1833:  100  (not 
Centronotus  Schneider  1801) 
Type  sp.  (O.  D.):  Murex  eurystomus  Swainson, 
1833  (=  ?M.  duplex  Roding,  1798),  Zool.  Illust. 
(2)3:  100,  pi.  3. 

Type  sp.  (S.  D.,  ICZN,  1970):  Murex  radix 
Gmelin,  1791,  Syst.  Nat.,  Ed.  13,  p.  3527.  (see 
ICZN  opinion  911,  1970,  Bull.  Zool.  Nomencl. 
27:  20,  wherein  M.  radix  was  designated  as  type 
to  supersede  all  others). 

CERASTOMA  Conrad,  1837:  264 
Type  sp.  (M.):  Murex  (Cerastoma)  nuttalli,  Con- 
rad. 1837:  J.  Acad.  Nat.  Sci.  Philadelphia  7:  264, 
pi.  20,  fig.  22. 

CERATOSTOMA  Hermannsen,  1846:  206 

New  name  for  Cerastoma  Conrad,  1837,  not 
Latreille,  1802. 

CHALMON  de  Gregorio.  1885:  28 

Type  sp.  (O.  D.):  Trophon  (Chalmon)  muricatus 
Montagu,  1802,  Test.  Brit.  1:  262.  pi.  9,  fig.  2. 

CHATHAMIDEA  Dell,  1956:  118 

Type  sp.  (O.  D.):  C.  expeditionis  Dell,  1956, 
Dominion  Mus.  Bull.  18:  118,  figs.  159,  160. 

CHICOMUREX  Arakawa,  1964:  361 

Type  sp.  (O.  D.):  Murex  superbus  Sowerby,  1889, 
Proc.  Zool.  Soc.  London  1889:  565,  pi.  28,  figs. 
10,  11. 

CHICOREUS  Montfort,  1810:  610 

Type  sp.  (fixed  by  ICZN  opin.  911.  1970.  Bull. 
Zool.  Nomencl.  27:  20):  Murex  ramosus  Linne, 
1758,  Syst.  Nat.  Ed.  10,  p.  747,  no.  448. 

CHOREOTYPHIS  Iredale,  1936:  324 

Type  sp.  (O.  D.):  Typhina  pavlova  Iredale,  1936, 
Rec.  Australian  Mus.  19(5):  324,  pi.  24,  fig.  12. 

CHORUS  Gray,  1847b:  136 

Type  sp.  (O.  D.):  Monoceros  giganteus  Lesson, 
1831,  Zool.  11(1):  403. 

CINCLIDOTYPHIS  DuShane,  1969:  343 

Type  sp.  (O.  D.):  C.  myrae  DuShane,  1969,  The 
Veliger,  11(4):  343.  p.  54,  figs.  1-3. 

COLUMBARIUM  von  Martens,  1881:  105 

Type  sp.  (O.  D.):  Pleurotoma  (Columbarium) 
spinicitictum  Martens,  1881;  Conchologische 
Mitteilungen,  2:  105,  pi.  21,  fig.  1-4. 

COLUZEA  Finlay  1927:  407 

Type  sp.  (O.  D.):  Fusus  spiralis  A.  Adams,  1856, 
Proc.  Zool.  Soc.  London  23:  221.  (For  comments 
see  Keen,   A.   M.,    1969,    Bull.   Zool.   Nomencl. 


26:  184.) 

COMPTELLA  Finlay,  1927:  424 

Type  sp.  (O.  D.):  Trophon  curtus  Murdoch,  1905, 
Trans.  New  Zealand  Inst.  37:  228. 

CONCHOLEPAS  "Klein"  Bruguiere.  1792:  535 
Type  sp.   (S.    D.   Lamarck,    1801):    Concholepas 
peruvianus   Lamarck,    1801    (=    Buccinum    con- 
cholepas Bruguiere,  1789),  Syst.  Anim.  s.  Vert, 
p.  70. 

CONCHOPATELLA  Herrmannsen.  1847:  291 
Listed  as  synonym  of  Concholepas  Lamarck,  1801 
(=  Concholepas  Bruguiere,  1792). 

CONCHULUS  Rafinesque.  1815:  142 

Introduced  as  a  synonym  oi  Concholepas  Lamarck, 
1801  (=  Concholepas  Bruguiere,  1792). 

CONOTHAIS  Kuroda,  1930:  1 

Type  sp.  (M.):  Conothais  citrina  Kuroda,  1930, 
Venus,  2(1):  1. 

CORALLINIA  Bucquoy  &   Dautzenberg  (in,   Buc- 
quoy,  Dautzenberg,  &  DoUfus),  1882:  24 
Type  sp.    (O.   D.):   Murex  aciculatus  Lamarck, 
1822,  Hist.  Nat.  Anim.  s.  Vert.  7:  176. 

CORALLIOBIA  H.  &  A.  Adams,  1853:  138 

Type  sp.  (M.):  Leptoconchus  (Coralliobia) 
fimbriata  H.  &  A.  Adams,  1853  (nomen  nudem) 
(=  Concholepas  [Coralliobia]  fimbriata  A. 
Adams,  1854),  Proc.  Zool.  Soc.  London  19:  93. 

CORALLIOFUSUS  Kuroda,  1953:  119 

Type  sp.  (O.  D.):  Coralliofusus  acus  Kuroda, 
1953,  Venus  17:  119,  figs.  3.  4. 

CORALLIOPHILA  H.  &  A.  Adams,  1853:  135 
Type  sp.  (S.  D.  Iredale,  1912):  Murex  neritoideus 
Chemnitz  (non-binominal)  (=  Fusus  neritoideus 
Lamarck,     1816),     Neues     Systematisches     Con- 
chylien-Cabinet  10:  280,  pi.  165,  figs.  1577,  1578. 

CRASPEDOTRITON  Dall,  1904:  119 

Type  sp.   (O.   D.):    Triton   convolutus  Broderip, 

1833,  Proc.  Zool.  Soc.  London  1:  7. 
CRASSILABRUM  Jousseaume.  1880:  335 

Type  sp.  (O.  D.):  Murex  crassilabrum  Sowerby, 

1834.  Conch.  Illust.  Murex.  pi.  59,  fig.  14. 
CRONIA  H.  &  A.  Adams,  1853:  128 

Type  sp.  (M.):  Purpura  amygdala  Kiener,  1835, 
Spec.  Gen.  Icon.  Coq.  Viv.  .  .  .  Pourpre,  p.  39, 
pi.  10,  fig.  26. 

CUMA  Swainson,  1840:  87,  307 

Type  sp.  (O.  D.):  Buccinum  tectum  Wood,  1828, 
Index  Testaceologicus  ...  A  catalog  of  shells, 
Suppl.  p.  12,  no.  13,  pi.  4,  fig.  13. 

CUMOPSIS  Rovereto,  1899:  105 

New  name  for  Cuma  Swainson,  1840,  not  Milne- 
Edwards,  1828-see  Cymia  Morch,  1860. 

CYTHAROMORULA  Kuroda,  1953:  183 

Type  sp.  (M.):  Cytharomorula  vexillum  Kuroda, 
1953,  Venus  17:  183. 

DALLIMUREX  Rehder,  1946:  142 

Type  sp.  (O.  D.):  Murex  nuttingi  Dall,  1896  (  = 
Murex  paz/ Crosse,  1869),  Bull.  Lab.  Nat.  Hist., 
State  Univ.  Iowa  4(1):  13,  pi.  1,  fig.  1. 

DENTOCENEBRA  Monterosato,  1917:  21 

Type  sp.  (O.  D.):  Ocenebra  corallinus  Scacchi, 
1836  (=  O.  aciculata  Lamarck,  1822),  Cat. 
Conch.  Regni  Neap.,  p.  11,  fig.  15. 

DERMOMUREX  Monterosato,  1890:  181 

New  name  for  Poweria  Monterosato,  1884,  not 
Bonaparte,  1841. 

DICATHAIS  Iredale,  1936:  325 

Type  sp.  (O.  D.):  Buccinum  orbita  Gmelin,  1791, 
Syst.  Nat.  Ed.  13,  Vermes,  p.  3490.  no.  183. 


282 


DISTICHOTYPHIS  Keen  &  Campbell,  1964:  56 
Type  sp.  (O.  D.):  Distichotvphis  vemae  Keen  & 
Campbell.  1964,  The  Veliger  7(1):  56-57,  pi.  11. 
figs.  45-47. 

DRUPA  Roding,  1798:  55 
Type  sp.  (S.  D.  Rovereto,  1899):  Drupa  morun, 
Roding,  1798,  Museum  Boltenianum  p.  55,  no. 
694. 

DRUPELLA  Thiele.  1925:  137 

Type  sp.  (S.  D.  Wenz,  1941):  Drupa  (Drupella) 
ochrostoma  (Blainville,  1832),  Nouv.  Ann.  Mus. 
Hist.  Nat.  Paris  ser.  3,  1:  205. 

DRUPINA  Dall,  1923:  303 

Type  sp.  (O.  D.):  Ricinula  digitata  Lamarck, 
1816  (=  Drupa  grossularia  Roding,  1798), 
Tableau  Encycl.  Meth.  pi.  395,  fig.  7a,  7b,  Liste, 
p.  2. 

tECPHORA  Conrad.  1843:  310 

Type  sp.  (O.  D.):  Fusus  quadricostatus  Say,  1824, 
J.  Acad.  Nat.  Sci.  Philadelphia  4:  127. 

EMOZAMIA  Iredale,  1929a:  185 

Type  sp.  (O.  D.):  Murex  licinus  Hedley  and  Pet- 
terd,  1906,  Rec.  Australian  Mus.  6:  219,  pi.  37, 
fig.  6. 

ENATIMENE  Iredale,  1929a:  185 

Type  sp.  (O.  D.):  Trophon  simplex  Hedley,  1903, 
Mem.  Australian  Mus.  4(1):  380. 

ENIXOTROPHON  Iredale,  1929a:  185 

Type  sp.  (O.  D.):  Trophon  carduelis  Watson, 
1882,  MoUusca  of  H.  M.  S.  Challenger  Expedi- 
tion, pt.  14.  p.  388. 

tENTACANTHUS  Ihering,  1907:  183 

Type  sp.  (M.):  Trophon  monoceros  Ihering,  1907, 
Anal.  Mus.  Nac.  Buenos  Aires  14:  183. 

tEOTYPHIS  Tembrock,  1963:  322 

Type  sp.  (O.  D.):  Typhis  sejunctus  Semper,  1861, 
Arch.  Verens.  Freunde  Naturg.  Mecklen.  15:  161. 

ERGALATAX  Iredale,  1931:  231 

Type  sp.  (O.  D.):  Ergalatax  recurrens  Iredale, 
1931  (=  Buccinum  contractum  Reeve,  1846), 
Rec.  Australian  Mus.  18:  231. 

EUPHYLLON  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Murex  monodon  Sowerby,  1825 
(=  Murex  comucervi  Roding,  1798),  Cat.  Shells 
Tankerville,  App.,  p.  19,  sp.  1703. 

EVOKESIA  Radwin  &  D'Attilio,  1972:  335 

Type  sp.  (O.  D.):  Sistrum  rufonotatum  Carpenter, 
1864,  Ann.  Mag.  Nat.  Hist.",  ser.  3,  14:. 48. 

EUPLEURA  H.  &  A.  Adams,  1853:  107 

Type  sp.  (S.  D.,  F.  C.  Baker,  1895):  Ranella 
caudata  Say,  1822,  J.  Acad.  Nat.  Sci.  Philadelphia 
2:  236. 

FAVARTIA  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Murex  breviculus  Sowerby, 
1834,  Conch.  Illust.,  pi.  63,  fig.  37. 

tFLEXOPTERON  Shuto,  1969:  112 

Type  sp.  (O.  D.):  Flexopteron  philippinensis 
Shuto,  1969,  Mem.  Fac.  Sci.  Kyushu  Univ.  (ser. 
Geol.)  19:  112. 

FORRERIA  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Murex  belcheri  Hinds  1844, 
Proc.  Zool.  Soc.  London  11:  128. 

FRONDOSARIA  Schluter,  1838:  20 
Type  sp.  (S.  D.,  E.  H.  Yokes,  1964):  Frondosaria 
inflata    (Lamarck,     1822)    (=     Murex    ramosus 
Linne,  1758),  Hist.  Nat.  Anim.  s.  Vert.  7:  160. 

FUEGOTROPHON  Powell,  1951:  157 

Type  sp.  (O.  D.):  Fusus  crispus  Gould,  1849  (  = 
Murex  pallidus    Broderip,    1833),    Proc.    Boston 


Soc.  Nat.  Hist.  3:  141. 
tFULGUROFUSUS  Grabau,  1904:  86 

Type  sp.  (O.  D.):  Fusus  quercollis  Harris,   1896, 
Bull.  Amer.  Paleo.  1:  200,  pi.  18,  fig.  9. 
FUSOMUREX  Coen,  1922:  69 
Type  sp.  (O.  D.):  Purpura  alucoides  Blainville, 
1829,  Faune  Francaise,  p.  128,  pi.  5b,  fig.  1. 

tGALEROPSIS  Hupe,  1860:  127 

Type  sp.  (O.  D.):  Galeropsis  lavenavana  Hupe, 
1860,  Rev.  Mag.  Zool.  12:  127. 

GALFRIDUS  Iredale,  1924:  271 

Type  sp.  (O.  D.):  Triton  (Cumia)  speciosum 
Angas,  1871,  Proc.  Zool.  Soc.  London,  89:  13,  pi. 
1,  fig.  1. 

GEMIXYSTUS  Iredale,  1929a:  185 

Type  sp.  (O.  D.):  Trophon  laminatus  Petterd, 
1884,  J.  Conch.,  4:  136,  pi.  22,  fig.  3. 

GENKAIMUREX  Kuroda,  1953:  120 

Type  sp.  (O.  D.):  Coralliophila  (Genkaimurex) 
varicosa  Kuroda,  1953  (=  Murex  fimbriatulum 
A.  Adams,  1863).  Venus  17:  120. 

GRACILIMUREX  Thiele,  1929:  289 

Type  sp.  (O.  D.):  Gracilimurex  bicolor  Thiele, 
1929,  Handbuch  Syst.  Weichtier.,  p.  289. 

GRACILIPURPURA  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Fusus  strigosus  Lamarck,  1822, 
Hist.  Nat.  Anim.  s.  Vert.  7:  130. 

HADRIANIA  Bucquoy  &  Dautzenberg,  1882:  16,  33 
Type  sp.  (O.  D.):  Murex  craticulatus  Brocchi, 
1814  (not  Linne,  1758)  (=  Hadriania  craticuloides 
E.  H.  Vokes,  1964),  Conch.  Foss.  Subapp.,  2: 
406,  pi.  7,  fig.  14. 

HANETIA  Jousseaume,  1880:  335 
Type  sp.  (O.  D.):  Murex  haneti  Petit,   1856,  J. 
Conchyl.  5:  90. 

tHARMATIA  Noszky,  1940:  28 

Type  sp.  (O.  D.):  Murex  (Harmatia)  stephani 
Noszky,  1940,  Ann.  Hist.  Nat.  Mus.  Hung.  Min. 
Geol.  33:  1-80. 

HAUSTELLARIA  Swainson,  1833:  pi.  100 

Type  sp.  (O.  D.):  Haustellaria  haustellum  Linne, 
1758,  Syst.  Nat.,  Ed.  10,  p.  746,  no.  493. 

HAUSTELLOTYPHIS  Jousseaume,  1880:  335 
Type  sp.  (O.  D.):  Typhis  cumingi  Broderip,  1833, 
Proc.  Comm.  Sci.  Corr.  Zool.  Soc.  London  2:  177. 

HAUSTELLUM  "Klein"  Bruguiere,  1792:  533 
Type   sp.    (S.    M.,    Schumacher,    1817),    Murex 
haustellum  Linne,  1758,  p.  746,  no.  213. 

HAUSTRUM  Perry,  1811:  pi.  44 
Type    sp.     (S.     D.     Iredale,     1915):     Buccinum 
haustrum    Martyn,     1788    (non-binominal)     (  = 
Buccinum    haustorium    Gmelin,     1791),     Univ. 
Conch.,  vol.  2,  fig.  9c. 

HERTLEINELLA  Berry,  1958:  95 

Type  sp.  (O.  D.):  Hertleinella  leucostephes  Berry, 
1958,  1(16):  95. 

tHETEROPURPURA  Jousseaume,  1880:  335 
Type  sp.  (O.  D.):  Murex  polymorphus  Brocchi, 
1814,  Conch.  Foss.  Subapp.  2:  415,  pi.  8,  figs. 
4a,  4b. 

tHEXACHORDA  Cossmann,  1903:  47 

Type  sp.  (O.  D.):  Murex  tenellus  Mayer-Eymar, 
1869,  J.  Conchyl.  17:  82,  pi.  3  fig.  5. 

HEXAPLEX  Perry,  1811:  pi.  8 

Type  sp.  (S.  D.  Jousseaume,  1880):  Murex 
cichoreum  Gmelin,  1791,  Syst.  Nat.,  Ed.  13,  1: 
3530. 

tHIPPOCAMPOIDES  Wade,  1916:  466 
Type    sp.     (O.     D.):    Hippocampoides    serratus 


283 


Wade,  1916.  Proc.  Acad.  Nat.  Sci.  Philadelphia 
68:  466.  pi.  24,  figs.  11-13. 

HIRTOMUREX  Coen.  1922:  69 

Type  sp.  (O.  D.):  Fusus  lamellosa  Philippi,  1836, 
Enum.  Moll.  Sicil.,  p.  204. 

tHIRTOTYPHIS  Jousseaume,  1880:  336 

Type  sp.  (O.  D.):  Murex  horridus  Brocchi,  1814, 
Conch.  Foss.  Subapp.,  2:  405,  pi.  7,  fig.  17. 

tHISPIDOFUSUS  Darragh,  1969:  67 

Type  sp.  (O.  D.):  Fusus  senticosus  Tate,  1888, 
Trans.  Roy.  Soc.  S.  Austr.,  10:  135,  pi.  7,  fig.  3. 

HISTRICOSCEPTRUM  Darragh,  1969:  87 

Type  sp.  (O.  D.):  Columbarium  atlantis  Clench  & 
Aguayo,  1938,  Mem.  Soc.  Cuba  Hist.  Nat.  vol. 
12(5):  382,  pi.  28,  fig.  1. 

HOMALOCANTHA  Morch,  1852:  95 

Type  sp.  (M.):  Murex  scorpio  Linne,  1758,  Syst. 
Nat.  Ed.  10,  p.  747,  no.  449. 

tINDOTYPHIS  Keen,  1944:  59 

Type  sp.  (O.  D.):  Laevityphis  (Indotyphis) 
hantamensis  (Oostingh,  1933),  De  Mijningenieur, 
Jaarg.  14,  p.  193. 

INERMICOSTA  Jousseaume,  1880:  335 
Type  sp.  (O.  D.):  Murex  fasciat us  Sowerby,  1841, 
Proc.  Zool.  Soc.  London  8:  144. 

lOPAS  H.  &  A.  Adams,  1853:  128 

Type  sp.  (S.  D.  Dall,  1909):  Purpura  sertum 
Lamarck,  1816  (=  Buccinum  sertum  Bruguiere, 
1789),  Tableau  Encycl.  Meth..  pi.  397,  fig.  2, 
Liste  p.  2. 

JANIA  Cossmann,  1892:  68 

Type  sp.  (O.  D.):  Murex  blainvillei  Payraudeau, 
1826,  Cat.  Moll.  Corse,  p.  149. 

JATON  Pusch,  1837:  135 

Type  sp.  (O.  D.):  Murex  decussatus  Gmelin, 
1791,  Syst.  Nat.  Ed.  13,  p.  3527,  no.  7. 

JATOVA  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Purpura  jatou  Adanson,  1757 
(non-binominal)  (=  Murex  decussatus  Gmelin, 
1791),  Hist.  Nat.  du  Senegal.  Coquillages,  p.  129, 
pi.  9,  fig.  21. 

KALYDON  Hutton,  1884:  220  (not  to  be  confused 

with    Calydon    J.    Thomson,    1864)    (Kalydon 

suppressed  —  ICZN   Opin.    911,    1970,   Bull. 

Zool.  Nomencl.  27:  20). 

Type  sp.  (O.  D.):  Fusus  plebeius  Hutton,   1873, 

Cat.  Mar.  Moll.  New  Zealand  p.  9. 

LAEVITYPHIS  Cossmann,  1903:  59 

Type  sp.  (O.  D.):  Typhis  coronarius  Deshayes, 
1865  (=  Typhis  muticus  J.  Sowerby,  1834), 
Descr.  Anim.  s.  Vert,  decouv.  Bassin  de  Paris,  p. 
335,  pi.  88,  figs.  11-13. 

LAMELLATIAXIS  Habe  &  Kosuge,  1970:  182 
Type     sp.     (O.     D.):     Latiaxis     (Lamellatiaxis) 
marumai  Habe  &  Kosuge,  1970,  Venus  24(4):  182. 

LATAXIENA  Jousseaume,  1883:  187 

Type  sp.  (T.):  Lataxiena  lataxiena  Jousseaume, 
1883  (==  Trophon  fimbriatus  Hinds,  1844),  Bull. 
Soc.  Zool.  Franc.  8:  187. 

LATIAXIS  Swainson,  1840:  82,  306 

Type  sp.  (M.):  Pyrula  mawae  "Gray"  Griffith  & 
Pidgeon,  1834,  (in,  Cuvier,  Regne  Animal) 
Mollusca  &  Radiata,  p.  599,  pi.  25,  figs.  3,  4. 

LATIMUREX  Coen,  1922:  70 

Type  sp.  (O.  D.):  Murex  meyendorffi  Calcara, 
1845,  Cenne  Moll.  Sicil.,  p.  38. 

LENITROPHON  Finlay,  1927:  424 
Type  sp.  (O.  D.):  Trophon  convexus  Suter,  1909, 
Rec.  Canterbury  Mus.  1(2):  126,  pi.  12,  fig.  4. 


LEPADOMUREX  Coen,  1922:  70 

Type  sp.  (O.  D.):  Purpura  brevis  Blainville,  1832, 
Nouv.  Ann.  Mus.  Hist.  Nat.  Paris,  vol.  1(2):  233. 

LEPSIA  Hutton,  1884:  223 

Type  sp.  (O.  D.):  Purpura  haustrum  Martyn, 
1788  {—  Buccinum  haustorium  Gmelin,  1791), 
Univ.  Conch.,  vol.  2,  fig.  9c. 

LEPSIELLA  Iredale,  1912:  223 

Type  sp.  (O.  D.):  Purpura  scobina  Quoy  & 
Gaimard,  1833,  Voyage  of  the  "Astrolabe,"  Zool. 
II,  p.  567. 

LEPSITHAIS  Finaly,  1928:  258 

Type  sp.  (O.  D.):  Polvtropa  squamata  Hutton, 
1878.  J.  Conchyl.  26:  19. 

LEPTOCONCHUS  Ruppell,  1834:  105 

Type  sp.  (S.  D.  Gray,  1847):  Leptoconchus  peroni 
Lamarck,  1818,  Hist.  Nat.  Anim.  s.  Vert.  5:  374. 

LINIAXIS  Laseron,  1955:  72 
Type   sp.    (O.    D.):    Liniaxis    elongata    Laseron, 
1955,   Proc.   Roy.   Zool.   Soc.   New  South  Wales 
1953-54:  72. 

LITOZAMIA  Iredale,  1929a:  185 

Type  sp.  (O.  D.):  Peristemia  rudolphi  Henn  & 
Brazier,  1894,  Proc.  Linn.  Soc.  New  South  Wales 
19:  166,  pi.  14,  fig.  1. 

tLOWENSTAMIA  Sohl,  1964a:  182 

Type  sp.  (O.  D.):  Lowenstamia  funiculus  Sohl, 
1964,  U.S.  Geol.  Surv.  Prof.  Paper  331b:  182,  pi. 
21,  figs.  23,  26. 

tLYROPURPURA  Jousseaume,  1880:  335 
Type  sp.  (O.  D.):  Murex  crassicostata  (Deshayes, 
1835,  Descrip.  Coq.  Foss.  Environ,  de  Paris  2: 
601,  pi.  82,  figs.  13,  14. 

tLYROTHYPHIS  Jousseaume,  1880:  336 
Type  sp.  (O.  D.):   Typhis  cuniculosus  Duchatel, 
(in   Bronn),    1848   (=   Murex  cuniculosus  Nyst, 
1836).  Mess.  Sci.  Arts  Belg.  4:  176. 

MACULOTRITON  Dall,  1904:  136 

Type  sp.  (O.  D.):  Triton  bracteata  Hinds,  1844, 
Proc.  Zool.  Soc.  London,  12:  132. 

MAGILOPSIS  Sowerby,  1919:  77 

Type  sp.  (O.  D.):  Leptoconchus  lamarcki  Des- 
hayes, 1863,  Cat.  Moll.  Conchyl.  L'ile  de  la 
Reunion  (Bourbon),  p.  127,  pi.  12,  figs.  1-3. 

MAGILUS  Montfort,  1810:  42 

Type  sp.  (O.  D.):  Magilus  antiquus  Montfort, 
1810,  2:  42. 

MANCINELLA  Link,  1807:  115 

Type  sp.  (T.):  Murex  mancinella  Linne,  1758  (  = 
Mancinella  aculeata  Link,  1807),  Syst.  Nat.  Ed. 
10,  p.  751,  no.  469. 

MARCHIA  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Murex  clavus  Kiener,  1843  (  = 
Murex  elongata  Lightfoot,  1786),  Spec.  Gen. 
Icon.  Coq.  Viv.  7:  111,  pi.  37,  fig.  2. 

MAXWELLIA  Baily,  1950:  9 

Type  sp.  (O.  D.):  Murex  gemma  Sowerby,  1879, 
Thes.  Conch.,  vol.  4,  Murex,  p.  32,  fig.  214. 

MENATHAIS  Iredale,  1937:  256 
Type  sp.  (O.  D.):  Purpura  pica  Blainville,   1832, 
Nouv.  Ann.  Mus.  Hist.  Nat.  Paris,  ser.  3,  1:  213. 

tMICRORHYTIS  Emerson,  1959:  6 

Type  sp.  (O.  D.):  Pterorytis  (Microrhytis)  pecki 
Emerson,  1959,  Amer.  Mus.  Novitates  1974. 

MICROTOMA  Swainson,  1840:  301 

Type  sp.  (S.  D.  Gray,  1847  [as  Microstoma]): 
Microtoma  persica  (Lamarck,  1799),  Mem.  Soc. 
Hist.  Nat.  Paris,  p.  71. 

MINNIMUREX  Woolacott,  1957:  115 


284 


Type  sp.  (O.  D.):  Minnimurex  phantom  Woola- 
cott,  1957,  Proc.  Roy.  Soc.  New  South  Wales, 
1955-56.  p.  115. 

MINORTROPHON  Finlay,  1927:  425 

Type  sp.  (O.  D.):  Daphnella  crassilirata  Suter, 
1908,  Trans.  Proc.  New  Zealand  Inst.  57:  425. 

tMIOCENEBRA  E.  H.  Yokes,  1963:  162 

Type  sp.  (O.  D.):  Tritonalia  (Miocenebra) 
silverdalense  E.  H.  Yokes,  1963,  Tulane  Stud. 
Geol.  1(4):  162,  pi.  2,  figs.  6a,  6b,  7a,  7b. 

MIPUS  De  Gregorio,  1885:  28 

Type  sp.  (O.  D.):  Trophon  gyratum  Hinds,  1844, 
Yoy.  H.  M.  S.  Sulphur,  Zoology,  2:  14,  pi.  1,  figs. 
14-15. 

MONOCEROS  Lamarck,  1822:  250  (not  Bloch  & 
Schneider,  1801) 
Type  sp.  (S.  D.,  herein):  Monoceros  imbricatum 
Lamarck,   1816  (=   Buccinum  monodon  Pallas, 
1774),  Tabl.  Encycl.  Meth.,  pi.  396,  figs,  la,  lb, 
Liste.  p.  2. 

MONSTROTYPHIS  Habe,  1961:  19  (appendix) 
Type  sp.  (O.  D.):   Typhis  (Typhinellus)  tosaensis 
Azuma,      1960,     Cat.      Shell-bearing     Mollusca 
Okinoshima,    Kashiwajima  .  .  .  (Tosa    Province), 
Shikoku,  Japan.,  p.  99,  pi.  2,  fig.  8. 

tMOREA  Conrad,  1860:  290 

Type  sp.  (M.):  Morea  cancellaria  Conrad,  1860, 
J.  Acad.  Nat.  Sci.,  Philadelphia  4:  290. 

MORULA  Schumacher,  1817:  68,  227 

Type  sp.  (M.):  Morula  papillosa  Schumacher, 
1817  (=  Drupa  uva  Roding,  1798),  Ess.  Yers. 
Test.,  pp.  68,  227. 

MORULINA  Dall,  1923:  303  (not  Borner,  1906) 
Type  sp.  (O.  D.):  Ricinula  mutica  Lamarck,  1816, 
Tabl.  Encycl. 

Tabl.  Encycl.  Meth.,  pi.  395,  figs.  2a,  2b,  Liste,  p. 
1. 

MORUNELLA  Emerson  &  Hertlein,  1964:  361 
Type    sp.    (O.    D.):    Buccinum    lugubre    C.    B. 
Adams,  1852,  Cat.  Shells  coll.  Panama  .  .  .,  p.  69. 

MUREX  Linne,  1758:  746 

Type  sp.  (S.  D.,  Montfort,  1810):  Murex  pecten 
Montfort,  1810,  (not  Lightfoot,  1786),  (=  Murex 
tribulus  Linne,  1758),  Conch.  Syst.  2:  619. 

MUREXIELLA  Clench  &  Perez  Farfante,  1945:  49 
Type  sp.  (O.  D.):  Murex  hidalgoi  Crosse,  1869,  J. 
Conchyl.  17:  408. 

MUREXSUL  Iredale,  1915:  471 

Type  sp.  (O.  D.):  Murex  octogonus  Quoy  & 
Gaimard,  1832,  Yoyage  .  .  .  I'Astrolabe  .  .  .  Paris, 
Zool.,  Mollusca,  2:  531,  pi.  36,  figs.  8,  9. 

MURICANTHUS  Swainson,  1840:  296 

New  name  for  Centronotus  Swainson,  1833,  not 
Schneider,  1801. 

MURICIDEA  Swainson,  1840:  64 

Type  sp.  (O.  D.):  Murex  magellanicus  Lamarck, 
1816  (—  Buccinum  geversianum  Pallas,  1774), 
Tabl.  Encycl.  Meth.,  pi.  419,  figs.  4a,  4b,  Liste, 
p.  5. 

MURICODRUPA  Iredale,  1918:  38 

Type  sp.  (O.  D.):  Purpura  fenestrata  Blainville, 
1832  (=  Murex  fvniculus  Wood,  1828),  Nouv. 
Ann.  Mus.  Hist.  Nat.  Paris  1:  221. 

MURICOPSIS  Bucquoy  &  Dautzenberg,  1882: 
16,  19 
Type  sp.  (O.  D.):  Murex  blainvillei  Payraudeau, 
1826,   Cat.    Descr.    Meth.    Annel.    Moll.    lie   de 
Corse,  p.  149. 

MURITHAIS  Grant  &  Gale,  1931:  729 


Type  sp.  (O.  D.):  Murex  trunculus  Linne,   1758, 
Syst.  Nat.  Ed.  10,  p.  747,  no.  447. 
tMUROTRITON  de  Gregorio,  1890:  97 

Type  sp.  (O.  D.):   Triton  grassator  de  Gregorio, 
1890,  Ann.  Geol.  Paleo.  7:  97. 
NAQUETIA  Jousseaume,  1880:  335 

Type  sp.   (O.   D.):  Murex  triqueter  Born,    1778, 
Index  Mus.  Caes.  Yindobon.,  p.  288. 
NAMAMUREX  Carrington  &  Kensley,  1969:  197 

Type    sp.     (O.     D.):    Namamurex    odontostoma 

Carrington  &  Kensley,   1969,  Ann.  S.  Afr.  Mus. 

52:  197. 
NASSA  Roding,  1798:  132 

Type  sp.  (S.  D.,  Dall  1909):  Nassa  picta  Roding, 
1798    (=    Buccinum    sertum    Bruguiere,     1789), 
Museum  Boltenianum,  p.  132. 
NEMOFUSUS  Cossmann,  1903:  195 

Type  sp.  (O.  D.):  Murex  fusulus  Brocchi,   1814, 
Conch.  Foss.  Subapp.  2:  409,  pi.  8,  fig.  9. 
NEORAPANA  Cooke,  1918:  7,  11 

Type  sp.    (O.    D.):   Purpura   muricata   Broderip, 

1832,  Proc.  Comm.  Sci.  Zool.  Soc.   London,   2: 

125,  126. 
NEOTHIAS  Iredale,  1912:  223 

Type  sp.  (O.  D.):  Purpura  smithi  Brazier,   1889, 

Mem.  Australian  Mus.  2:  pi.  4,  figs.   1-4,  7-12, 

21.  22. 
tNEOTYPHlS  Yella,  1961:  385 

Type  sp.  (O.  D.):  Typhis  tepunga  Fleming,  1943, 

Trans.  Roy.  Soc.  New  Zealand  73(3):  205,  pi.  30, 

fig.  21. 
tNEURARHYTIS  Olsson  &  Harbison,  1953:  252 

Type  sp.  (O.  D.):  Purpura  (Pterorhytis)  fluviana 

(Dall,  1903),  Trans.  Wagner  Free  Inst.  Sci.  3(6): 

1633,  pi.  60,  figs.  20,  21. 
NIPPONOTROPHON  Kuroda  &  Habe,  1971:  233 
(Japanese),  152  (English) 

Type  sp.    (O.   D.):   Boreotrophon   echinus   Dall, 

1920.  Proc.  U.S.  Natl.  Mus.  54:  232. 
NODULOTROPHON  Habe  &  Ito,  1965:  32-33 

Typesp.  (O.  D.):  Trophon  dalli.  Kobelt,  1878  (in 

Kiister)  Martini  &  Chemnitz  Conchylien-Cabinet, 

pt.  275,  p.  289,  pi.  74,  figs.  1,  2. 
NOTHOTYPHIS  Fleming,  1962:  109,  119 

Type    sp.    (O.D.):    Pterynotus    (Nothotyphis) 

norfolkensis  Fleming,  1962,  Trans.  Roy.  Soc.  New 

Zealand  2(14):  109,  119. 
NUCELLA  Roding,  1798:  131 

Type   sp.    (S.    D.    Winckworth,    1945):    Nucella 

theobromus  Roding,  1798  (=  Buccinum  lapillus 

Linne.  1758),  Mus.  Bolten.  p.  131. 
OCINEBRELLUS  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Murex  eurypteron  Reeve,  1845, 

Conch.  Icon.,  vol.  3,  Murex,  sp.  176,  pi.  34,  fig. 

176a,  176b. 
OCENEBRA  Gray,  1847a:  269 

Type  sp.    (M.):   Murex   erinaceus   Linne,    1758, 

Syst.  Nat.  Ed.  10,  p.  748,  no.  451. 
OCINEBRINA  Jousseaum,  1880:  335 

Type  sp.  (O.  D.):  Fusus  corallinus  Scacchi,  1836 

(=    Murex    aciculatus    Lamarck,     1822),     Cat. 

Conch.  Neap.,  p.  11. 
OLLAPHON  Iredale,  1929a:  186 

Type  sp.  (O.  D.):  Trophon  molorthus  Hedley  & 

May,  1908,  Rec.  Australian  Mus.  7:  122. 
OPPOMORUS  Iredale,  1937:  258 

Type    sp.    (O.    D.):    Morula    nodulifera    Menke, 

1829,  Conch.  Samml.  Malsburg,  p.  33. 
ORANIA  Pallary,  1900:  285 


285 


Type  sp.  (O.  D.):  Pseudomurex  spadae  Libassi, 
1859,  Atti  Acad.  Palermo,  3:  43,  fig.  29. 

PAGODULA  Monterosato,  1884:  116 

Type  sp.  (O.  D.):  Murex  vaginata  Cristofori  & 
Jan,  1832,  Cat.  .  .  .  rerum  Nat.  Mus.  Extant. 
Josephi  de  Cristofori,  Sect.  11(1),  Conch.  Fossili, 
p.  11. 

tPANAMUREX  Woodring,  1959:  217 

Type  sp.  (O.  D.):  Murex  gatunensis  Brown  & 
Pilsbry,  1911,  Proc.  Acad.  Nat.  Sci.  Philadelphia 
63:  354,  pi.  26,  fig.  2. 

PARATROPHON  Finlay,  1927:  424 

Type  sp.  (O.  D.):  Polvtropa  cheesemani  Hutton, 
1882,  New  Zealand  J.  Sci.  1:  69. 

PASCULA  Dall.  1908:  311,  312 

Type  sp.  (O.  D.):  Trophon  (Pascula)  citricus  Dall, 
1908.  Bull.  Mus.  Comp.  Zool.  43(6):  312. 

PATELLIPURPURA  Dall,  1909:  50 
Type  sp.  (O.  D.):  Buccinum  patulum  Linne,  1758, 
Syst.  Nat.,  Ed.  10,  p.  739,  no.  402. 

PAZIELLA  Jousseaume,  1880:  335 
Type  sp.  (O.  D.):  Murex  pazi  Crosse,    1869,  J. 
Conchyl.,  17:  183. 

PAZINOTUS  E.  H.  Yokes,  1970b:  27 
Type  sp.  (O.  D.):  Eupleura  stimpsoni  Dall,  1889, 
Bull.  Mus.  Comp.  Zool.  18:  204. 

PENTADACTYLUS  "Klein"  Bruguiere,  1792:  520 
(not  Schultze,  1760) 
Type  sp.  (S.  D.,  P.  C.  Baker,  1895):  Pentadactylus 
ricinus  Lamarck  (=  Murex  ricinus  Linne,   1758, 
Syst.  Nat.,  Ed.  10,  p.  750,  no.  464.) 

tPERITROPHON  Marwick,  1931:  119 

Type  sp.  (O.D.):  Peritrophon  decoratus  Marwick, 
1931,  Paleont.  Bull.  New  Zealand  no.  13:  119. 

PEROTYPHIS  Jousseaume,  1880:  336  (error  for 
Pterotyphis  —  q.v.) 
Type  sp.  (O.D.):  Typhis  pinnatus  Broderip,  1833, 
Proc.  Comm.  Sci.  Corr.  Zool.  Soc.  London  2:  178. 

PHYLLOCOMA  Tapparone-Canefri.  1881:  44 
Type  sp.   (O.   D.):    Triton   convolutus  Broderip, 
1833,  Proc.  Zool.  Soc.  London  1:  7. 

PHYLLONOTUS  Swainson,  1833:  pi.  100 
Type  sp.  (S.  D.  Swainson,  1833  —  pi.  109): 
Murex  imperialis  (var.a)  Swainson,  1833  (  = 
Murex  imperialis.  Swainson,  1831)  (not  Fischer 
de  Waldheim,  1807),  (=  Murex  margaritensis 
Abbott,  1958),  Zool.  Illust.,  ser.  2,  3:  pi.  100. 

PHRYGIOMUREX  Dall,  1904:  137 
Type  sp.  (O.  D.):  Triton  sculptilis  Reeve,   1844, 
Proc.  Zool.  Soc.  London  12:  118-119. 

tPILSBRYTYPHIS  Woodring,  1959:  220 

Type  sp.  (O.  D.):  Typhis  gabbi  Brown  &  Pilsbry, 
1911,  Proc.  Acad.  Nat.  Sci.  Philadelphia  63:  354, 
pi.  26,  fig.  6. 

PINAXIA  H.  &  A.  Adams,  1853:  132 
Type  sp.  (M.):  Pinaxia  coronata  H.  &  A.  Adams, 
1853   (nomen   nudem)    (=    Pinaxia   coronata   A. 
Adams,  1853,  Proc.  Zool.  Soc.  London  19:  185). 

PINON  de  Gregorio,  1885:  28 

Type  sp.  (O.  D.):  Trophon  (Pinon)  vaginatus 
Cristofori  &  Jan,  1832,  Cat.  rerum  Nat.  Mus. 
Sect.  11(1),  Conch.  Foss.,  p.  11. 

tPIRGOS  de  Gregorio,  1885:  28 

Type  sp.  (S.  D.,  Cossmann,  1904):  Fusus 
alveolatus  J.  Sowerby,  1823,  Min.  Conch.  5:  9. 

PIRTUS  de  Gregorio,  1884:  257 

Type  sp.  (O.  D.):  Murex  (Pirtus)  fiatus  de 
Gregorio,  1884,  Bull.  Soc.  Malac.  Ital.  10:  257. 

PLANITHAIS  "Bayle"  Fischer,  1884:  645 


Type  sp.  (O.  D.):  Purpura  planospira  Lamarck, 
1822,  Hist.  Nat.  Anim.  s.  Vert.  7:  240. 

PLICOPURPURA  Cossmann,  1903:  69 

New  name  for  Purpurella  Daii,  1871,  not 
Robineau-Desvoidy,  1853. 

POIRIERIA  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Murex  zelandicus  Quoy  & 
Gaimard,  1833,  Voy.  Astrolabe,  Zool.  2:  529,  pi. 
36.  figs.  5-7. 

POLYPLEX  Perry,  1811:  pi.  9 
Type  sp.  (S.  D.,  ICZN  Opinion  911,  1969  —  see 
Bull.  Zool.  Nomencl.,  27:  20):  Polyplex  bulbosa 
Perry,  1811,  Conchology,  pi.  9. 

POLYTROPA  Swainson,  1840:  305 

Type  sp.  (S.  D.,  Gray,  1847):  Buccinum  lapillus 
Linne,  1758.  Syst.  Nat.,  Ed.  10,  p.  739,  no.  403. 

POROPTERON  Jousseaume,  1880:  335 
Type  sp.  (O.  D.):  Murex  tubifer  Bruguiere,  1972, 
(an  apparent  error  as  Jousseaume  also  designated 
this  species  as  type  of  Typhis  Montfort) 
Type  sp.  (S.  D.  Jousseaume,  1881):  Murex 
uncinarius  Lamarck,  1822,  Hist.  Nat.  Anim.  s. 
Vert.  7:  166. 

POWERIA  Monterosato,  1884:  113  (not 
Bonaparte,  1841) 
Type  sp.  (M.):  Poweria  scalarina  Bivona,  1832  (  — 
Murex  scalaroides  Blainville,  1826),  Effem.  Lett. 
Sicii.,  p.  22. 

PROTOTYPHIS  Ponder,  1972:  221 

Type  sp.  (O.  D.):  Typhis  angasi  Crosse,  1863,  J 
Conchyl.  11:  86,  pi.  1,  fig.  2. 

PROVEXILLUM  Hedley.  1918:  79 

New  name  for  Vexilla  Swainson,  1840,  not  Vexil- 
lum  Roding,  1798. 

tPSEUDOMOREA  Cossmann,  1925:  265 
Type  sp.  (M.):  Morea  marylandica  Gardner,  1916, 
Maryland  Geol.  Survey,  Upper  Cretaceous,  Syst. 
Paleo.  Mollusca,  p.  371. 

PSEUDOMUREX  Monterosato,  1872:  15,  33 

Type  sp.  (O.  D.):  Murex  bracteata  Brocchi,  1814, 
Moll.  Foss.  Subapp.,  p.  409,  pi.  9,  fig.  3. 

tPSEUDORAPA  Holzapfel,  1888:  111 

Type  sp.  (O.  D.):  Murex  pleurotomoides  Muller, 
1851,  vol.  1,  Mon.  2,  p.  24,  pi.  3,  fig.  31. 

PSEUDOSALPINX  Olsson  &  Harbison,  et  al., 
1953:  254 
Type  sp.  (O.  D.):  Urosalpinx  floridana  (Conrad, 
1837)   (=   Murex  ostrearum  Conrad,    1846),    J. 
Acad.  Nat.  Sci.  Philadelphia  7:  265. 

PTEROCHELUS  Jousseaume,  1880:  335 
Type  sp.  (O.  D.):  Murex  acanthopterus  Lamarck, 
1816,  Tabl.  Encycl.  Meth.  pi.  417,  figs.  2a,  2b, 
Liste,  p.  5. 

PTEROPURPURA  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Murex  macropterus  Deshayes, 
1839,  Rev.  Zool.  Soc.  Cuv.  2:  360. 

PTERORYTIS  Conrad,  1862:  17 

Type  sp.  (O.  D.):  Murex  umbrifer  Conrad,  1832, 
Foss.  Shells  Tert.  form.  N.  Amer.   1:   17,  pi.  3, 

fig.  1. 
PTEROTYPHIS  Jousseaume,   1881:  338  (Emenda- 
tion for  Perotvphis  Jousseaume,  1880) 
PTERYMUREX  Rovereto,  1899:  105 

New  name  for  Pteronotus  Swainson,    1833,   not 

Rafinesque,  1815. 
tPTERYNOPSIS  E.  H.  Vokes,  1972:  1 

Type  sp.  (O.  D.):  Pterynopsis  prosopeion  E.  H. 

Vokes,  1972  (=  Murex  nysti  von  Koenen,  1867, 

not  Roualt,  1850),  Bull.  Inst.  R.  Sci.  Nat.  Belg. 


286 


48(9):  2. 

PTERYNOTUS  Swainson.  1833:  pi.  100 

Type  sp.  (S.  D.  Swainson.  1833,  pi.  122):  Murex 
pinnatus  Swainson,  1822  (=  Purpura  alatus 
Roding.  1798),  App.  Cat.  Bligh,  p.  17. 

PURPURA  Bruguiere.  1789:  xv 

Type  sp.  (S.  D.,  Montfort,  1810):  Buccinum 
pepsicum  Linne,  1758,  Syst.  Nat..  Ed.  10,  p.  738, 
no.  401. 

PURPURELLA  Dall,  1871:  110  (not  Bellardi,  1882, 
nor  Robineau-Desvoidy,  1853) 
Type  sp.  (O.  D.):  Purpura  columellaris  Lamarck, 
1816,  Tabl.  Encycl.  Meth..  pi.  398,  figs.  3a,  3b, 
Liste.  p.  2. 

tPURPURELLA  Bellardi.  1882:  193  (not  Dall,  1871. 
nor  Robineau-Desvoidy,  1853) 
Type  sp.  (M.):  Purpurella  canaliculata  Bellardi, 
1882,  Moll.  Terr.  Terz.  Piem.  Ligur.  pt.  3,  p.  193, 
pi.  11,  fig.  35. 

PURPURELLUS  Jousseaume,  1880:  335 

Type  sp.  (O.  D.):  Murex  gambiensis  Reeve,  1845, 
Conch.  Icon.  vol.  3.  Murex.  pi.  16,  sp.  65. 

tPURPURINA  Cox,  1961:  10 

Type  sp.  (O.  D.):  Purpurina  yanreyensis  Cox, 
1961,  Bull.  Australian  Bur.  Min.  Res.  Geol. 
Geophys.,  pp.  10,  33,  pi.  7,  figs.  6a,  6b. 

QUOYULA  Iredale.  1912:  221 

Type  sp.  (O.  D):  Purpura  monodonta  Quoy  & 
Gaimard,  1833,  Voy.  Astrolabe.  Zool.  2,  p.  561, 
pi.  37.  figs.  9.  11. 

RAPA  "Klein"  Bruguiere.  1792:  533 
Type  sp.  (S.  D..  Herrmannsen.  1848):  Bulla  rapa 
(Linne,   1767)  (=  Murex  rapa  Linne.   1758.  not 
Gmelin.   1791).  Syst.  Nat.  Ed.   12.  p.   1184.  no. 
384. 

RAPANA  Schumacher.  1817:  214 
Type    sp.    (M.):    Rapana  foliacea    Schumacher. 
1817   (=   Buccinum   bezoar  Linne,    1758),    Ess. 
Vers.  Test.,  p.  214. 

RAPANUS  "Schum."  Sowerby,  1839:  92 

Type  sp.  (M.):  Pyrula  papyracea  [Lamarck,  1816] 
(=  Rapa  rapa  Bruguiere,  1792),  Tabl.  Encycl. 
Meth.  pi.  436,  fig.  1. 

RAPELLA  Swainson,  1840:  82,  307 
Type  sp.  (M.):  Rapella  papracia  (sic)  (Lamarck, 
1816),  Tabl.  Encycl.  Meth.  pi.  436,  fig.  1,  Liste, 
p.  8. 

RHINOCANTHA  H.  &  A.  Adams,  1853:  72 
Type  sp.    (S.    D.,    E.    H.    Yokes,    1964):   Murex 
brandaris  Linne,  1758,  Syst.  Nat.,  Ed.  10,  p.  747, 
no.  446. 

RHIZOCHILUS  Steenstrup.  1850:  75 

Type  sp.  (M.):  Rhizochilus  antipathum  Steen- 
strup. 1850,  Overs.  K.  Danske  Vidensk.  Selsk. 
Forh.  1850:  75. 

RHIZOPHORIMUREX  Oyama.  1950:  10 

Type  sp.  (O.  D.):  Murex  capuchinus  (sic) 
Lamarck,  1822,  Hist.  Nat.  Anim.  s.  Vert.  7:  164. 

RHOMBOTHAIS  Woolacott,  1954:  38 

Type  sp.  (O.  D.):  Rhombothais  arbutum  Wool- 
acott, 1954,  Proc.  Roy.  Soc.  Zool.  Soc.  New  South 
Wales,  1952-53:  38,  pi.  3.  figs.  1,  2. 

RICINELLA  Schumacher.  1817:  72 

Type  sp.  (S.  D.,  Iredale,  1937):  Ricinella  pur- 
purata  Schumacher,  1817,  Ess.  Syst.  Vers.  Test., 
p.  72. 

RICINULA  Lamarck,  1816:  1 

Type  sp.  (S.  D.,  Children.  1823):  Ricinula  huridu 
(sic)  Lamarck.  1816,  Tabl.  Encycl.  Meth..  pi.  395, 


figs,  la,  lb. 

RISOMUREX  Olsson  &  McGinty,  1958:  40 

Typesp.  (O.  D.):  Engina  schrammi  Crosse,  1863, 
J.  Conchyl.  11:  86,  pi.  1,  fig.  2. 

ROPERIA  Dall,  1898:  5 
Type  sp.  (O.  D.):  Fusus  roperi  Dall,   1898,  The 
Nautilus  12(1):  5. 

RUDOLPHA  Schumacher,  1817:  63,  210 

Type  sp.  (O.  D.):  Buccinum  mo«oceros  Chemnitz, 
1788  (non-bionominal)  (=  Buccinum  monodon 
Pallas,  1774),  Neues  Systematisches  Conchylien- 
Cabinet  10:  197.  pi.  154.  figs.  1469,  1470. 

tRUGOTYPHlS  Vella,  1961:  376 

Typesp.  (O.  D.):  Typhis  francescae  Finlay.  1924. 
Trans.  Proc.  New  Zealand  Inst.  55:  465,  pi.  49, 
figs.  6a,  6b. 

tSARGANA  Stephenson,  1923:  377 
Type  sp.  (O.  D.):  Rapana  stantoni  Stephenson, 
1923,  North  Carolina  Geol.  Surv..  5:  377. 

tSCALASPIRA  Conrad.  1862:  560 

Type  sp.  (M.):  Fusus  strumosa  Conrad,  1862, 
Proc.  Acad.  Nat.  Sci.  Philadelphia  14:  560. 

SEMIRICINULA  von  Martens,  1903:  95 
Type  sp.  (M.):  Purpura  muricina  Blainville.  1832, 
Nouv.  Ann.  Mus.  Hist.  Nat.  Paris  1:  218. 

tSEMITYPHIS  K.  Martin.  1931:  31 

Type  sp.  (M.):  Semitvphis  incisus  K.  Martin, 
1931,  Wetens.  Meded.'l8:  31,  pi.  5,  figs,  la,  lb. 

tSERRATIFUSUS  Darragh.  1969:  89 

Type  sp.  (O.  D.):  Fusus  crasspedotus  Tate,  1888, 
Trans.  Roy.  Soc.  South  Australia  10:  134.  pi.  8. 
fig.  4. 

SHASKYUS  Burch  &  Campbell.  1963:  203 
Type  sp.  (O.  D.):  Murex  festivus  Hinds.    1844. 
Proc.  Zool.  Soc.  London  11:  127. 

SIPHONOCHELUS  Jousseaume.  1880:  335 
Type  sp.  (O.  D.):   Typhis  avenatus  (sic)   Hinds, 
1843  (=   T.   arcuatu's  Hinds.   1843),  Proc.  Zool. 
Soc.  London  11:  19. 

SIRATUS  Jousseaume,  1880:  335 
Type  sp.  (O.  D.):  Purpura  sirat  Adanson,   1757 
(non-binominal)  (=  Murex  senegalensis  Gmelin, 
1791),  Hist.  Nat.  Senegal,  p.  125. 

SISTRUM  Montfort,  1810:  595 
Type  sp.   (M.):  Sistrum  album  Montfort,    1810, 
Conch.  Syst.  2:  595. 

SPINIDRUPA  Habe  &  Kosuge,  1966:  330 

Type  sp.  (O.  D.):  Murex  euracantha  A.  Adams, 
1851,  Proc.  Zool.  Soc.  London  18:  268. 

SPINOSTOMA  Coen,  1943:  90 
Type  sp.  (S.  D.  herein):  Murex  nuttalli  Conrad, 
1837,   J.   Acad.   Nat.   Sci.    Philadelphia   7:    264, 
pi.  20,  tig.  22. 

STRAMONITA  Schumacher,  1817:  226 

Type  sp.  (S.  D.  Gray,  1847):  Buccinum  haemas- 
toma  Linne,  1867,  Syst.  Nat.,  Ed.  12,  p.  1202, 
no.  566. 

STRAMONITROPHON  Powell,  1951:  156 

Type  sp.  (O.  D.):  Buccinum  laciniatus  Martyn, 
1788  (non-binominal)  (=  Buccinum  laciniatum 
"Martyn"  Diilwyn,  1817),  Univ.  Conch.  2:42. 

SUBPTERYNOTUS  Olsson  &  Harbison.  1953:  246 
Type  sp.  (O.  D.):  Murex  textilis  Gabb,  1873, 
Trans.  Amer.  Philos.  Soc.  (n.s.)  15(1):  202. 

TAKIA  Kuroda,  1953:  190 

Type  sp.  (O.  D.):  Murex  inermis  Sowerby,  1841 
(not  Philippi,  1836)  (=  Dermomurex  [Takia] 
infrons  E.  H.  Vokes,  1974),  Venus  17(4):  190. 

TALITYPHIS  Jousseaume.  1882:  338 


287 


Type  sp.  (O.  D.):  Typhis  expansus  Sowerby,  1874, 

Proc.  Zool.  Soc.  LxJndon  42:  719,  pi.  59,  fig.  4. 
TARANTELLAXIS  Habe,  1970:  85 

Type   sp.    (M.):    Tarantellaxis    kuroharai   Habe, 

1970,  Venus  29(3):  85. 
tTAURASIA  Bellardi,  1882:  194 

Type  sp.  (O.  D.):  Purpura  subfusiformis  Orbigny, 

1952.  Prodrome  Paleont  ...  3:  15. 
TENGUELLA  Arakawa.  1965:  123 

Type  sp.  (O.  D.):  Morula  granulata  Duclos,  1924 

(sic)  (correct  date  Duclos,  1832).  Ann.  Sci.  Nat. 

26(101):  111. 
TEREFUNDUS  Finlay,  1927:  425 

Type   sp.    (O.    D.):    Trophon    crispulatus    Suter, 

1908,  Proc.  Malac.  Soc.  London  8:   178,  pi.   7, 

fig.  2. 
TERNARIA  Coen,  1943:  89 

Type  sp.    (S.    D..   E.   H.    Yokes,    1964):   Murex 

eurypteron   Reeve.    1845,   Conch.    Icon.,   vol.    3, 

Murex.  pi.  34,  sp.  176. 
THAIS  Roding,  1798:  54 

Type  sp.  (S.  D.  Stewart,  1926):  Thais  lena  Roding, 

1798  (=  Merita  nodosa  Linne,  1758,  p.  777.) 
THAISELLA  Clench,  1947:  69 

Type  sp.   (O.  D.):  Purpura  trinitatensis  Guppy, 

1869.  Proc.  Sci.  Assoc.  Trinidad  1:  366. 
THALESSA  H.  &  A.  Adams.  1853:  127 

Type  sp.  (S.  D.,  Cossmann.  1903):  Purpura  hypo- 

castaneum  "Linne."  Cossmann.  1903  (=  Murex 

hippocastanum  "Linne"  Auct.)  (=  Murex  hippo- 

castanum  Gmelin,  1791.  p.  3539.) 
tTIMBELLUS  de  Gregorio.  1885:  275 

Typesp.  (O.  D.):  Murex  latifolius  Bellardi.  1872, 

Moll.  Terr.  Terz.  Piem.  Ligur.  pt.  1,  p.  54,  pi.  4, 

fig.  5. 
tTIMOTHIA  Palmer,  1938:  3 

New  name  for  A  Id  rich  ia  Palmer,  1937,  not  Coquil- 

let.  1894.  nor  Vaughan,  1900. 
TOLEMA  Iredale,  1929a:  186 

Type  sp.  (O.  D.):  Purpura  sertata  Hedley,   1903, 

Mem.  Australian  Mus.  4:  382,  figs.  95,  96.  (ICZN 

ruling.  Opinion  911,  1970.  Bull.  Zool.  Nomencl. 

27:  20  fixed  the  type  as  Tolema  australis  Laseron, 

1955). 
TORVAMUREX  Iredale,  1936:  323 

Type  sp.  (O.  D.):  Triplex  denudatus  Perry,  1811, 

Conchology  .  .  .,  pi.  7,  fig.  2. 
TRACHYPOLLIA  Woodring,  1928:  268 

Type  sp.  (O.  D.):  Trachvpollia  sclera  Woodring, 

1928,  Carnegie  Inst.  Wash.  Publ.  385:  269,  pi.  16, 
figs.  7,  8. 

TRANSTRAFER  Iredale,  1929b:  290 

Type  sp.  (O.  D.):   Transtrafer  longmani  Iredale, 

1929,  Mem.  Queensland  Mus.  9:  290. 
TRIALATELLA  Berry,  1964:  149 

Type  sp.  (O.  D.):  Trialatella  cunninghamae  Berry, 
1964,  Leaflets  in  Malacology  1(24):  149. 

TRIBULUS  "Klein"  Bruguiere,  1792:  530 

Type  sp.  (S.  D.,  Wenz,  1941):  Mancinella  {Tri- 
bulus)  planospira  Lamarck,  1822,  Hist.  Nat. 
Anim.  s.  Vert.  7:  240. 

TRIGONOTYPHIS  Jousseaume,  1881:  339 

Type  sp.  (O.  D.):  Typhis  Jimbriatus  A.  Adams, 
1854.  Proc.  Zool.  Soc.  London  21:  71. 

TRIPLEX  Perry.  1810:  M7 

Type  sp.  (M.):  Triplex  foliatus  Perry.  1810  (  = 
Murex  palmarosae  Lamarck.  1822).  The  Arcana, 
or  the  Museum  Nat.  Hist.,  p.  M7.  (Triplex  foliatus 
Perry  has  been  suppressed  by  the  ICZN,  Opinion 


911,  1970,  Bull.  Zool.  Nomencl.  27:  20). 
TRIPTEROTYPHIS  Pilsbry  &  Lowe,  1932:  78 

Type  sp.  (O.  D.):  Typhis  lowei  Pilsbry,  1931,  The 

Nautilus  45(2):  72. 
TRIREMIS  "Bayle"  P.  Fischer,  1884:  641 

Type  sp.   (M.):  Murex  gambiensis   Reeve,    1845, 

Conch.  Icon.,  vol.  3,  Murex  pi.  16,  sp.  65. 
TRITONALIA  "Fleming"  Gray,    1847b:    122   (not 
Fleming,  1828) 

Type  sp.  (O.  D.):  Murex  erinaceus  Linne,   1758, 

Syst.  Nat.,  Ed.  10.  p.  748.  no.  451. 
TROCHIA  Swainson,  1840:  302 

Type    sp.     (M.):     Trochia    sulcata    "Lamarck," 

Swainson.  1840  (=  Buccinum  cingulatum  Linne, 

1771).  Treat.  Malac.  p.  302. 
TROMINA  Dall.  1918a:  137 

Type  sp.   (O.   D.):  Fusus  unicarinatus  Philippi, 

1868.  Malak.  Bl.  15:  223. 
TROPHON  Montfort.  1810:  483 

Type  sp.  (O.  D.):  Trophon  magellanicus  Gmelin, 

1791    (=    Buccinum   geversianus   Pallas,    1774), 

Syst.  Nat.,  Ed.  13,  p.  3548,  no.  80. 
TROPHONOPSIS  Bucquoy  &  Dautzenberg, 
1882:  40 

Type  sp.    (O.    D.):   Murex   muricatus   Montagu, 

1803,  Test.  Brit..  1:  262.  pi.  9,  fig.  2. 
TRUBATSA  Dall.  1889:  215 

Type  sp.  (S.  D..  Keen,  1944):  Typhis  (Trubatsa) 

longicomis  Dall  (in,  Agassiz),   1888,  The  Three 

Cruises  of  the  "Blake,"  2:  70,  fig.  294. 
TRUNCULARIA  Monterosato,  1917:  20  (not 
Wiegmann,  1832) 

Type  sp.  (O.  D.):  Murex  trunculus  Linne,   1758, 

Syst.  Nat.,  Ed.  10,  p.  747,  no.  447. 
TRUNCULARIOPSIS  Cossmann,  1921:  79 

New  name  for   Truncularia  Monterosato,    1917, 

not  Wiegmann,  1832. 
TUBICAUDA  Jousseaume,  1880:  335 

Type  sp.   (O.   D.):   Murex  brevispina   Lamarck, 

1822.  Hist.  Nat.  Anim.  s.  Vert.  7:  159. 
TYPHINA  Jousseaume.  1880:  335 

Type  sp.  (O.  D.):  Typhis  belcheri  Broderip.  1833, 

Proc.  Comm.  Sci.  Zool.  Soc.  London  2:  178. 
TYPHINELLUS  Jousseaume.  1880:  335 

Type  sp.  (O.  D.):  Typhis  sowerbiyi  (sic)  Broderip, 

1833  (=   Typhis  sowerbii  Broderip,  1833),  Proc. 

Comm.  Sci.  Zool.  Soc.  London  2:  178. 
TYPHISALA  Jousseaume,  1881:  339 

Type  sp.  (O.  D.):  Typhis  grandis  A.  Adams,  1855, 

Proc.  Zool.  Soc.  London  22:  41. 
TYPHISOPSIS  Jousseaume,  1880:  335 

Type   sp.    (O.    D.):    Typhis   coronatus   Broderip, 

1833.  Proc.  Comm.  Sci.  Zool.  Soc.  London  2:  178. 
UNICORNUS  Montfort,  1810:  454 

Type   sp.    (O.    D.):    Unicornus    typus   Montfort, 

1810    (=    Buccinum    monodon    Pallas,     1774), 

Conch.  Syst.  2:  454,  pi.  114. 
UROSALPINX  Stimpson.  1865:  58 

Type  sp.   (O.   D.):  Fusus  cinereus  Say,   1822,  J. 

Acad.  Nat.  Sci.  Philadelphia  2:  236. 
USILLA  H.  Adams,  1860:  369 

Type  sp.  (O.  D.):  Vexilla  nigro-fusca  Pease,  1860 

(=   Vexilla  fusconigra  Pease,   1860),  Proc.  Zool. 

Soc.  London  27:  141. 
tUTTLEYA  Marwick,  1934:  19 

Type  sp.  (O.  D.):  Uttleya  arcana  Marwick,  1934, 

Proc.  Malac.  Soc.  London  21:  19. 
tVESANULA  Finlay.  1926:  245 

Type  sp.  (M.):  Trophon  chaskanon  Finlay,  1926, 


288 


Trans.  New  Zealand  Inst.  56:  245. 

VEXILLA  Swainson,  1840:  300 

Type  sp.  (M.):  Vexilla  picta  Swainson,  1840  (  = 
Murex  vexillum  Gmelin.  1791),  Treat.  Malac, 
p.  300. 

VIATOR  E.  H.  Yokes,  1974:  4 

Type  sp.  (O.  D.):  Viator  antonius  E.  H.  Yokes, 
1974.  J.  Malac.  Soc.  Australia  3(1):  4. 

VITULARIA  Swainson,  1840:  297 

Type  sp.  (M.):  Vitidaria  tuberculata  Swainson, 
1840  (=  Murex  miliaris  Gmelin,  1791),  Treat. 
Malac,  p.  297. 

VITULINA  Swainson,  1840:  64 

Type  sp.  (O.  D.):  Murex  vitulina  Lamarck,  1816 
(=  Murex  miliaris  Gmelin,  1791),  Tabl.  Encycl. 
Meth.,  pi.  419,  figs,  la,  lb,  Liste,  p.  5. 

tWIDNINGlA  Ludbrook,  1941:  95 

Type  sp.  (O.  D.):  Widningia  crassiplicata  Lud- 
brook, 1941,  Trans.  Roy.  Soc.  South  Australia 
65(1):  95. 

XANTHOCHORUS  P.  Fischer,  1884:  639 

Type  sp.  (M.):  Trophon  xanthostoma  Broderip, 
1833,  Proc.  Zool.  Soc.  London  1:  8. 


XENOTROPHON  Iredale,  1929a:  184 

Type  sp.  (O.  D.):  Trophon  euschema  Iredale, 
1929,  Rec.  Australian  Mus.  17:  184,  pi.  40,  fig.  3. 

XYMENE  Iredale.  1915:  471 

Type  sp.  (O.  D.):  Fusus  plebius  Hutton,  1873, 
Cat.  Mar.  Moll.  New  Zealand,  p.  9. 

XYMENELLA  Finlay,  1927:  424 

Type  sp.  (O.  D.):  Trophon  pusillus  Suter,  1907, 
Trans.  New  Zealand  Inst.  39:  253,  pi.  19,  fig.  10. 

XYMENOPSIS  Powell.  1951:  158 

Type  sp.  (O.  D.):  Fusus  liratus  "Couthouy" 
Gould.  1849,  Proc.  Boston  Soc.  Nat.  Hist.  3:  141. 

tYASILA  Olsson,  1930:  59 

Type  sp.  (O.  D.):  Yasila  paytensis  Olsson,  1930, 
Bull.  Amer.  Paleo.  17(62):  59. 

ZACATROPHON  Hertlein  &  Strong,  1951:  86 
Type  sp.  (O.  D.):  Trophon  (Zacatrophon)  beebei, 
Hertlein  &  Strong,  1951,  Zoologica  36:  86. 

ZEATROPHON  Finlay,  1927:  424 

Type  sp.  (O.  D.):  Fusus  ambiguus  Philippi,  1844, 
Abbild.  Beschreib.  Conch..  Fusus,  p.  107.  pi.  1. 
fig.  2. 


GENERIC  REFERENCES 


Adams.  H..  1860.  On  two  new  genera  of  acephalous 
mollusks.  Proc.  Zool.  Soc.  London  28:  369-371. 

Adams,  H.  and  A.,  1853-1858.  The  Genera  of 
Recent  Mollusca,  Yan  Yoorst,  London,  vol.  1, 
484  pp. 

Arakawa,  K.  Y..  1964.  A  study  on  the  Radulae  of 
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[1957]:  112-117,  10  figs. 


Department  of  Marine  Invertebrates, 
Natural  History  Museum,  San  Diego, 
California  92112. 


v5^^^ 


AU8  191970 

BLOOD  CIRCULATION  IN  FOUR  SPECIES  OF  BARNACLES 
(LEPAS,  CONCHODERMA:  LEPADIDAE) 

BRYAN  R.  BURNETT 


TRANSACTIONS 

OF  THE  SAN   DIEGO 
SOCIETY   OF 
NATURAL  HISTORY 

VOL.  17,  NO.  21         20  JUNE  1975 


BLOOD  CIRCULATION  IN  FOUR  SPECIES  OF  BARNACLES 
ilEPAS,  CONCHODERMA:  LEPADIDAE) 


BRYAN  R.  BURNETT 


ABSTRACT. — Circulatory  morphologies  of  the  primitive  lepadomorphans  Lepas  anatifera,  L.  pectinata 
pacifica.  L.  fasicularis  and  Conchoderma  virgatum  are  similar,  but  major  differences  appear  in  vessel 
caliber  and  refinement  of  the  basic  system.  The  smaller  species  (L.  fasicularis  and  L.  pectinata  pacifica) 
have  larger  vessels  for  their  body  size  than  the  larger  species  (Z,.  anatifera  and  C.  virgatum).  Circulatory 
organization  of  the  lepadids  is  simpler  than  that  of  Pollicipes  polymerus  (Scalpellidae)  and  Balanus 
tintinnahulum  (Balanidae).  The  lepadid  rostral  vessel,  which  is  morphologically  similar  to  that  of  P. 
polyments,  is  interpreted  as  a  vestige  of  the  heart.  Pump  function  can  be  attributed  to  the  rostral  sinus 
(the  blood  pump),  which  is  apparently  a  remnant  of  the  pericardial  sinus.  Transfer  of  hemolymph 
pumping  from  the  heart  to  the  rostral  sinus  probably  occurred  with  the  development  of  the  peduncle. 

Detailed  accounts  of  cirriped  circulation  may  be  found  in  Cannon  (1947)  and  Burnett 
(1972).  The  circulatory  systems  of  Lithotrya  valentiana  and  Pollicipes  polymerus 
(Cannon,  1947)  had  been  considered  to  represent  the  general  condition  for  thoracican 
Cirripedia  (e.g.  Maynard,  1960).  However,  Burnett  (1972)  showed  that  the  circulatory 
system  of  the  pedunculate  barnacle  Pollicipes  polymerus  was  unlike  that  of  other 
Crustacea.  In  order  to  obtain  a  more  complete  understanding  of  circulatory  relationships 
in  the  Cirripedia,  I  studied  the  circulatory  systems  of  four  species  of  Lepadidae:  Lepas 
anatifera,  L.  pectinata  pacifica,  L.  fasicularis  and  Conchoderma  virgatum. 

MATERIALS  AND  METHODS 

The  three  species  o^  Lepas  were  collected  from  debris  that  washed  ashore  at  Scripps 
Institution  of  Oceanography,  La  Jolla,  California  in  the  summers  of  1972  and  1973.  The 
Conchoderma  virgatum  were  collected  from  a  Pacific  Ridley  sea  turtle  {Lepidochelys 
olivacea)  captured  off  La  Jolla.  Living  specimens  were  injected  with  yellow  (MV-122)  or 
maroon  (MV-118)  Microfil  (Canton  Bio-Medical  Products,  Inc.  P.O.  Box  2017,  Boulder, 
Colorado  80302),  either  into  the  peduncle  or  through  the  adductor  scutorum  into  the 
rostral  sinus,  following  techniques  developed  in  an  earlier  study  (Burnett,  1972).  The 
amount  injected  ranged  from  0.5  to  2.0  ml  based  on  the  size  of  the  animal.  In  each 
species,  the  rostral  valve  at  the  posterior-most  part  of  the  peduncular  vessel  usually  did 
not  hold  under  the  pressure  exerted  from  the  Microfil  injections  into  the  peduncle;  almost 
always  a  significant  amount  of  Microfil  entered  the  body  via  the  peduncular  vessel.  The 
rostral  valve  in  the  lepadids  is  more  delicate  than  that  of  P.  polymerus:  consequently  their 
vessels  are  more  prone  to  rupture  and  distort,  which  makes  it  difficult  to  trace  circulatory 
pathways,  especially  with  the  peripheral-collecting  circulation.  In  order  to  determine 
vessel  wall  structure,  portions  of  the  gut  vessels  were  removed  (while  they  still  had 
solidified  Microfil  in  the  vessel  lumina)  and  embedded  in  Spurr  (Polysciences,  Inc.  Paul 
Valley  Industrial  Park,  Warrington,  Penna.  18976).  Sections,  2[j.m  thick,  were  made  with 
a  glass  knife  on  a  Porter-Blum  JB-4  microtome. 

Body  movements  of  L.  fasicularis  were  observed  through  a  dissection  microscope  by 
shining  a  light  through  the  thin  walled  capitulum. 

CIRCULATORY  MORPHOLOGY 

Basically,  I  shall  follow  Burnett  (1972)  in  dividing  the  barnacle  circulatory  system 
into  three  arbitrary  divisions:  1)  the  circulation  of  the  peduncle  and  mantle,  2)  the 
distributive  circulation  and  3)  the  peripheral-collecting  circulation. 

SAN  DIEGO  SOC.  NAT.  HIST..  TRANS.  17(21):  293-304,  20  JUNE  1975 


294 


AD  SCUT 
AF  AD  SCUT 


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^;5^6»<*  •»  ^ » y,»  ij^^iMati<ya»,f°fe^^£flF  mancirc. 


>=»  o'  ^ 


SCUT  VES 


ADSCUT 


OW  FR, 


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Figure  1 .  Mantle  circulation  in  L.  anatifera.  a.  View  of  the    carina  with  the  carinal  vessel  situated  along  the 

midline  of  the  valve,  b.  View  of  the  right  side  of  the  mantle  and  a  portion  of  the  peduncle.  Only  the  intervalve 

circulation  is  shown,  with  the  arrows  representing  the  direction  of  hemolymph  flow.  Basically,  this  pattern  of 

circulation  is  present  in  L.  pectinata  pacijlca  and  C.  virgatum.  Abbrevations  are  explained  in  the  Appendix. 

Figure  2.  Mantle  circulation  (right  side)  in  L.  fasicularis.  Arrows  show  the  direction  of  flow  with  the  heavier 

arrows  indicating  major  hemolymph  flow. 

Figure  3.  The  origin  and  associated  circulation  of  the  carinal  vessel  in  L.  fasicularis. 

Figure  4.  A  portion  of  the  tergal  plexus  from  L.  pectinata  pacifica. 

Figure  5.  The  double  circulation  of  the  mantle  in  C  virgatum. 

Figure  6.  A  portion  of  the  tergal  plexus  from  L.  anatifera. 


295 


Circulation  of  the  peduncle  and  mantle. — The  peduncular  vessel  extends  the  full 
length  of  the  peduncle  without  giving  off  any  branches,  and  ends  with  a  gradual 
enlargement  at  the  basal  disc.  From  the  basal  disc,  the  hemolymph  percolates  towards 
the  mantle,  and  it  appears  that  the  entire  peduncle  is  a  single  sinus. 

Hemolymph  from  the  peduncle  enters  the  mantle  circulation  by  two  pathways  (Fig. 
1).  In  one  route,  blood  is  directed  into  the  mantle  circulation  from  the  posterior-most  part 
of  the  peduncle  by  a  series  of  short  parallel  vessels  (Fig.  lb).  From  these  vessels  the  blood 
moves  through  a  plexus  toward  the  ovigerous  frena  and  eventually  into  the  paired  scutal 
vessels.  Lepas  fasicularis  has  enlarged  vessels  entering  the  mantle  in  the  area  between  the 
scutal  and  carinal  regions  (Figs.  2,  3). 

The  other  pathway  by  which  the  blood  enters  the  mantle  is  through  the  carinal  vessel 
(Figs.  1-3).  In  Lithotrya  valentiana.  Cannon  (1947)  described  a  pair  of  vessels  in  the 
mantle  region  between  the  terga  and  carina,  but  I  doubt  these  are  homologous  to  the 
carinal  vessel  of  the  Lepadidae,  which  is  unpaired.  The  carinal  vessel  extends  the  full 
length  of  the  carina,  and  gives  off  smaller  vessels  along  its  entire  length.  Almost  all  of  the 
tergal  area  and  a  good  portion  of  the  scutal  area  of  the  mantle  is  supplied  by  this  vessel. 

In  lepadids,  the  ovigerous  frena  (Figs.  1,  2)  are  highly  vascularized  with  a  circulation 
similar  to  the  rest  of  the  mantle.  A  vessel,  connecting  the  scutal  vessel  on  each  side  of  the 
mantle,  borders  the  distal  margin  of  each  ovigerous  frenum.  In  L.  fasicularis,  the 
ovigerous  frena  are  bilobate  (Fig.  2),  with  a  large  vessel  extending  along  the  distal  margin 
of  each  lobe.  These  vessels  join  and  the  resulting  vessel  connects  to  the  scutal  vessels, 
which  in  turn  enter  the  body.  In  contrast  to  the  situation  in  P.  polymerus,  the  mantle 
knobs  and  the  circulation  associated  with  the  mantle  muscles  are  not  present  in  the 
lepadids. 

The  paired  scutal  vessels  partially  circle  the  adductor  scutorum  at  the  muscle's 
insertion  on  the  two  scutal  plates  (Figs.  1,  2)  in  a  manner  similar  to  that  found  in  P. 
polymerus.  The  scutal  valve  (Fig.  1)  lies  just  inside  the  entrance  of  the  scutal  vessel  into 
the  body. 

The  circulation  of  the  mantle  varies  between  species  (Figs.  3-6),  with  the  plexuses 
appearing  random  in  L.  anatifera  (Fig.  6)  to  fairly  organized  in  L.  pectinate  pacifica  (Fig. 
4).  The  mantle  circulation  of  C.  virgatum  (Fig.  5)  differs  from  Lepas  in  being  essentially  a 
double  system  in  which  plexuses  are  associated  with  both  the  external  and  internal 
cuticles  of  the  capitulum.  Scattered  connections  exist  between  these  two  plexuses. 

In  L.  anatifera,  the  vessels  between  the  capitular  plates  enlarge  somewhat;  in  C. 
virgatum  the  plexal  vessels  appear  uniform  throughout  the  mantle,  but  enlarge  as  they 
approach  the  scutal  vessels. 

Distributive  circulation. — Near  the  points  where  the  adductor  scutorum  inserts  on 
the  scuta,  the  two  scutal  vessels  enter  the  body  from  the  mantle  and  enlarge  to  form  the 
paired  scutal  sinuses  (Figs.  7-10).  In  all  four  species,  as  in  P.  polymerus,  the  scutal 
sinuses  are  located  on  each  side  of  the  rostral  sinus,  in  close  proximity  to  the  adductor 
scutorum.  The  precise  position  of  the  scutal  sinuses  varies  from  species  to  species:  in  L. 
anatifera  they  are  mostly  posterior  to  the  adductor  scutorum;  in  L.  fasicularis  they  are 
anterior;  and  in  L.  pectinata  pacifica  and  C.  virgatum  they  are  dorsal.  Their  shape  and 
extent  also  varies. 

The  adductor  scutorum  receives  blood  from  the  scutal  sinuses  in  all  species.  The 
afferent  circulation  to  this  muscle  is  located  immediately  posterior  to  the  scutal  valves. 
From  the  scutal  sinuses  hemolymph  enters  the  vessels  of  the  gut,  gastric  gland,  and  the 
maxillary  gland. 

On  the  gut,  the  gastric  plexus  continues  around  most  of  the  cephalic  portion  of  the 
gut  with  little  variation  in  vessel  caliber  (Figs.  7-10).  In  the  posterior  part  of  the  cephalic 
gut,  the  paired  inferior  gastric  vessels  continue  from  vessels  of  the  gastric  plexus.  As  this 
pair  of  vessels  continues  posteriorly,  branches  of  the  gastric  gland  plexus  also  combine 
with  the  inferior  gastric  vessels. 

The  paired  inferior  gastric  vessels  join  on  the  ventral  surface  of  the  thoracic  gut 
above  the  first  pair  of  cirri.  This  combined  vessel  (the  posterior  inferior  gastric  vessel) 
continues  posteriorly  and   descends  to  contact  the  epineural  sinus  by  one   or   more 


296 


CEPH.  TR.  M. 


INF  GAS.  VES. 


CAST  GL. 

SUB.  VES. 


THOR.  TR  M.  I 


POST  INF  GAS  VES. 

EPI.SIN, 


-AD  SCUT 


CEPH.TR.  M. 

INF  GAS.  VES. 


POST  INFGAS  VES. 


SUB.  VES 

EPISIN. 


SCUT  SIN 


Figure  7.  Distributive  circulation  as  seen  from  the  left  side  of  the  body  in  I.  anatifera.  Numbers  1-6  refer  to  the 
positions  of  the  respective  cirri. 

Figure  8.  Distributive  circulation  in  L.  fasicularis. 

branches.  In  L.  anatifera,  the  posterior  inferior  gastric  vessel  is  reduced  and  has  only  one 
contact  with  the  epineural  sinus.  In  all  lepadids,  the  epineural  sinus,  which  surrounds  the 
nerve  cord  at  the  base  of  the  cirri,  receives  blood  from  two  additional  sources:  1)  the 
scutal  sinuses  through  the  maxillary  gland,  to  connect  to  the  anterior  part  of  the 
epineural  sinus,  and  2)  a  connection  by  the  subintestinal  vessel  from  the  gastric  gland 


297 


CEPH  TR  M 


GAS.VES. 


/NFGAS.VES. 

POST  SIN. 


AD.SCUT 


SUBVES 

THORTR.M.  182 


Figure  9.  Distributive  circulation  in  L.  pectinata  pacifica. 
Figure  10.  Distributive  circulation  in  C.  virgatum. 

plexus.  Blood  from  the  epineural  sinus  goes  to  the  cirri,  penis  and  oral  cone. 

In  L.  anatifera  the  subintestinal  vessel  originates  as  a  pair  of  vessels  among  the 
plexus  surrounding  the  gastric  gland.  These  two  vessels  collect  blood  from  the  gastric 
gland  plexus  and  enlarge  as  they  descend  toward  the  epineural  sinus.  The  subintestinal 
vessel  is  then  formed  by  the  combining  of  the  two  vessels  just  anterior  to  the  first  thoracic 
transverse  muscle.  On  the  anterior  part  of  the  epineural  sinus,  the  subintestinal  vessel 


298 


enters  slightly  dorsal  to  the  maxillary  gland  connections.  In  the  other  three  species  the 
subintestinal  vessel  consists  of  one  or  two  short  vessels  connecting  the  gastric  gland  plexus 
to  the  epineural  sinus. 

The  distributive  circulation  of  Z..  fasicularis  is  more  grossly  constructed  in  contrast  to 
L.  atiatifem.  The  posterior  inferior  gastric  vessel  unites  with  the  epineural  sinus  to  form  a 
large  posterior  sinus.  The  afferents  to  the  oral  cone,  which  originate  from  the  epineural 
sinus,  are  divided  into  two  vessels,  the  afferent  mandibular  vessel  going  directly  to  the 
mandibles,  and  the  afferent  oral  cone  vessel  to  the  rest  of  the  oral  cone.  The  afferent 
filamentary  vessels  to  the  four  filamentary  appendages  at  the  base  of  each  first  cirrus 
originate  on  the  anterior-most  part  of  the  epineural  sinus. 

Circulatory  morphology  in  L.  pectinata  paciftca  appears  most  similar  to  that  of  L. 
fasicularis.  The  union  of  the  posterior  inferior  gastric  vessel  to  the  epineural  sinus  is  so 
extensive  that  the  two  make  up  a  single  sinus  (the  posterior  sinus)  posterior  to  the  second 
thoracic  segment.  The  gastric  gland  plexus  does  not  extend  directly  around  the  cephalic 
transverse  muscle  and  the  large  sinus  at  the  base  of  the  first  cirrus  is  not  present  as  it  is  in 
L.  fasicularis. 

The  distributive  system  of  C.  virgatum  is  similar  to  that  of  Lepas  (Fig.  10).  The  gut 
plexus  is  strongly  directionally  oriented.  The  connection  of  the  posterior  inferior  gastric 
vessel  to  the  epineural  sinus  is  by  two  or  three  large  caliber  vessels.  Plexal  circulation  of 
the  gastric  gland  appears  more  haphazard  than  in  L.  anatifera.  The  dorsal  part  of  the 
gastric  circulation  is  connected  to  the  inferior  gastric  vessels  by  a  varying  number  of  short 
vessels. 

In  P.  polymenis,  the  cirri  are  too  darkly  pigmented  to  observe  their  circulatory 
morphology,  but  the  opposite  holds  with  the  lepadids.  Figure  11  shows  the  circulation  of 
three  segments  of  a  ramus  from  L.  anatifera,  which  is  similar  to  that  of  the  other  species 
being  considered  here. 

The  afferent  circulation  in  a  ramus  of  a  cirrus  continues  distally  from  the  epineural 
sinus  and  is  in  close  contact  with  the  flexor  muscle.  In  each  segment  of  a  ramus,  the 
circumflexor  muscle  circulation  originates  from  the  afferent  vessel  and  surrounds  the 
flexor  muscle  in  a  sheet-like  sinus.  This  circulation  connects  to  the  efferent  circulation  of 
the  ramus  by  a  steadily  constricting  sinus.  There  may  be  a  valve  at  the  contact  point  with 
the  efferent  cirral  vessel.  The  efferent  cirral  vessel  progresses  down  the  outside  margin  of 
the  ramus  to  eventually  connect  with  the  peripheral-collecting  circulation. 

The  general  morphology  of  circulation  in  the  lepadid  filamentary  appendage  is 
similar  to  that  of  P.  polymerus  (Burnett,  1972).  There  are  two  vessels  (the  filamentary 
vessels)  on  opposite  sides  of  the  filamentary  appendage  that  parallel  the  main  axis  of  the 
appendage  (Fig.  12).  From  the  afferent  filamentary  vessel,  a  sheet-like  sinus  arises  on 
each  side  of  the  vessel,  and  each  extends  in  a  semicircle  around  the  filamentary 
appendage  to  connect  the  efferent  filamentary  vessel. 

The  Lepadidae  have  two  types  of  filamentary  appendages:  type  I  receives  hemolymph 
from  thq  afferent  circulation  to  the  cirri;  type  II  receives  blood  from  the  peripheral 
circulation  and  will  be  discussed  below.  Each  species  shows  a  different  arrangement  and 
number  of  type  I  filamentary  appendages.  Lepas  anatifera  has  a  type  I  appendage  at  the 
base  of  the  first  cirrus.  Lepas  pectinata  paciftca  also  has  one  in  the  same  location,  but  it 
is  reduced.  There  are  four  such  appendages  in  L.  fasicularis,  which  form  a  star  pattern 
where  they  originate  at  the  base  of  first  cirrus.  Conchoderma  virgatum  has  filamentary 
appendages  of  the  first  type  at  the  base  of  the  first,  third,  fourth  and  fifth  cirri. 
Interestingly,  C.  virgatum  has  an  additional  filamentary  appendage  at  the  base  of  the  first 
cirrus  that  receives  blood  from  the  efferent  circulation  of  that  cirrus  (a  type  II  filamentary 
appendage).  This  is  the  only  case  where  a  type  II  filamentary  appendage  occurs  on  a 
cirral  base. 

Peripheral-collecting  circulation.  (Figs.  13-18). — In  P.  polymenis,  I  described  three 
circulations  of  the  body:  distributive,  peripheral  and  collecting.  The  lepadids,  however, 
have  only  two  distinctive  circulations  of  the  body,  the  distributive  and  the  return.  In  order 
to  maintain  uniformity  in  nomenclature,  I  shall  call  the  return  circulation  of  the  lepadids 
the  peripheral-collecting  circulation.  Cannon  (1947)  also  described  the  return  circulation 


299 


CIRCUMFL.M.CIRC. 


0-5mm 


CEPHTR.MCIRC. 


TEST  PL 
PER  CIRC 


ROSr  VES 


GAS.VES. 


AD.  SCUT 


Figure  11.  Circulation  of  three  segments  of  a  ramus  in  L.  anatifera.  a.  View  of  the  posteriorly  facing  side  of  the 

ramus,  b.  anteriorly  facing  side. 

Figure  12.  A  filamentary  appendage  from  the  base  of  the  first  cirrus  in  L.  anatifera  showing  the  circulatory 
pattern.  This  arrangement  is  basic  to  all  filamentary  appendages  thus  far  observed,  a.  Longitudinal  view,  b. 
cross  section. 

Figure  13.  The  peripheral-collecting  circulation  of  L.  anatifera  in  an  illustration  similar  to  Fig.  7.  The  superior 
gastric  vessel  along  most  of  its  length  abutts  directly  against  the  inferior  gastric  vessel  (see  Fig.  7). 


300 


PRO  SIM 


POST  VAL 


0-5  mm 


Figure  14.  Variation  in  the  superior  gastric  vessel  of  L.  anatifera.  See  text  for  description. 

Figure  15.  The  peripheral-collecting  circulation  as  seen  from  the  left  side  of  the  body  of/,,  anatifera,  a.  the 
ln)rdcr  of  the  prosomal  sinus. 

Figure  16.  Close-up  of  the  peripheral-collecting  circulation  of  the  cephalic  region  in  L  fasicularis  showing  the 
thin  layer  of  the  peripheral  circulation  along  with  the  testicular  plexus. 

Figure  17.  The  rostral  vessel  and  its  associated  sinuses  in  L.  anatifera. 

Figure  18.  The  distribution  of  the  thin  layer  of  muscle  surrounding  the  prosomal  sinus  in  C.  virgatum. 


301 


in  Lithotrya  valentiana  as  essentially  a  peripheral-collecting  system. 

The  peripheral-collecting  circulation  in  the  Lepadidae  was  much  more  difficult  to 
trace  due  to  the  weakness  of  the  rostral  valve  and  a  consequent  filling  of  the  prosoma  with 
Microfil.  However,  in  a  few£.  anatifera  I  was  able  to  trace  this  circulation,  although  fine 
details  were  usually  obscurred. 

The  major  source  of  hemolymph  to  the  peripheral-collecting  circulation  comes  from 
the  efferent  cirral  vessels  and  the  return  flow  from  the  penis.  There  are  two  possible 
routes  for  this  hemolymph  after  it  leaves  the  cirri.  In  one  pathway,  vessels  from  the 
posterior  cirri  (5  and  6)  and  penis  join  to  form  the  paired  superior  gastric  vessels  (Fig. 
13).  These  vessels  occupy  a  ventrolateral  position  on  each  side  of  the  thoracic  gut  and 
decrease  in  caliber  from  their  posterior  origin.  The  superior  gastric  vessels  give  rise  to  a 
plexus  covering  almost  the  entire  thoracic  gut.  Essentially  the  same  pattern  is  shown  in  P. 
polvmerns  (Burnett,  1972).  In  one  individual  (of  five  £.  anatifera)  the  efferent  circulations 
from  the  left  side  of  cirri  2  through  6  contributed  to  the  superior  gastric  vessel  (which  in 
this  case  was  divided  into  two  vessels;  Fig.  14).  On  the  right  side,  the  morphology  was  as 
described  above. 

The  efferent  cirral  circulation  also  contributes  to  the  peripheral  circulation  of  the 
thoracic  region  (Fig.  15).  In  the  dorsal  part  of  the  thorax  the  peripheral  circulation  is 
derived  from  the  plexus  of  the  thoracic  gut  circulation.  The  two  peripheral  circulations 
combine  and  their  hemolymph  flows  anteriorly.  At  the  cephalic-thoracic  border,  a  vessel 
emanating  from  the  thoracic  peripheral  circulation  enters  the  cephalic  filamentary 
appendage. 

There  is  a  peripheral  connection  between  the  thoracic  and  cephalic  peripheral 
circulations,  but  as  this  area  is  remote  from  the  site  of  injection,  the  Microfil  rarely 
formed  a  continuous  band  from  the  thoracic  to  the  cephalic  peripheral-collecting 
circulations.  The  cephalic  filamentary  appendage  (a  type  II  filamentary  appendage), 
however,  serves  as  a  less  resistant  connection  between  the  two  halves  of  the  peripheral- 
collecting  circulations.  The  cephalic  transverse  muscle,  in  contrast  to  that  of  P. 
polymerus,  is  surrounded  by  hemolymph  from  the  cephalic  peripheral  circulation  (Fig. 
13). 

The  peripheral-collecting  system  of  the  prosoma  has  two  regions.  In  the  posterior 
part,  the  circulation  is  divided  into  a  plexus  that  surrounds  the  testes  (the  testicular 
plexus).  This  is  similar  to  the  peripheral-collecting  circulation  of  the  thoracic  region.  In 
L.  fasicularis,  this  plexus  is  more  grossly  constructed  than  in  L.  anatifera  and  in  both  a 
thin  peripheral  circulation  arises  from  connections  with  the  testicular  plexus. 

The  testicular  plexus  and  peripheral  circulation  connect  anteriorly  to  the  prosomal 
sinus  (Figs.  13,  17),  which  is  a  half  bowl-shaped  sinus  occupying  the  anterodorsal  part  of 
the  body.  This  sinus  is  completely  covered  by  a  thin  blanket  of  muscle  (Fig.  18)  that  is 
sandwiched  between  the  sinus  and  the  cuticle  of  the  prosoma. 

As  in  P.  polymerus,  the  prosomal  sinus  is  connected  by  a  pair  of  round  openings  (the 
prosomal  valves.  Fig.  17)  to  the  rostral  vessel.  The  morphology  of  the  region  appears  to  be 
as  in  P.  polymerus,  except  the  valve  flaps  of  the  rostral  vessel  do  not  appear  to  be  present 
(this  is  probably  due  to  vessel  distortion  so  frequently  observed  in  the  injections  of  the 
lepadids). 

The  rostral  sinus  (Figs.  13,  17)  also  has  a  morphology  similar  to  that  of  P. 
polymerus.  This  sinus  receives  blood  from  the  oral  cone,  the  adductor  scutorum,  and 
perhaps  the  prosomal  sinus  via  the  rostral  vessel. 

By  shining  a  light  through  L.  fasicularis,  I  observed  that  the  cuticle  between  the 
adductor  scutorum  and  the  oral  cone  pulsates  every  3-4  seconds  at  22°C.  Such 
movements  probably  result  from  the  contraction  of  the  rostral  sinus  muscles.  However, 
for  the  rostral  sinus  to  operate  as  a  pump,  a  valve  should  be  located  between  the  rostral 
sinus  and  the  anterior  oral  cone;  none  was  found. 

It  appears  that  the  rostral  sinus  pumps  hemolymph  through  the  rostral  vessel  and 
into  the  peduncle.  The  direction  of  flow  is  deduced  from  the  position  of  the  valves,  partial 
injections  of  Microfil,  and  from  my  studies  with  P.  polymerus  (Burnett,  1972). 


302 

0(pr.) 


CARDVAL  ...„.-.,..- ...™^..,. 

ROSTVAL. 
ROSTSIN.      \  ^PRQVAL.(pr) 


Figure  19.  Comparison  of  the  rostral  vessel  (right)  with  the  heart  of  Calanus  finmarchicus  (left  —  redrawn  from 

Lowe.  1935). 

The  structure  of  the  vessel  wall  of  the  cephalic  gut  is  like  that  of  the  midsagittal 
vessels  in  P.  polymenis.  Light  microscopy  shows  an  intima  containing  large  branching 
fibers  that  become  more  diffuse  a  short  distance  from  the  intima.  The  spaces  with  no 
apparent  circulation  ventral  to  the  gut  are  occupied  by  the  seminal  vesicles.  These  organs 
appear  to  have  little  circulation  associated  with  them. 

The  distributive  circulation  of  the  four  species  differs  in  vessel  number  and  caliber: 
the  smaller  species  {L.  fasicularis  and  L.  pectinata  pacifica)  have  fewer,  larger  caliber 
vessels  than  the  larger  species,  suggesting  that  small  barnacles  have  less  complex 
circulation.  Conchodenna  virgatum  and  L.  fasicularis  are  of  similar  size,  but  the  former 
has  more  complex  circulation. 

The  blood  pump. — The  location  of  the  hemolymph  pump  in  barnacles  has  been  in 
dispute  (Fyhn  et  al.,  1973).  From  a  study  of  serial  sections  of  Lithotrya  valentiana. 
Cannon  (1947),  placed  it  between  the  adductor  scutorum  and  the  base  of  the  oral  cone 
(the  rostral  sinus).  He  called  this  sinus  the  "blood  pump"  instead  of  a  heart  because  the 
muscles  are  located  within  the  sinus  rather  than  encircling  it.  In  Balanus  balanoides, 
Gutmann  (1960)  argued  that  circulation  takes  place  as  a  result  of  muscular  activity  or 
during  cirral  extension  and  retraction:  during  periods  of  inactivity  contraction  of  muscles 
in  the  prosoma  propel  the  blood.  However,  in  inactive  barnacles  Blatchford  (1970) 
observed  movements  in  the  region  of  the  rostral  sinus  that  he  ascribed  to  circulatory 
movements. 

I  found  by  my  observations  on  L.  fasicularis  that  the  rostral  sinus  probably  acts  as  a 
blood  pump  for  this  species  and  for  the  other  lepadid  species. 

DISCUSSION 

Newman  et  al.  (1969)  postulated  that  the  thoracican  Cirripedial  were  derived  from  an 
ascothoracican-like  maxillopodian  ancestor  and  therefore  are  closely  allied  to  the 
Copepoda.  I  noted  (1972)  the  similarity  of  the  heart  of  the  copepod  Calanus  finmarchicus 
to  the  rostral  vessel  of  P.  polymerus  (Fig.  19).  The  lepadid  rostral  vessel  with  its 
connection  to  the  prosomal  sinus  has  the  same  arrangement  as  in  P.  polymerus.  The 
argument  supporting  the  rostral  vessel  as  being  a  vestigial  heart  is  1)  the  rostral  vessel  is 
essentially  in  a  dorsal  position  and  is  properly  oriented,  2)  the  openings  into  the  rostral 
vessel  correspond  to  the  positions  of  the  ostia  in  the  copepod  Calanus  finmarchicus 
(Lowe,  1935),  and  3)  the  rostral  valve  is  homologous  to  the  cardioarterial  valve  in  the 
copepod  heart. 

Since  the  rostral  vessel  lacks  musculature,  I  infer  that  during  the  evolution  of  the 
Cirripedia,  heart  function  was  shifted  from  the  heart  (rostral  vessel)  to  the  rostral  sinus  (an 
original  part  of  the  pericardial  sinus).  Why  would  there  be  a  shift  of  heart  function  in  the 
barnacles?  In  pedunculate  barnacles  contraction  of  the  peduncle  forces  a  large  pulse  of 
hemolymph  into  the  body.  Apparently,  the  only  large  sinus  positioned  to  receive  and  store 
this  extra  hemolymph  is  the  prosomal  sinus.  The  prosomal  sinus  probably  not  only  acts  as 
the  main  venous  sinus  for  the  body,  but  also  is  involved  in  maintaining  equilibrium 
between  the  peduncle  and  the  body. 

In  the  primitive  thoracic  cirriped,  the  heart  was  suspended  in  a  sinus  that  was 
subject  to  increasingly  high  pressures  as  the  peduncle  became  more  dynamic.  The  net 


303 


effect  of  these  increasingly  high  pressures  would  be  collapse  of  the  heart.  A  shift  of  blood 
pumping  from  the  heart  to  a  sinus  where  the  muscles  are  intrinsically  located  would  solve 
the  problem  of  collapse. 

The  reason  for  splitting  the  primitive  pericardial  sinus  into  the  prosomal  and  rostral 
sinuses  is  difficult  to  postulate.  Perhaps  this  separation  was  present  prior  to  the 
development  of  a  peduncle  and  the  consequent  loss  of  heart  musculature.  Such  a 
separation  would  be  necessary  if  part  of  the  pericardial  sinus  was  to  act  as  a  reservoir  and 
part  as  a  pumping  organ. 

Regardless  of  the  state  of  the  peduncle,  it  must  always  receive  oxygenated  blood. 
This  is  accomplished  by  a  continuous  beating  of  the  rostral  sinus  in  which,  no  matter 
what  the  length  of  the  peduncle,  a  constant  volume  of  blood  is  pumped  into  the  peduncle 
from  the  rostral  sinus.  Peduncular  extension  is  probably  mostly  mediated  by  hemolymph 
from  the  prosomal  sinus  that  was  originally  squeezed  out  of  this  sinus  by  contraction  of 
the  prosomal  muscles. 

ACKNOWLEDGEMENTS 

I  wish  to  thank  Dr.  Robert  R.  Hessler  of  Scripps  Institution  of  Oceanography  for  his  review  of  the 
manuscript  and  helpful  suggestions  during  this  study.  I  also  wish  to  thank  Dr.  Carl  Hubbs  of  Scripps  Institution 
of  Oceanography  for  supplying  the  Conchoderma  virgatum,  and  Dr.  William  Newman  of  Scripps  Institution  of 
Oceanography  for  his  review  of  the  manuscript. 

LITERATURE  CITED 

Blatchford,  J.G. 

1970.   Possible  circulatory  mechanism  in  an  operculate  barnacle.  Comp.  Biochem.  Physiol.,  34:911-915. 
Burnett,  B.R. 

1972.  Aspects  of  the  circulatory  system  oi  Pollicipes  polymerus  J.B.  Sowerby  (Cirripedia:  Thoracica).  J. 
Morph.,  136:79-107. 

Cannon,  H.G. 

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Scripps  Institution  of  Oceanography.  P.O.  Box  1529,  La  Jolla,  California  92037. 


APPENDIX 
ABBREVIATIONS 

AD.  SCUT.  adductor  scutorum 

AF.  AD.  SCUT.  afferent  of  the  adductor  scutorum 

AF.  CIR.  VES.  afferent  cirral  vessel 

AF.  FIL.  VES.  afferent  filamentary  vessel 

AF.  MAN.  CIRC.  afferent  mantle  circulation 

AF.  MD.  afferent  mandible  circulation 

AF.  OR.  C.  VES.  afferent  oral  cone  vessel 

AN.  anus 

ANT.  OR.  C.  VES.  anterior  oral  cone  vessel 


304 


CAR. 

CAR.  VES. 

CARD.  VAL. 

CEPH.  FIL.  AP. 

CEPH.  PER.  CIRC. 

CEPH.  TR.  M. 

CEPH.  TR.  M.  CIRC. 

CIR.  1 

CIRCUM.  FL.  M.  CIRC. 

CON.  VES. 


carina 

carinal  vessel 

cardioarterial  valve 

cephalic  filamentary  appendage 

cephalic  peripheral  circulation 

cephalic  transverse  muscle 

cephalic  transverse  muscle  circulation 

cirrus  1 

circumflexor  muscle  circulation 

connective  muscle 


EF.  CIR.  VES. 
EF.  FIL.  VES. 
EPI.  SIN. 
EX.  CUT. 


efferent  cirral  vessel 
efferent  filamentary  vessel 
epineural  sinus 
exterior  cuticle 


FIL.  AP. 


filamentary  appendage 


G. 

gut 

CAST.  GL. 

gastric  gland 

GAST.  PL. 

gastric  plexus 

INF.  GAS.  VES. 

inferior  gastric  vessel 

IV.  CIRC. 

intervalve  circulation 

M. 

mantle 

MAX.  GL.  PL. 

maxillary  gland  plexus 

O. 

ostium 

O.  (pr) 

ostium  (paired) 

OR.  C. 

oral  cone 

OV.  FR. 

ovigerous  frenum 

P. 

penis 

PED. 

peduncle 

PED.  VES. 

peduncular  vessel 

PER.  CIRC. 

peripheral  circulation 

POST.  INF.  GAS.  VES. 

posterior  inferior  gastric  vessel 

POST.  OR.  C.  VES. 

posterior  oral  cone  vessel 

POST.  SIN. 

posterior  sinus 

PRO.  M. 

prosomal  muscle 

PRO.  SIN. 

prosomal  sinus 

PRO.  VAL. 

prosomal  valve 

PRO.  VAL.  (pr) 

prosomal  valve  (paired) 

ROST.  M. 

rostral  muscle 

ROST.  SIN. 

rostral  sinus 

ROST.  VAL. 

rostral  valve 

ROST.  VES. 

rostral  vessel 

SCUT.  PL. 

scutal  plexus 

SCUT.  SIN. 

scutal  sinus 

SCUT.  VAL. 

scutal  valve 

SCUT.  VES 

scutal  vessel 

SUB.  VES. 

subintestinal  vessel 

SUP.  GAS.  VES. 

superior  gastric  vessel 

SUR.  CIRC. 

surface  circulation 

TER.  PL. 

tergal  plexus 

TEST.  PL. 

testicular  plexus 

THOR.  FIL.  AP. 

thoracic  filamentary  appendage 

THOR.  PER.  CIRC. 

thoracic  peripheral  cjrculation 

THOR.  TR.  M. 

thoracic  transverse  muscle 

VAL. 


valve 


79(J5    uB3 


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MAR  9  - 1984 

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