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HARVARD    UNIVERSITY 

Library  of  the 

Museum  of 

Comparative  Zoology 


TRANSACTIONS 

OF  THE 

SAN  DIEGO  SOCIETY  OF  NATURAL  HISTORY 

VOLUME  20 
1982-1985 

CONTENTS 

1.  Late  Wisconsinan  and  Holocene  fauna  from  Smith  Creek  Canyon,  Snake 
Range.  Nevada.  By  Jim  I.  Mead.  Robert  S.  Thompson  and  Thomas  R. 

Van  De\ender.  6  August  1982   1-26 

2.  Fossil  decapod  crustaceans  from  the  Lower  Cretaceous.  Glen  Rose  Limestone 

of  Central  Texas.  By  Gale  A.  Bishop,  25  January  1983 27-55 

3.  A  new  subspecies  of  Euphyes  vestris  (Boisduval)  from  southern  California 
(Lepidoptera:  Hesperiidae).  By  John  W.  Brown  and  William  W.  McGuire. 

24  June  1983 ' 57-68 

4.  Two  new  iodeid  isopods  from  Baja  California  and  the  Gulf  of  California 
(Mexico)  and  an  analysis  of  the  evolutionary  history  of  the  genus  Colidotea 
(Crustacea:  Isopoda:  Idoteidae).  By  Richard  C.  Brusca.  24  June  1983  ....         69-79 

5.  Descriptions  of  five  new  muricacean  gastropods  and  comments  on  two 
additional  species,  in  the  Families  Muricidae  and  Coralliophilidae: 
(Mollusca).    By   Anthony   D'Attilio  and   Barbara   W.    Myers,    18  January 

1984 '.         81-94 

6.  The  fossil  Leptostracan  Rhahdouraea  ^?A7/r;  (Malzahn.  1958).  By  Frederick 

R.  Schram  and  Eric  Mabahn.  18  January  1984 95-98 

7.  Phylogen\,  evolution  and  biogeography  of  the  marine  isopod  Subfamily 
Idoteinae  (Crustacea:  Isopoda:  Idoteidae).  By  Richard  C.  Brusca,  18 
January  1984 99-134 

8.  Rhamdia  reddelli,  new  species,  the  first  blind  pimelodid  catfish  from  Middle 
America,  with  a  key  to  the  Mexican  species.  B\  Robert  Rush  Miller.  18 
January  1984 '. ' 135-144 

9.  A  complete  specimen  of  Peachella  brevispina  Palmer  —  an  unusual  olenellid 
trilobite  (Arthropoda:  Olenellida)  from  the  lower  Cambrian  of  California. 

By  James  H.  Stitt  and  R.  L.  Clark.  20  June  1984 145-150 

10.  Type  specimens  of  amphibians  and  reptiles  in  the  San  Diego  Natural  History 
Museum.  By  Gregory  K.  Pregill  and  James  E.  Berrian.  20  June  1984 151-164 

11.  Imocaris  tuhenulata,  n.  gen.,  n.  sp.  (Crustacea:  Decapoda)  from  the  upper 
Mississippian    Imo    Formation.   Arkansas.    By   Frederick    R.   Schram  and 

Royal  H.  Mapes.  20  June  1984 165-168 

12.  New  material  of  Hydrodamalis  custae  (Mammalia:  Dugongidae)  from  the 
Miocene  and  Pliocene  of  San  Diego  County,  California.  By  Daryl  P. 
Domning  and  Thomas  A.  Demere,  20  November  1984 169-188 

13.  Fossil  Syncarida.  By  Frederick  R.  Schram,  20  November  1984 189-246 

14.  The  late  Wisconsinan  vertebrate  fauna  from  Deadman  Cave,  southern 
Arizona.  Bv  Jim  1.  Mead.  Edward  L.  Roth.  Thomas  R.  Van  De\enderand 

Da\id  W.  Steadman.  20  November  1984 247-276 

15.  A   Pliocene  flora   from   Chula   Vista,  San   Diego  County,  California.   By 

Daniel  I.  A.xclrod  and  J  homas  A.  Demere,  20  November  1984 277-300 

16.  Relationships  within  Eumalacostracan  Crustacea.  By  Frederick  R.  Schram. 

20  Noxcmber  1984   301-312 

17.  Historv  and  status  of  the  avifauna  of  Isla  Guadalupe.  Mexico.  By  Joseph  R. 

Jehl,  Jr.  and  William  I.  Everett,  30  January  1985 313-336 


i^ 


TRANSACTIONS 
OF  THE  SAN  DIEGO 
SOCIETY  OF 
NATURAL  HISTORY 


Volume  20  Number  1  pp.  1-26     6  August  1982 


Late  Wisconsinan  and  Holocene  Fauna  from  Smith  Creek  Canyon, 
Snake  Range,  Nevada 

Jim  I.  Mead,  Robert  S.  Thompson,  and  Thomas  R.  Van  Devender 

Laboratory  of  Paleoenvironmental  Studies,  Department  of  Geosciences, 
University  of  Arizona,  Tucson  85721 

Abstract.  During  the  late  Pleistocene,  montane  glaciers  in  the  Snake  Range,  eastern  Nevada 
reached  an  elevation  as  low  as  2900  m  and  pluvial  Lake  Bonneville  rose  to  approximately  1580  m. 
only  130  m  below  the  entrance  of  the  east-facing  Smith  Creek  Canyon.  It  is  not  known  whether  the 
two  events  coincided.  Packrat  midden  macrofossils  indicate  that  bristlecone  pine  (Piiuis  lont^'acva). 
limber  pine  (P.  flexilis),  and  other  subalpine  taxa  dominated  the  plant  communities  in  Smith  Creek 
Canyon  through  the  late  Wisconsinan.  We  report  here  2  fish,  4  anurans,  9  lizards,  8  snakes,  and  15 
small  mammals  recovered  from  15  packrat  middens  and  a  pollen  profile  from  cave  fill.  This  assemblage 
adds  15  amphibians  and  reptiles  and  7  mammals  to  the  approximately  46  terrestrial  animals  previously 
known  from  the  late  Pleistocene  and  early  Holocene  of  the  canyon.  Dung  pellets  of  the  locally  extir- 
pated pika  (Ochotona  cf.  princeps)  were  found  in  five  packrat  middens.  A  single  tooth  of  the  heather 
vole  (Phenacomys  cf.  intermedius)  from  Smith  Creek  Cave  is  the  first  late  Pleistocene  record  for  this 
genus  in  the  Great  Basin.  We  review  and  update  the  late  Pleistocene  and  Holocene  fauna  from  4  caves 
and  2  shelters  in  Smith  Creek  Canyon. 

Introduction 

The  Snake  Range  of  White  Pine  County,  eastern  Nevada,  is  a  north-south  trending 
mountain  80  km  long.  Smith  Creek  Canyon,  a  deep  canyon  on  the  eastern  face  of  this 
Great  Basin  range,  opens  onto  the  Lake  Bonneville  playa  in  the  Snake  Valley  of  Utah. 
During  the  late  Pleistocene,  montane  glaciers  reached  elevations  as  low  as  2900  m  in 
the  Snake  Range  (Drewes  1958),  and  pluvial  Lake  Bonneville  rose  to  a  level  of  ap- 
proximately 1580  m,  only  130  m  below  the  entrance  of  Smith  Creek  Canyon.  If  the 
late  Wisconsinan  glacial  maximum  coincided  with  the  high  stand  of  Lake  Bonneville, 
biotic  communities  in  this  canyon  would  have  been  restricted  to  an  elevational  range 
of  less  than  1310  m  (Fig.  I). 

Fossil  localities. — Late  Pleistocene  vertebrate  fossils  from  Smith  Creek  Canyon, 
specifically  Smith  Creek  Cave,  were  reported  by  M.  Harrington  { 1934)  from  the  South- 
west Museum  and  by  others  since.  In  1955,  T.  E.  Downs  and  associates  from  the 
Natural  History  Museum  of  Los  Angeles  County  (LACM),  California,  screened  some 
of  the  cave  sediments.  Field  notes  of  the  LACM  indicate  that  the  cave  contained  little 
or  no  stratification  (Brattstrom  1976).  Howard  (1935,  1952)  described  a  new  species  of 
extinct  eagle  (Spizaetiis  willctti)  and  a  teratorn  (Teratornis  incrcilihilis)  and  listed  the 
Smith  Creek  Cave  avifauna,  unfortunately  giving  the  stratigraphic  associations  only  as 
from  the  older  deposits  of  the  lower  levels.  Stock  (1936)  described  a  new  species  of 
extinct  mountain  goat  (Oreamnos  harringtoni),  but  again  did  not  discuss  placement 
within  the  stratigraphic  sequence.  Brattstrom  (1958,  1976)  reported  amphibians  and 
reptiles  from  the  cave  sediments  and  Goodrich  (1965)  updated  the  list  of  the  entire 
fauna  (amphibians,  reptiles,  birds,  and  mammals);  again  the  stratigraphy  and  temporal 
associations  were  only  scantily  discussed. 

Bryan  ( I979«)  made  further  excavations  in  the  cave  in  search  of  evidence  for  Early 
Man  and  for  the  first  time  described  the  stratigraphy  within  a  portion  of  the  cave. 


Figure  1 .     Map  of  Nevada  and  bordering  states  showing  Smith  Creek  Canyon  ( I )  and  Gatecliff  Shelter  (2) 
along  with  Great  Salt  Lake  (black)  and  Lake  Bonneville  (stippled)  at  the  1580  m  elevation. 


Miller  (1979)  identified  the  mammalian  fauna  recovered  during  Bryan's  excavations, 
providing  another  updated  listing  of  the  fauna;  some  stratigraphic  associations  were 
given.  Both  Bryan  {\979a)  and  Miller  (1979)  emphasized  the  deposits  and  fauna  asso- 
ciated with  Man,  those  units  dating  less  than  11500  B.P.  Thus  no  comprehensive 
stratigraphic  analysis  of  the  fauna  has  been  published. 

In  1977  and  1978  we  visited  Smith  Creek  Canyon  to  study  fossil  packrat  (Neotoma) 
middens  and  to  collect  a  pollen  profile  from  the  Smith  Creek  Cave  sediments.  Plant 
remains  from  some  of  these  middens  are  discussed  in  a  previous  report  (Thompson 
1979),  and  other  assemblages  are  still  being  analyzed  (RST).  The  packrat  midden  fossils 
document  that  bristlecone  pine  {Pinus  longaeva),  limber  pine  {P.  fle.xilis),  and  other 
subalpine  taxa  dominated  the  plant  communities  in  Smith  Creek  Canyon  throughout 
the  late  Wisconsinan.  The  Smith  Creek  Cave  pollen  profile  provided  little  paleoenvi- 
ronmental  information  and  could  not  be  dated  directly.  In  this  report  we  describe  the 
fish,  amphibians,  reptiles,  and  mammals  recovered  from  15  packrat  middens  from 
Smith  Creek  Canyon  and  in  association  with  the  pollen  profile,  and  the  herpetofauna 
recovered  by  Bryan  (\919a,  Miller  1979)  in  his  excavation  in  Smith  Creek  Cave.  We 
also  review  the  entire  local  fauna  recovered  from  all  archaeological  and  paleontological 


investigations  in  Smith  Creek  Canyon  and  attempt  to  place  these  records  in  the  correct 
paleoenvironmental  and  chronological  context. 

Local  setting. — The  lower  concourse  of  Smith  Creek  Canyon  is  a  steep-walled 
east-west  trending  canyon  incised  into  Paleozoic  limestone.  In  this  part  of  the  canyon 
there  are  numerous  caves,  including  those  described  below  in  which  we  have  found 
our  fossil  materials.  The  upper  reaches  of  Smith  Creek  Canyon  are  lined  with  other 
Paleozoic  sedimentary  rocks  and  Mesozoic  intrusives.  We  have  not  located  any  caves 
or  fossil  packrat  middens  in  this  area.  The  intermittent  waters  of  Smith  Creek  are 
primarily  derived  from  the  elevated  plateau  surrounding  Mount  Moriah  (3673  m). 

Although  the  vegetation  in  Smith  Creek  Canyon  is  broadly  divided  into  elevational 
zones,  the  major  slope  and  aspect  contrasts  between  the  north  and  south  facing  slopes 
create  numerous  microenvironments  for  plants  of  higher  and  lower  elevations.  At  the 
canyon  entrance  the  plant  community  is  dominated  by  shadscale  (A triplex  confertifolia) 
and  other  xerophytes,  including  spiny  hopsage  {Grayia  spinosa),  greasewood  (Sarcoh- 
atus  venniciilatiis),  rabbitbush  {Chrysothamniis  nanseosns).  Mormon  tea  (Ephedra 
nevadensis),  horsebrush  (Tetradyinia  axillaris),  Harriman  yucca  (Yucca  harriinaniae), 
and  bud-sage  (Artemisia  spinescens).  These  same  taxa  are  dominant  in  the  Snake 
Valley  to  the  east  and  also  are  common  on  xeric  slopes  throughout  Smith  Creek  Can- 
yon. 

Sagebrush  (Artemisia  tridentata,  A.  nova)  occurs  both  in  nearly  pure  stands  in  the 
canyon  bottom  and  as  a  common  understory  element  in  the  pinyon-juniper  woodlands 
and  forested  communities.  Utah  juniper  (Juniperus  osteosperma)  and  single  needle 
pinyon  (Finns  monophylla)  occur  throughout  the  limestone  walled  part  of  the  canyon 
and  form  denser  stands  on  the  north  facing  slopes  and  on  alluvium.  Little  leaf  mountain 
mahogany  (Cercocarpus  intricatus),  greasebush  (Forsellesia  nevadensis),  skunkbush 
(Rhus  trilohata),  and  joint-fir  (Ephedra  viridis)  are  common  in  the  pinyon-juniper 
woodland  and  are  often  dominant  on  xeric  slopes  and  on  limestone  substrates. 

Montane  and  subalpine  conifers,  including  white  fir  (Abies  concolor),  Douglas  fir 
(Pseudotsuga  menziesii),  Engelmann  spruce  (Picea  engelmannii),  ponderosa  pine 
(Pinus  ponderosa),  limber  pine,  and  bristlecone  pine,  are  present  at  relatively  low 
elevations  in  mesic  niches  on  the  north  facing  slope  and  along  the  upper  concourse  of 
Smith  Creek.  Other  common  riparian  plants  include  narrowleaf  cottonwood  (Populus 
angustifolia),  willows  (Salix  spp.),  water  birch  (Betula  occidentalis),  chokecherry 
(Pruniis  virginiana).  Rocky  Mountain  maple  (Acer  glabrum).  Rocky  Mountain  juniper 
(Juniperus  scopulorum),  and  wild  rose  (Rosa  woodsii).  The  higher  elevations  of  the 
northern  Snake  Range  support  groves  of  quaking  aspen  (Populus  tremuloides)  and 
stands  of  bristlecone  pine  and  limber  pine  on  rocky  outcrops. 

Although  the  fossil  localities  we  investigated  occur  in  a  narrow  elevational  range 
(1860  m  to  2060  m),  the  variations  in  slope  and  aspect  at  these  sites  place  them  in 
different  environmental  settings  (Table  1).  Smith  Creek  Cave  (1950  m  elev.)  and  Ladder 
Cave  (2060  m  elev.)  are  on  a  steep  south  facing  slope  at  the  entrance  to  the  canyon 
and  are  surrounded  by  a  xeric  pinyon-juniper  woodland  with  an  abundance  of  xero- 
phytic  plants  (see  Thompson  1979).  Streamview  Shelter  is  on  a  protected  north  facing 
slope  near  the  canyon  bottom  (1860  m  elev.),  and  Council  Hall  Cave  is  higher  (2040 
m  elev.)  on  the  same  slope.  These  two  sites  are  in  a  relatively  dense  pinyon-juniper 
woodland,  with  fewer  xerophytes  than  on  the  opposing  slope.  Montane  and  subalpine 
conifers  occur  in  protected  niches  near  both  Council  Hall  Cave  and  Streamview  Shel- 
ter. Amy's  Shelter  and  the  Kachina  Cave  are  in  the  bottom  of  the  canyon  by  the  creek. 
These  two  sites  probably  were  buried  by  alluvium  during  the  last  full-glacial,  only  to 
be  exposed  during  the  latest  phases  of  the  late  glacial  or  early  Holocene. 

Methods 

The  Smith  Creek  Canyon  localities  included  within  this  report  are  Smith  Creek, 
Ladder,  Council  Hall,  and  Kachina  caves.  Amy's  Shcilcr  and  the  packrat  midden 
rockshelter  we  named  Streamview. 


Table  i.  Paleontological  and  archaeological  sites  in  Smith  Creek  Canyon,  Snake  Range,  and  the  adjacent 
Snake  Valley,  Nevada,  that  are  discussed  in  text. 


Locality 

Slope 

Eleva- 
tion 

Type  of  deposit 

Age  range 
(yrB.P.) 

Present  status 

Smith  Creek 
Cave 

South- 
facing 

1950  m 

Archaeological 
and  non- 
archaeological 
cave 

sediments;  4 
packrat 
middens 

^12  500  to  1000 

Relatively  xeric  site,  sparse 
pinyon-juniper  woodland, 
active  talus  formation,  high 
above  canyon  bottom 

Ladder  Cave 

South- 
facing 

2060  m 

6  packrat 
middens 

27  000  to  1 1  000 

Relatively  xeric  site,  sparse 
pinyon-juniper  woodland, 
active  talus  formation,  high 
above  canyon  bottom 

Amy's 
Rockshelter 

North- 
facing 

1740  m 

Archaeological 
cave  sediments 

5000  to  1500 

In  canyon  bottom  near 
entrance,  relatively  arid, 
scattered  juniper  with 
sagebrush-shadscale  shrub 

Streamview 
Rockshelter 

North- 
facing 

1860  m 

3  packrat 
middens 

17  000  to  6500 

Ca.  15  m  above  canyon 
bottom,  pinyon-juniper 

Kachina  Cave 

North- 
facing 

1770  m 

Archaeological 
cave  sediments 

ca.  4500  to  present 

Along  streamside  of 

intermittent  Smith  Creek, 
pinyon-juniper  and  riparian 
vegetation 

Council  Hall 
Cave 

North- 
facing 

2040  m 

Cave  sediments; 
2  packrat 
middens 

24  000  to  4000 

High  above  cayon  bottom, 
relatively  mesic  site, 
pinyon-juniper  with 
scattered  subalpine 
conifers,  little  active  talus 
formation 

Garrison 

North- 
facing 

1640  m 

Small  overhang; 
2  packrat 
middens 

13  500  to  12  000 

Small  rock  outcrop  in  the 
middle  of  Snake  Valley, 
desertscrub,  little  active 
talus 

Packrat  middens. — All  fossil  packrat  middens  in  this  report  were  well-indurated 
with  urine.  To  insure  that  stratigraphic  units  of  different  ages  were  not  mixed  during 
sampling,  each  midden  was  examined  carefully  prior  to  collection.  Approximately  two 
kilograms  were  removed  from  each  stratigraphic  unit  and  packaged  in  the  field.  In  the 
laboratory  approximately  one  kilogram  of  each  unit  was  soaked  in  water  until  the 
cementing  urine  dissolved.  The  disaggregated  samples  were  then  washed  through  20- 
mesh  screens  and  oven  dried.  The  fossils  were  hand-sorted,  and  after  identification, 
plant  remains  were  selected  for  radiocarbon  dating  and  pretreated  with  10%  HCI  in  an 
ultrasonic  cleaner.  Radiocarbon  dates  from  the  packrat  middens  and  the  cave  sedi- 
ments (discussed  below)  are  presented  in  Table  2. 

Four  of  the  5  packrat  middens  recovered  from  Smith  Creek  Cave  will  be  discussed. 
Radiocarbon  dating  of  these  middens  indicates  a  time  range  from  1 1  650  ±  280  to 
13  340  ±  430  yr  B.P.  (years  before  present.  Table  2).  Ladder  Cave,  a  much  smaller 
cave,  is  located  immediately  above  Smith  Creek  Cave.  We  discuss  6  packrat  middens 
from  this  cave,  dating  from  1 1  080  ±  115  to  27  280  ±  970  yr  B.P.  (Table  2).  Two  packrat 
middens  from  Council  Hall  Cave  (Table  2)  provided  faunal  remains  of  middle  Holocene 
age  (4220  ±  60  to  6120  ±  80  yr  B.P.).  Three  middens  reported  from  the  Streamview 
locality  date  from  the  middle  Holocene  (6490  ±  190  yr  B.P.),  late  glacial  (1 1  010  ±  400 
yr  B.P.),  and  the  end  of  the  last  full-glacial  (17  350  ±  435  yr  B.P.).  Amy's  Shelter  is 
in  the  bottom  of  the  canyon  across  the  canyon  from  Smith  Creek  Cave.  This  site  and 
Kachina  Cave  are  included  here  to  add  the  results  of  Miller's  (1979)  faunal  study. 

Smith   Creek   Cave  stratigraphy.— The   stratigraphy  currently  known  from  the 


Table  2.  Radiocarbon  dated  packrat  middens  and  cave  sediments  from  Smith  Creek  Canyon,  Snake  Range, 
Nevada  (Thompson  1979,  Thompson  and  Mead  1982).  Associated  plant  material:  A)  Dung  and/or 
unidentified  plants;  B)  Ephaha  vindis\  C)  Juniperns  communis:  D)  J.  osteosperma;  E)  Picea  enfielmannii; 
F)  Pinu.s  fli-.xilis;  G)  P.  lonf^iwva:  H)  P.  monophvlla:  I)  Artemisia  spp.;  J)  Afriplex  confertifolia;  K) 
Cercocarpus  intricatus;  L)  Chamuchatiaria  millijoliiim:  M)  Chrysothumnus  sp.;  N)  Foresellesiu  neva- 
densis;  O)  Ribes  montif^eniim;  P)  Symphoricarpos  sp.;  Q)  Rhus  triolhala. 


Radiocarbon  age 

Material 

Plant 

Locality 

Lab  no. 

(yr  B.P.) 

dated 

associates 

Council  Hall  Cave  (CHC) 

la 

Wk-157 

6120 

+ 

80 

D 

BKNHQ 

lb 

Wk-158 

4220 

-t- 

60 

D 

BHKMQ 

Ladder  Cave  (LC) 

1 

Wk-151 

11  200 

± 

200 

G 

DIKN 

2a 

WK-152 

27  280 

-+- 

970 

P 

BDINO 

2b 

A-2092 

17  960 

-H 

110 

A 

GO 

3 

Wk-154 

13  230 

-+- 

110 

G 

DILNO 

4 

Wk-155 

12  100 

± 

150 

G 

IKNO 

6 

Wk-156 

11080 

-+- 

115 

G 

BDIKN 

Smith  Creek  Cave  (SCC) 

1 

Gx-5861 

11660 

-+- 

245 

A 

FGKP 

3 

Gx-5862 

1 1  650 

^ 

280 

G 

MNP 

4 

Gx-5863 

12  235 

H- 

395 

G 

CIKN 

5 

A-2094 

13  340 

•+- 

430 

A 

GILM 

(Reddish-brown  Silt:  SCC  Sed.) 

Tx-1639 

28  650 

^: 

760 

(see 

text) 

Streamview  Shelter  (STV) 

1 

A-2095 

11010 

^ 

400 

FG 

INP 

2 

Gx-5866 

17  350 

-+■ 

435 

G 

CEJKN 

3 

Gx-5867 

6490 

-+- 

190 

D 

BCGJK 

northwestern  sections  of  Smith  Creek  Cave  was  estabHshed  from  the  excavation  of 
Test  Pits  2  and  3  (Bryan  \979a).  The  stratigraphic  units  established  by  Bryan  will  be 
followed  in  this  report.  Bryan  recognized  3  stratigraphic  "zones"  in  the  rear  of  Smith 
Creek  Cave,  the  lowest  unit  being  the  Cemented  White  Silt  Zone.  In  the  northwestern 
section  of  the  cave,  this  unit  was  eroded  away  before  the  deposition  of  the  second 
stratigraphic  zone — the  Reddish-brown  Silt  Zone.  Bryan  noted  that  the  erosional  dis- 
conformity  between  the  Cemented  White  Silt  Zone  and  the  Reddish-brown  Silt  unit  is 
near  vertical.  The  reddish  silt  of  the  latter  unit  is  believed  to  be  of  probable  eolian 
deposition,  although  it  is  just  as  likely  that  water  from  a  nearby  ceiling  conduit  may 
have  caused  the  erosion  of  the  first  stratigraphic  unit  and  at  least  a  partial  deposition 
of  the  Reddish-brown  Silt.  This  latter  unit  is  very  fossiliferous  (partially  a  raptor  ac- 
cumulation) and  is  stained  reddish-brown  by  the  surrounding  silt  (dry  7/6  lOYR.  yellow; 
damp  5/8  7.5YR,  strong  brown,  Munsell  color;  our  observations).  No  organic  remains, 
including  charcoal,  were  observed  in  the  Reddish-brown  Silt  by  Bryan  (\979a)  or  us. 
Permineralization  has  occurred  with  all  of  the  red-stained  bones  (Miller  1979).  Younger 
contaminating  bones  are  white,  not  mineralized,  and  easily  spotted. 

Bryan  ( 1979a)  reported  that  "a  sample  of  unidentified  red-stained  bone  scrap  [from 
Test  Pit  2]  yielded  a  collagen  date  of  28,650  ±  760  years  B.P.  (Tx- 1639)."  A  discrepancy 
arises  in  that  Valastro  (1977)  reported  that  the  Tx-1639  '^C  date  is  from  a  charcoal 
sample  from  Test  Pits  2  and  5;  Test  Pit  5  is  at  the  mouth  of  the  cave.  We  did  not  find 
charcoal  in  the  unit  in  Test  Pit  2.  Because  the  red-stained  bones  in  the  Reddish-brown 
Silt  in  Test  Pit  2  is  permineralized  with  the  loss  of  most  bulk  organic  constituents 
(Miller  1979),  we  assume  that  the  radiocarbon  date  from  the  unit  is  at  best  a  very  rough 
estimate  of  its  true  age  and  that  the  material  dated  was  probably  bone  scraps,  not 
charcoal.  Further  radiocarbon  dating  of  this  unit  is  in  order. 

Bryan  indicated  probable  temporal  correlations  between  the  Reddish-brown  Silt 
and  the  Laminated  Pink  Silt  and  Rubble  Zone  recognized  in  the  excavation  of  the 
deposit  at  the  mouth  of  the  cave.  Unlike  the  Reddish-brown  Silt,  this  middle  unit  at 
the  cave  entrance  rarely  contains  bones  (Bryan  1979a).  Although  no  radiometric  dates 
were  obtained  from  the  Laminated  Pink  Silt  and  Rubble  Zone,  a  '^C  date  of  12  600  ± 
170  B.P.  (A-1565)  was  obtained  at  the  lower  boundary  of  the  above  unit — Bristlecone 


Table  3.  Wisconsinan  and  early  Holocene  fish,  amphibians,  and  reptiles  from  Smith  Creek  Canyon.  G)  = 
previously  reported  by  Goodrich  (1965).  B)  =  previously  reported  by  Brattstrom  (1958,  1976).  1)  =  now 
living  within  50  miles;  2)  =  not  living  locally  but  elsewhere  in  Great  Basin;  3)  =  not  presently  within  the 
Great  Basin;  ?  =  age  assignment  in  question. 


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Fish 

Salmo  clarki 
Gila  atraria 

Amphibians 

Scaphiopiis  cf.  intermontanus  X 

5.  cf.  hammondi  (B)  3 

fiw/o  boreas  2 

Bufo  cf.  woodhousei  1 

Bm/o  sp.  (B) 
/?a«a  sp. 

Reptiles 

Crotaphytus  collaris  X 

Crotaphytiis  wislizeni  X 

Crotaphytus  sp. 

Sceloporus  magister  2 

Sceloporus  occidentalis  or  iindulatus         X 

Sceloporus  graciosus  (B)  X 

Sceloporus  sp. 

f/Za  stansburiana  (B)  X 

Phrynosoma  platyrhinos  (B)  X 

Phrynosoma  douglassi  2 

Phrynosoma  sp.  (G) 

Cnemidophorus  cf.  //^r/5  X 

Coluber  constrictor  (G,  B)  X 

Masticophis  flagellum  (B)  2 

Pituophis  melanoleucus  (G,  B)  X 

Lumpropeltis  getulus  (G,  B)  X 

Lampropeltis  pyromelana  X 

Lampropeltis  triangulum  3 

Rhinocheilus  lecontei  X 

Thamnophis  sp.  (G) 

Hypsiglena  torquata  (G,  B)  X 

Crotalus  cf.  viridis  (G)  X 

Crotalus  viridis  (B) 


X 


X 


X 


X 


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X 

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X 

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9 

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Pine  and  Sheep  Dung  Zone  (Bryan  197%/).  Presumably  this  radiocarbon  date  provides 
a  minimum  age  estimate  for  the  Laminated  Pink  SiU  and  Rubble  Zone  and  by  corre- 
lation the  Reddish-brown  Silt  unit. 

The  third  depositional  unit  Bryan  described  from  the  rear  of  the  cave  is  the  Grey 
Silt,  Rubble,  and  Dung  Zone.  Organic  remains  are  very  common  within  this  unit.  All 
bones  incorporated  in  this  uppermost  unit  are  various  shades  of  white  without  per- 
mineralization.  Occasional  mixing  of  the  red-stained  bones  by  bioturbation  has  oc- 
curred near  the  lower  boundary  of  the  unit.  Although  this  unit  has  not  been  radiocarbon 


dated,  it  certainly  is  younger  than  10  000  yr  B.P.  and  the  upper  portion  probably  is 
less  than  3000  years  old  (Bryan  1979^/). 

Results 

Table  3  is  a  chronological  check-list  of  late  Pleistocene  and  Holocene  age  fish, 
amphibians,  and  reptiles  from  Smith  Creek  Canyon.  Many  of  these  taxa  have  not  been 
reported  previously  from  the  late  Pleistocene  of  the  Great  Basin.  Table  4  is  an  updated 
check-list  of  fossil  mammals  known  from  Smith  Creek  Canyon.  Although  previous 
reports  have  described  the  mammals  (Harrington  1934.  Stock  1936.  Goodrich  1965. 
Miller  1979).  this  table  shows  their  stratigraphic  provenience,  not  adequately  given 
before . 

Following  is  an  annotated  list  of  the  fossil  fish,  amphibians,  reptiles,  and  mammals 
that  we  recovered  from  packrat  middens  and  from  one  sedimentary  layer  (Reddish- 
brown  Silt)  in  Smith  Creek  Cave.  Abbreviations  for  the  fossil  localities  are  listed  in 
Table  2.  The  number  in  parentheses  refers  to  the  quantity  of  that  element.  We  follow 
the  nomenclature  and  ordering,  unless  otherwise  stated,  of  Smith  (1978)  for  the  fish. 
of  Stebbins  (1966)  for  the  amphibians  and  reptiles,  and  of  Jones  et  al.  (1979)  for  the 
mammals. 

Class  Osteichthyes 

Family  Salmonidae 

Salmo  clarki  (Cut-throat  Trout) 

Material. — SCC  Sed.:  R  angular.  R  dentary.  vertebrae  (25). 

Remarks. — The  right  angular  resembles  that  of  a  small  individual  of  the  Great 
Basin,  not  of  the  Colorado  River.  The  right  dentary  is  of  a  specimen  of  approximately 
100  mm  in  length.  All  the  vertebrae  are  of  small  individuals.  The  skeletal  elements 
were  identified  as  Salmo  rather  than  Salvelinus  malma  (Dolly  Varden.  the  other  trout 
in  the  Great  Basin)  because  ( 1)  the  angle  between  the  coronoid  and  post-dorsal  process 
of  the  angular  is  near  90°,  and  (2)  the  dentary  lacks  the  deep  groove  under  the  tooth 
platform.  The  small  size  of  the  specimens  suggest  that  they  were  from  a  creek  (Smith 
Creek?),  not  a  lake  (Lake  Bonneville)  (Gerald  R.  Smith  identifications  and  personal 
communication  1981). 

Distribution. — Salmo  clarki,  which  occurred  in  Lake  Bonneville  during  the  Pleis- 
tocene, is  the  widespread  trout  of  the  Great  Basin  and  Intermountain  Region  (Smith 
et  al.  1968.  Smith  1978). 

Family  Cyprinidae 
Gila  atraria  (Utah  Chub) 

Material. —SCC  Sed.:  Basioccipital  and  vertebrae  (3). 

Remarks.— The  basioccipital  with  a  pharyngeal  process  has  the  shape  and  angles 
of  Gila  atraria,  not  Richardsonius  helteatiis  (Redside  Shiner),  which  has  a  less  ovoid 
haemal  canal  and  a  less  obtuse  angle  between  the  cranial  part  o\'  the  bone  and  its 
pharnygeal  process.  The  basioccipital  belonged  to  a  fish  about  1  10  mm  long  (Gerald  R. 
Smith  identification  and  personal  communication  1981). 

Distribution.— Gila  atraria  is  native  to  the  Bonneville  and  upper  Snake  River 
drainages  and  occurred  in  Lake  Bonneville  during  the  Pleistocene  (Smith  et  al.  1968. 
Smith  1978). 

Class  Amphibia 

Order  Salientia 

Family  Pelobatidae 

Scaphiopus  cf.  intermontanus  (Great  Basin  Spadefoot  Toad) 

Material.—SCC  1:  tibiofibula;  SCC  Sed.:  tibiofibulae  (13).  radio-ulnae  (3). 
Remarks. — No  comparative  material  was  available  for  Scaphiopus  intermontanus 
although  we  have  an  excellent  series  of  the  closely  related  S.  luimmondi. 


Distribution. — Only  this  species  of  spadefoot  toad  presently  occurs  within  wood- 
lands and  sagebrush  areas  in  the  Great  Basin  (as  well  as  in  the  Snake  Range);  however, 
S.  cf.  hammondi  has  been  identified  from  an  unprovenienced  level  in  Smith  Creek 
Cave  (Brattstrom  1976).  ^ 

Family  Bufonidae 
Biifo  bore  as  (Western  Toad) 

Material. — SCC  Sed.:  sacral  vertebra,  R  ilium. 

Distribution. — Bufo  horeas  presently  occurs  throughout  most  of  the  Great  Basin 
except  the  drier  eastern  part  along  the  Nevada-Utah  border;  it  has  not  been  found  in 
the  Snake  Range. 

Bufo  cf.  woodhousei  (Woodhouse's  Toad) 

Material.— SCC  Sed.:  tibiofibula. 

Remarks. — Martin  (1973)  used  the  ratio  of  the  tibiofibula,  minimum  width  relative 
to  length,  to  differentiate  species  of  Bufo  {B.  boreas:  690-760;  B.  cognatus:  840-1050; 
B.  hemiophrys:  950-1160;  B.  microscaphus:  670-850;  and  B.  woodhousei:  770-950). 
The  ratio  of  the  Smith  Creek  Cave  sediment  fossil  is  1013  (1.84/18.16  mm  x  10  000), 
being  relatively  stout  and  thick  in  the  middle.  Although  our  fossil  could  be  identified 
as  B.  cognatus  based  upon  the  tibiofibula  ratio,  we  refer  our  specimen  to  B.  wood- 
housei because  of  its  present  closer  distribution;  B.  cognatus  does  not  live  in  the  Great 
Basin.  Additional  fossils  of  Bufo  are  needed  to  refine  our  identification. 

Distribution. — Bufo  woodhousei  presently  does  not  inhabit  much  of  the  Great 
Basin;  the  western  edge  of  its  range  is  just  north  and  east  of  the  Snake  Range. 

Bufo  sp.  (toad) 

Material. — SCC  Sed.:  tibiofibulae  (3),  radio-ulna. 

Remarks. — We  were  unable  to  identify  the  fragmented  fossils  to  species. 

Family  Ranidae 
Rana  sp.  (frog) 

Material. — SCC  Sed.:  tibiofibulae  (4),  coracoid,  atlas. 
Remarks. — We  were  unable  to  assign  the  specimens  to  species. 
Distribution. — Presently  R.  pretiosa  and  R.  pipiens  occur  within  the  Great  Basin 
(Stebbins  1966). 

Class  Reptilia 

Order  Squamata 

Suborder  Sauria 

Family  Iguanidae 

Crotaphytus  collaris  (Collared  Lizard) 

Material. — SCC  Sed.:  L  dentaries  (3),  R  dentary. 

Remarks. — C.  collaris  and  C.  wislizeni  can  be  separated  from  most  other  iguanid 
lizards  by  their  larger  size  and  their  dental  characters.  The  individual  teeth  of  C. 
collaris  are  relatively  wide  anteroposteriorly  as  compared  to  those  of  C.  wislizeni,  the 
posterior  teeth  strongly  tricuspid,  the  anterior  tending  toward  blunt  spikes,  some  with 
a  slight  posterior  curve.  In  C.  wislizeni  the  anterior  three-quarters  of  the  teeth  are 
sharp,  recurved  simple  cusps  with  only  a  few  posterior  teeth  tricusped. 

Distribution. — The  Collared  Lizard  occurs  throughout  the  Great  Basin  in  a  variety 
of  mountain  and  rocky  habitats. 

Crotaphytus  wislizeni  (Leopard  Lizard) 

Material. — SCC  Sed.:  L  dentaries  (2),  L  maxillae  (3).  R  dentaries  (3),  R  maxillae 
(4). 


42- 


40- 


122 
1 


Figure  2.  Present  distributions  of  Sceloporus  magister  (L's),  Lampropehis  triangiiliim  (parallel  lines), 
and  L.  pyroineUina  (mixed  lines)  (from  Stebbins  1966).  ( I)  Fossil  localities  and  Snake  Range.  The  distribution 
of  L.  triangulum  overlaps  the  entire  range  of  L.  pyromelana  only  in  Utah.  The  distribution  of  .V.  magister 
overlaps  only  the  southern  tip  of  both  species  of  Lamprupeltis  in  Utah  and  all  of  L.  pyronwlanti  in  Arizona. 

Remarks. — The  distinguishing  characters  are  listed  under  C.  collaris. 

Distribution. — Like  the  Collared  Lizard,  the  Leopard  Lizard  occurs  throughout 
the  Great  Basin,  although  usually  in  desertscrub  communities  on  the  finer  alluvial 
habitats  of  the  valleys. 


Crotuphytus  sp.  (Collared  or  Leopard  Lizard) 

Material. — SCC  Sed.:  L  dentaries  (2),  L  maxillae  (3),  R  dentaries  (2),  R  maxilla. 
Remarks. — We  were  unable  to  assign  these  elements  to  species. 

Sceloporus  magister  (Desert  Spiny  Lizard) 

Material. —SCC  Sed.:  L  dentary. 

Remarks. — The  Desert  Spiny  Lizard  can  be  differentiated  from  other  sceloporine 
lizards  of  the  Great  Basin  by  its  larger  size. 

Distribution. — The  range  of  S.  magister  presently  ends  southwest  of  the  Snake 
Range  (Fig.  2).  Although  no  other  spiny  lizards  reach  the  size  of  5.  magister  in  the 
Great  Basin,  other  very  similar  sized  species  occur  farther  south  in  southern  Arizona 
(S.  clarki  and  S.  Jarrovi). 


10 

Sceloporus  occidentalis  or  undiilcitiis  (Western  or  Eastern  Fence  Lizard) 

Material. — LC  2a:  epidermal  scale;  SCC  Sed.:  L  dentaries  (6),  R  dentaries  (2),  R 
maxillae  (5),  frontal. 

Remarks. — There  are  no  satisfactory  dental  criteria  for  separating  the  closely  re- 
lated S.  unditlatiis  and  S.  occidentalis;  both  species  have  larger  races  at  their  northern 
distribution.  Since  the  latter  species  now  lives  within  the  Snake  Range,  we  refer  our 
fossils  to  it.  The  nearest  population  oi S.  undidatus  is  to  the  east  in  Utah.  Both  species 
possibly  occurred  within  the  mountain  range  in  the  late  Pleistocene. 

Sceloporus  graciosus  (Sagebrush  Lizard) 

Material.— SCC  Sed.:  R  dentaries  (2),  frontal. 

Remarks. — The  skeletal  elements  of  5.  graciosus  can  be  differentiated  from  those 
of  other  sceloporine  lizards  of  the  Great  Basin  by  their  distinctly  smaller  adult  size  and 
their  slender  pointed  teeth  with  weakly  developed  secondary  cusps. 

Distribution. — S.  graciosus  now  occurs  throughout  the  Great  Basin  including  the 
Snake  Range. 

Sceloporus  sp.  (spiny  lizard) 

Material. — SCC  Sed.:  L  dentaries  (2),  L  maxilla.  R  dentary,  R  maxillae  (2). 
Remarks. — We  were  unable  to  assign  these  elements  to  species. 

Uta  stansburiana  (Sideblotched  Lizard) 

Material. — LC  modern:  L  dentary. 

Remarks. — The  single  element  compared  well  with  modern  Uta  stansburiana.  The 
lizard  presently  lives  in  Smith  Creek  Canyon,  and  the  only  skeletal  specimen  recovered 
in  our  study  came  from  a  modern  packrat  midden. 

Phrynosoma  douglassi  (Short-horned  Lizard) 

Material.— hC  2a:  frontal;  LC  3:  L  maxilla;  SCC  Sed.:  L  dentary,  R  maxillae  (2), 
frontal. 

Remarks. — The  recovered  dentary,  maxillae,  and  frontals  could  not  be  distin- 
guished from  those  of  modern  P.  douglassi.  In  bone  and  dental  characters  of  the 
frontal,  dentary.  and  maxilla,  P.  douglassi  are  distinct  from  P.  platyrhinos  (Robinson 
and  Van  Devender  1973).  P.  douglassii  has  tall  Sceloporus-WkQ  teeth  with  well-devel- 
oped secondary  cusps  and  a  rounded  bottom  to  the  dentary.  P.  platyrhinos  has  short 
peg-like  teeth  with  reduced  secondary  cusps. 

Phrynosoma  platyrhinos  (Desert  Horned  Lizard) 

Mrt/mV//.— SCC  Sed.:  L  dentary. 

Distribution. — Both  Phrynosoma  platyrhinos  and  P.  douglassi  live  within  the 
Great  Basin  today,  the  latter  only  at  higher  elevations  on  a  few  mountain  ranges  in 
northeastern  Nevada  (Stebbins  1966). 

Family  Teiidae 
Cnemidophorus  cf.  //^t,'/7.v  (Western  Whiptail  Lizard) 

Material. —SCC  Sed.:  R  dentaries  (4). 

Remarks. — We  refer  our  specimens  to  C.  cf.  tigris  because  it  is  the  only  species 
of  whiptail  lizard  now  living  within  the  interior  Great  Basin.  Modern  C.  tigris  and  the 
fossils  have  very  large  dentaries.  Other  species  of  whiptail  (e.g.,  C.  hurti)  with  den- 
taries of  similar  size  now  occur  far  south  of  the  Great  Basin. 


Suborder  Serpentes 

Family  Cokibridae 

Pituophis  fucldnolenciis  {Gt>pher  Snake) 

Material. see  Sed.:  palatine,  vertebrae  (23). 

Ri'inarks. — The  vertebrae  o'i  Fituoplus  inelanolcucus  are  most  similar  to  those  of 
Elaphe.  We  used  the  criteria  described  by  Auffenberg  (1963)  to  differentiate  the  two 
species. 

Distribution. — The  Gopher  Snake  is  very  common  throughout  the  Great  Basin. 

Lampropeltis  pyromeliimi  (Sonoran  Mountain  Kingsnake) 

Material.— sec  Sed.:  vertebrae  (29). 

Remarks. — The  vertebrae  of  Lampropeltis  pyromelana  are  typical  of  kingsnakes, 
with  well-developed  subcentral  ridges.  We  differentiate  L.  pyromelana  from  L.  trian- 
gidum  by  two  criteria:  (1)  the  cotyle  and  condyle  are  proportionally  larger  in  L.  py- 
romelana at  all  stages  of  growth,  and  (2)  the  accessory  processes  are  fairly  pointed  on 
L.  pyromelana  whereas  they  are  longer  and  globose  (as  viewed  anteriorly)  on  L.  trian- 
gulum.  Both  species  have  low  neural  spines  unlike  L.  getuUis  or  Rhinoeheilns  leeontei. 

Distribution. — The  Sonoran  Mountain  Kingsnake  presently  occurs  only  in  a  few 
relictual  populations  in  the  Great  Basin  including  the  Snake  Range  (Fig.  2).  It  com- 
monly occurs  in  the  central  and  southern  Wasatch  Mountains  and  the  mountainous 
region  farther  south  and  west. 

Lampropeltis  triangulum  (Milksnake) 

Material. — SCC  Sed.:  vertebra. 

Remarks. — The  distinguishing  criteria  were  discussed  under  L.  pyromelana. 

Distribution. — The  Milksnake  occurs  on  the  eastern  periphery  of  the  Great  Basin 
in  the  Wasatch  Mountains  and  farther  south  in  Utah  but  not  as  far  west  as  the  Snake 
Range  (Fig.  2). 

Rhinoeheilus  leeontei  (Long-nosed  Snake) 

Material. — LC  modern:  vertebra;  SCC  Sed.:  vertebrae  (13) 

Remarks. — The  vertebrae  of  Rhinoeheilus  are  quite  characteristic  and  can  readily 
be  identified  (Hill  1971,  Van  Devender  and  Mead  1978). 

Distribution.— The  Long-nosed  Snake  lives  throughout  the  Great  Basin  except  for 
the  northwesternmost  region. 

Thamnophis  sp.  (garter  snake) 

Material.— SCC  Sed.:  vertebrae  (3). 

Remarks. — We  were  unable  to  assign  the  specimens  to  species. 

Distribution.— Presently  T.  elegans  (Wandering  Garter  Snake)  lives  within  the 
Great  Basin,  including  the  Snake  Range,  and  T.  sirtalis  (Common  Garter  Snake)  bor- 
ders parts  of  the  basin  (Stebbins  1966). 

Hypsiglena  torquata  (Night  Snake) 

Material.— CHC  lb:  vertebra;  SCC  Sed.:  L  dentary.  vertebrae  (36). 

Remarks.— H.  torquata  has  small  generalized  colubrid  vertebrae,  which  are  very 
similar  to  those  of  other  small  snakes  such  as  Sonora  semianulata.  Identification  cri- 
teria used  here  were  those  used  by  Van  Devender  and  Mead  (1978). 

Distribution. — The  Night  Snake  occurs  over  most  of  the  Great  Basin. 


12 


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16 


Family  Viperidae 
Crotalus  cf.  viridis  (Western  Rattlesnake) 

Material. see  Sed.:  vertebrae  (5) 

Distribution. — We  refer  our  specimens  to  C.  viridis  because  they  are  from  a  me- 
dium-sized rattlesnake  and  this  species  presently  inhabits  the  Great  Basin.  C.  mitchelli 
is  very  similar  but  occurs  no  farther  north  than  the  Mohave  Desert. 

Class  Mammalia 

Order  Chiroptera 

Genus  and  species  indeterminate 

Material. — SCC  Sed.:  isolated  teeth,  mandible. 

Remarks. — The  isolated  teeth  and  the  fragment  of  a  mandible  did  not  allow  generic 
identification. 

Distribution. — Many  species  of  bats  occur  in  the  Great  Basin  (Barbour  and  Davis 
1969). 

Order  Lagomorpha 

Family  Ochotonidae 

Ochotona  cf.  princeps  (pika) 

Material.— SCC  4:  dung  pellets  (17),  RM>;  SCC  5:  dung  pellets  (2),  RP;  SCC 
Sed.:  LP3;  STV  1:  dung  pellet;  STV  2:  dung  pellets  (>50);  STV  3:  dung  pellets  (3). 

Remarks. — The  RP^  from  SCC  5  is  referred  to  Ochotona,  but  in  view  of  its  worn 
state,  it  could  possibly  be  a  Sylvilagus  idahoensis.  Miller  (1979)  reported  that  pika 
remains  were  recovered  in  all  stratigraphic  units  in  Smith  Creek  Cave;  the  youngest 
occurrence,  however,  cannot  be  determined.  Pika  also  was  recovered  from  Council 
Hall  Cave;  unfortunately,  the  remains  (not  described)  were  not  reported  in  relation  to 
the  testpits,  stratigraphy,  or  associated  radiocarbon  age  (Miller  1979,  Bryan  \919b). 
Small  mammal  bones  {Ochotonal)  occurred  throughout  the  upper  two  meters  (above 
a  23  900  ±  970  yr  B.P.  radiocarbon  date,  GaK-5100)  of  organic  layers  in  Test  Pit  2 
(Bryan  1979/?). 

Two  packrat  middens  in  Smith  Creek  Cave  and  three  middens  in  Streamview  Shelter 
contained  lagomorph  dung  pellets  referable  to  Ochotona  cf.  princeps  (Fig.  3).  Dung 
pellets  from  each  packrat  midden  were  measured  (length  and  width)  and  the  measure- 
ments compared  with  those  from  samples  from  modern  Ochotona  princeps,  Sylvilagus 
idahoensis,  and  S.  nuttallii.  The  advantage  of  the  preserved  dung  pellets  from  the 
packrat  middens  over  the  skeletal  fragments  from  the  cave  sediments  is  that  presum- 
ably Ochotona  lived  at  the  fossil  site  and  was  not  brought  there  via  a  raptor  stomach. 
Pika  (skeletal  remains)  has  been  identified  previously  from  mountain  ranges  outside  its 
present  distribution  and  from  elevations  below  its  current  lower  limit  in  ranges  it  now 
inhabits  (Fig.  4;  Grayson  1977,  1981,  in  press  a,  b.  Miller  1979).  To  our  knowledge,  this  is 
the  first  record  of  late  Pleistocene  age  dung  of  Ochotona  south  of  the  permafrost  in 
North  America  (Guthrie  1973).  A  more  detailed  account  of  the  fossil  Ochotona  in  the 
Great  Basin  is  in  progress. 

Distribution. — Pika  does  not  now  live  in  the  Snake  Range  or  in  any  nearby  moun- 
tains (Hall  1946,  Hall  and  Kelson  1959),  although  it  does  inhabit  the  mountainous 
regions  on  the  west,  north,  and  east  sides  of  the  Great  Basin  (Fig.  4).  Only  O.  p. 
nevadensis  (Ruby  Mountains  region)  and  O.  p.  tutelata  (Toquima  Range  and  Desatoya 
Mountains)  live  in  restricted  relictual  localities  within  the  interior  Great  Basin. 

Family  Leporidae 
Sylvilagus  {^Brachylagus)  idahoensis  (Pygmy  Rabbit) 

Material.— SCC  Sed.:  M,,. 

Remarks. — S.  idahoensis  has  not  been  previously  recorded  from  Smith  Creek 
Canyon  (Miller  1979).  In  the  central  Great  Basin  at  Gatecliff  Shelter,  the  Pygmy  Rabbit 


17 


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Figure  3.  Measurements  (width  and  length)  of  lagomorph  dung  pellets,  (a)  Ochotona  cf.  princeps.  fossil, 
this  report;  (b)  O.  princeps,  modern,  California;  (c)  Sylvilugus  idahoensis,  modern,  Nevada:  (d)  S.  nuttallii, 
modern,  Nevada. 


was  recovered  throughout  the  deposit  (Grayson  ///  press  h).  North  of  the  Great  Basin, 
S.  idahoensis  is  recorded  in  late  Pleistocene  context  at  Jaguar  Cave  (Guilday  and 
Adams  1967),  Wasden  site  (Owl  Cave,  Guilday  1969),  and  Moonshiner  Cave  (Anderson 
1974).  South  of  the  Great  Basin  it  is  reported  from  Tule  Springs,  Nevada  (Mawby  1967) 
and  Isleta  Cave,  New  Mexico  (Harris  1977). 

Distribution. — The  Pygmy  Rabbit  presently  occurs  across  the  northern  and  east- 
central  Great  Basin  (Hall  1946). 

Sylvilagus  sp.  (rabbit) 

Material. —see  Sed.:  RP^ 

Remarks. — The  specimen  could  not  be  identified  to  species. 


Lepiis  sp.  (hare) 

Material.— sec  Sed.:  RM,. 

Remarks. — The  specimen  could  not  be  identified  to  species. 


18 


Figure  4.     Distribution  of  Ochotona  princeps  on  isolated  mountains  within  and  bordering  the  Great  Basin 
(from  Hail  1946,  Hall  and  Kelson  1959).  (1)  Fossil  localities  of  Smith  Creek  Canyon  and  Garrison. 


Order  Rodentia 

Family  Sciuridae 

Eiitamias  minimus  (Least  Chipmunk) 

Material.— sec  Sed.:  R  mandible  P4Mi_3. 

Remarks. — This  is  the  smallest  chipmunk  in  Nevada.  Assignment  of  our  specimens 
to  this  species  was  on  the  basis  of  size.  Although  the  other  species  of  Eutamias  of  the 
Great  Basin  are  restricted  to  the  vicinity  of  coniferous  trees,  E.  minimus  may  live  in 
sagebrush  {Artemisia  tridentata)  at  both  high  and  low  elevations  (Hall  1946). 

Eutamias  cf.  umhrinus  (Uinta  Chipmunk) 

Material.— sec  Sed.:  frontal. 

Remarks. — Neither  of  the  two  species  of  Eutamias  have  been  reported  from  Smith 
Creek  Canyon  (Miller  1979).  Our  specimen  compared  most  favorably  with  modern  E. 
umhrinus. 

Distribution. — The  Uinta  Chipmunk  is  a  medium-sized  chipmunk  that  presently 
occurs  in  the  Snake  Range  (Hall  1946). 


19 

EutiiDiias  sp.  (chipmunk) 
Material. see  Sed.:  LM,  (2).  RM,.  RM'  (3),  LM-',  RM^ 

Mannota  flavivcntris  (Yellow-bellied  Marmot) 

Material. see  Sed.:  LI. 

Remarks. — Although  we  are  assuming  our  specimen  is  M.flaviventris,  the  isolated 
incisor  does  not  allow  for  specific  identification.  The  Hoary  Marmot,  Mannota  cali- 
gata,  cannot  be  ruled  out  definitively  based  upon  our  specimen,  though  its  occurrence 
in  the  Great  Basin  in  the  late  Pleistocene  seems  unlikely. 

Distribution. — Marmota  jlaviventris  presently  occurs  in  the  Snake  Range  at  ele- 
vations higher  than  Smith  Creek  Cave. 

Ammospermophilus  lencnrns  (White-tailed  Antelope  Squirrel) 

Material.— Le  2a:  frontal;  SCC  Sed.:  L  mandible  M,_.^,. 

Remarks. — The  White-tailed  Antelope  Squirrel  has  not  previously  been  reported 
from  Smith  Creek  Canyon  (Miller  1979).  Our  specimens  compare  with  modern  repre- 
sentatives of  y4.  lencnrns. 

Distribution. — Ammospermophilns  lencnrns  presently  lives  in  most  of  the  Great 
Basin,  including  the  area  of  the  Snake  Range  (Hall  1946). 

Spermophilns  cf.  richardsonii  (Richardson's  Ground  Squirrel) 

Material.— 'Le  1:  RM^;  LC  4:  LM,. 

Remarks. — Spermophilns  richardsonii  has  not  been  recovered  previously  from 
Smith  Creek  Canyon,  though  it  has  been  recorded  from  the  more  northern  localities 
of  Jaguar  Cave  (Guilday  and  Adams  1967)  and  Moonshiner  Cave  (Anderson  1974), 
Idaho.  Both  Davis  (1939)  and  Hall  (1946)  have  expressed  the  belief  that  S.  richardsonii 
must  have  occupied  a  much  wider  range  over  an  ecological  area  now  filled  by  S. 
beldingi  and  S.  armatns.  On  the  basis  of  the  Smith  Creek  Canyon  fossils  of  S.  cf. 
richardsonii,  we  concur  with  them.  Our  two  specimens  compared  most  favorably  v\ith 
modern  S.  richardsonii. 

Distrihntion. — Richardson's  Ground  Squirrel  does  not  occur  presently  in  the 
Snake  Range.  The  present  distribution  of  5'.  r.  nevadensis  in  the  Great  Basin  is  centered 
in  the  Independence  Mountains  of  northern  Nevada,  and  it  occurs  no  farther  south 
than  the  Roberts  Mountains  (Hall  1946). 

Spermophilns  cf.  beldingi  (Belding's  Ground  Squirrel) 

Material.— Le  2b:  L  maxilla  M'. 

Remarks. — Spermophilns  beldingi  has  not  been  reported  previously  from  Smith 
Creek  Canyon,  and  the  only  other  published  fossil  or  subfossil  record  for  Belding"s 
Ground  Squirrel  in  the  Great  Basin  is  from  Stratum  3  at  Gatecliff  Shelter  (Grayson  //; 
press  h).  Our  single  specimen  compared  most  favorably  with  modern  ,S.  beldingi. 

Distribution. — Spermophilns  beldingi  is  ecologically  and  physically  very  similar 
to  S.  richardsonii.  Belding's  Ground  Squirrel  presently  does  not  occur  in  the  Snake 
Range  or  in  any  immediate  mountain  range.  Its  present  distribution  centers  in  the 
northern  Sierra  Nevada  and  in  the  higher  mountains  of  north  central  and  northern 
Great  Basin,  though  it  occurs  farther  south  than  .S'.  richardsonii.  dow  n  to  the  TcHiuima 
Range  (Hall  1946)'] 

Spermophilns  cf.  lateralis  (Golden-mantled  Ground  Squirrel) 

Material. — LC  modern:  RM,;  LC  2a:  L  maxilla  M'"''.  L  mandible  M,.;,;  SCC  1: 
R  mandible  P,M,;  SCC  Sed.:  R  mandible  M,_,.  RM,.  LM'.  RM,_.,. 

Remarks. — Although  S.  lateralis  was  not  previously  reported  from  Smith  Creek 
Canyon  (Miller  1979).  it  is  fairly  widespread  in  the  Smith  Creek  Canyon  fossil  localities 


20 


and  apparently  was  common  in  the  canyon  throughout  most  of  the  late  Pleistocene. 
Although  we  have  found  this  to  be  the  most  common  species  of  Spermophiliis,  Miller 
(1979)  considered  that  S.  cf.  tcnvnsendii  was  the  most  prevalent  in  the  Smith  Creek 
Cave  record.  Miller  (1979)  stated  that  Goodrich  (1965)  referred  his  specimens  from 
Smith  Creek  Cave  to  S.  cf.  townsendii;  although  he  considered  that  species,  he  did 
not  definitely  refer  his  specimens  there.  However,  the  presence  of  Townsend's  Ground 
Squirrel  in  Smith  Creek  Canyon  in  the  late  Pleistocene  seems  plausible. 

Spermophilus  sp.  (ground  squirrel) 
Material.— \.C  2b:  RM,;  SCC  Sed.:  P  (3),  RMg,  RM^. 

Family  Geomyidae 
Thomomys  sp.  (pocket  gopher) 

Material. —SCC  Sed.:  LM». 

Remarks. — We  were  unable  to  assign  our  single  specimen  to  a  species. 

Family  Heteromyidae 
Perognathus  sp.  (pocket  mouse) 

Material. —SCC  1:  L  mandible  P4  (3),  R  mandible,  R  maxilla  M^;  SCC  Sed.:  L 
mandible,  R  maxilla  M*  (2);  STV  2:  L  maxilla  (2),  R  maxilla. 

Remarks. — We  were  unable  to  assign  these  specimens  to  species. 

Dipodomys  sp.  (kangaroo  rat) 

Material.— SCC  Sed.:  LP. 

Remarks. — We  were  unable  to  assign  this  specimen  to  species. 

Family  Cricetidae 
cf.  Peromysciis  (white-footed  mouse) 

Material.— CnC  lb:  RM'-^;  SCC  I:  LM^;  SCC  Sed.:  LM',  RM'-^;  STV  2:  LM,, 
RM'-^:  STV  3:  LM,_2. 

Remarks. — We  have  not  identified  these  specimens  to  species  because  we  do  not 
have  available  a  sufficient  comparative  collection  of  Reithrodontomys  and  Peromyscus 
from  Nevada.  Presently  Reithrodontomys  megalotis,  Peromyscus  eremicus,  P.  crini- 
tus,  P.  maniculatus,  P.  hoy  Hi,  and  P.  truei  live  in  or  near  the  Great  Basin.  Previous 
work  at  Smith  Creek  Cave  has  produced  specimens  assigned  to  Peromyscus  sp.,  but 
Reithrodontomys  and  Onychomys  have  not  been  identified  from  the  fossil  localities  in 
Smith  Creek  Canyon,  though  they  very  well  may  be  included. 

Neotoma  lepida  (Desert  Packrat) 

Material.— CHC  la:  LM' ^  CHC  lb:  LM,  (2);  SCC  Sed.:  LM,  (2),  RM,  (2),  RM' 
(3),  LM'  (8);  STV  2:  RM^. 

Neotoma  cinerea  (Bushy-tailed  Packrat) 

Material.— CHC  la:  LM',  RM'  (2);  CHC  lb:  LM,,  LM';  SCC  1:  lower  leg  assem- 
blage with  hide;  SCC  Sed.:  RM,,  RM'  (7). 

Remarks. — Neotoma  lepida  and  N.  cinerea  are  found  living  in  Smith  Creek  Can- 
yon. Their  middens,  both  modern  and  fossil,  can  be  found  throughout  the  Great  Basin. 
The  excellent  preservation  and  the  advantages  of  having  a  packrat  midden  are  illus- 
trated by  the  partial  mummy  of  the  hind  quarters  of  the  Bushy-tailed  Packrat  (ca. 
1 1  600  yr  B.P.).  Molars  of  N.  cinerea  are  generally  larger  than  those  of  N.  lepida. 


21 


L 


J 


Figure  5.     Occlusial  (left)  and  lingual  (right)  views  of  the  Phenacoinys  cf.  intermedins  RM,  from  Smith 
Creek  Cave.  Bar  =  2  mm. 


Neotoma  lepida  or  cinerea  (Desert  or  Bushy-tailed  Packrat) 

Material.— CHC  la:  RM^;  CHC  lb:  L&RM^  (5).  LM.,  (2).  M^  (2).  M,  (2);  SCC 
Sed.:  LM,  (2),  RM,  (2),  L&RM2  (17),  L&RM^  (8),  M-^  (8),  M3  (8). 

Neotoma  sp.  (packrat) 
Material.— SCC  Sed.:  M3;  STV  2:  M  fragment. 

Phenacomys  cf.  intermedius  (Heather  Vole) 

Material.— SCC  Sed.:  RM,. 

Remarks. — Only  one  specimen  was  identifiable  as  Phenacomys  cf.  intermedius. 
Criteria  for  identification  were  the  presence  of  tooth  roots,  lack  of  cementum,  and 
distinctive  occlusal  pattern  (Fig.  5;  Guilday  and  Parmalee  1972).  The  isolated  molar 
does  not  allow  for  specific  identification. 

Distribution. — The  Heather  Vole  presently  does  not  live  in  the  Great  Basin  (Hall 
1946).  Our  specimen  from  Smith  Creek  Cave  is  the  first  reported  late  Pleistocene 
Phenacomys  from  the  Great  Basin.  Grayson  (1981)  has  recovered  a  specimen  from 
Gatecliff  Shelter  dating  approximately  5300  yr  B.P.,  which  illustrates  the  middle  Holo- 
cene  survival  of  this  presently  extirpated  mammal. 

Microtus  cf.  longicaudus  (Long-tailed  Vole) 

Material.— SCC  Sed.:  RM,  (7),  LM,  (8),  R  mandible:  STV  2:  LM,. 

Remarks. — Our  fossil  specimens  appear  most  similar  to  the  Long-tailed  Vole, 
although  we  find  it  difficult  to  differentiate  M.  longicaudus  from  M.  montanus.  Of  the 
15  fossil  M,'s  examined,  14  had  five  closed  alternating  triangles,  1  had  six  triangles. 
Only  M.  montanus  has  been  reported  from  Smith  Creek  Cave  (Goodrich  1965,  Miller 
1979).  Goodrich  compared  his  fossils  with  M.  californicus,  M.  montanus,  M.  town- 
sendii,  M.  oregoni,  and  M.  pennsylvanicus:  M.  longicaudus  was  not  compared. 

Distribution. — Microtus  montanus  and  M.  longicaudus  are  the  only  species  of 
meadow  vole  in  Nevada  (Hall  1946).  Normally,  M.  montanus  is  found  in  the  valleys 
and  M.  longicaudus  in  the  mountains. 

Microtus  sp.  (meadow  vole) 

Material.— \.C  2b:  RM,_.,;  SCC  4:  RM,;  SCC  Sed.:  LM'  (3),  RM'  (3),  LM^  (4), 
LM2  (4),  RM3  (2).  LM3  (2),  RW. 


22 


Order  Carnivora 

Family  Mustelidae 

Spilogale  piitoiius  (Spotted  Skunk) 

Material.—SCC  Sed.:  L  mandible  C,P..^4M,_2. 

Remarks. — This  is  the  only  place  where  we  disagree  with  the  classification  of 
Jones  et  al.  (1979).  We  follow  Kurten  and  Anderson  (1980)  and  group  the  Western 
Spotted  Skunk  (Spilogale  gracilis)  with  the  Eastern  Spotted  Skunk  {S.  putorius).  Our 
specimen  from  Smith  Creek  Cave  has  an  alveolar  length  (anterior  edge  of  C,  to  pos- 
terior edge  of  Mo)  of  19.0  mm  and  an  occlusial  length  of  the  Mo  of  2.1  mm.  These 
measurements  compare  well  with  two  modern  specimens  of  5.  putorius  from  the  Ruby 
Mountains,  Elko  and  White  Pine  Counties,  Nevada. 

Distribution. — The  Spotted  Skunk  is  common  throughout  the  Great  Basin. 

Order  Artiodactyla 
Genus  and  species  indeterminate 

Material. — CHC  la:  dung  pellets  ( 15);  SCC  1:  keratinous  hoof  fragment;  SCC  Sed.: 
I  fragment. 

Remarks. — Because  of  fragmented  state  of  preservation,  we  were  unable  to  iden- 
tify further  these  specimens. 

Discussion  and  Summary 

We  report  here  the  occurrence  of  2  fish,  4  anurans,  9  lizards,  8  snakes,  and  17 
small  mammals.  This  assemblage  adds  15  amphibians  and  reptiles  and  7  mammals  to 
the  approximately  46  animals  previously  known  from  the  late  Pleistocene  and  early 
Holocene  of  Smith  Creek  Canyon  (Goodrich  1965,  Miller  1979).  Over  half  of  the  large 
herbivores  {Equus  spp.,  Camelops  sp.,  IHemiauchenia  sp.,  Oreamnos  harringtoni, 
and  IBreameryx  sp.)  and  the  two  carnivores  {Martes  nobilis  and  IFelis  atrox)  reported 
from  Smith  Creek  Canyon  are  extinct  (Miller  1979).  Our  data  add  no  large  herbivore 
or  carnivore  species  to  the  local  fauna.  Of  the  extant  species  from  the  fossil  sites.  16 
are  not  recorded  from  the  Snake  Range  or  in  the  immediate  valleys  (Bufo  boreas, 
Sceloporus  magister,  Phrynosoma  douglassi,  Lampropeltis  triangulum,  Masticophis 
flagellum,  Ochotona  princeps,  Spermophilus  cf.  richardsonii,  S.  cf.  beldingi,  Phena- 
comys  cf.  intermedius,  Vulpes  velox,  Ursus  sp.,  Mustela  vison,  Martes  sp.,  Felis  onca, 
Cervus  elaphus,  and  IBison  sp.). 

Few  large  herbivores  presently  occur  in  the  Snake  Range.  Oris  canadensis  was 
reintroduced  in  the  middle  1900s.  The  historic  distributions  of  Cervus  elaphus  and 
Bison  bison  in  the  eastern  Great  Basin  are  not  well  known,  though  Cervus  has  been 
sighted  in  White  Pine  County  (Hall  1946).  The  late  Pleistocene  occurrences  of  Ca- 
melops sp.,  IHemiauchenia  sp.,  Cervus  elaphus,  Odocoileus  sp.,  Equus  (both  large 
and  small  species),  ?Brcameryx  sp.,  Antilocapra  americana,  Oreamnos  harringtoni, 
Ovis  canadensis,  and  'IBison  in  Smith  Creek  Canyon  are  fairly  well  documented,  al- 
though the  exact  timing  (late  or  middle  Wisconsinan)  of  these  occurrences  is  not  under- 
stood and  is  discussed  below.  We  question  the  tentative  identification  of  Oreamnos 
americanus  reported  from  Council  Hall  Cave  (Miller  1979).  Although  the  estimated 
age  is  not  reported  for  the  modern  mountain  goat  specimen  (it  is  not  known  historically 
from  the  Great  Basin),  we  speculate  that  the  record,  if  Oreamnos,  is  of  the  extinct 
Oreamnos  harringtoni. 

If  our  reconstruction  of  the  Smith  Creek  Canyon  fauna-flora  assemblage  is  correct, 
during  the  late  Pleistocene,  montane  glaciers  on  Mount  Moriah  (3673  m  elevation) 
moved  down  into  the  upper  reaches  of  Smith  Creek  Canyon  to  an  estimated  elevation 
of  2900  m  (Drews  1958).  This  was  also  a  time  when  an  open  forest  with  Pinus  longaeva, 
P.  fiexilis,  and  Picea  engehnannii  was  present  on  the  coarse  talus  and  rock  outcrops 
of  the  limestone  entrance  region  of  the  canyon.  Alluvial  substrates  in  the  canyon 
bottom  probably  supported  a  mosaic  o{ Artemisia  spp.,  shrubs,  meadows,  and  riparian 


23 


Table  5.   Fauna  from  two  packrat  middens  on  a  rock  outcrop  in  the  middle  of  Snake  Valley  (1640  m 
elevation),  near  Garrison.  Utah.  (D)  =  dung.  (B)  =  bone  or  tooth. 


Garrison  No.  1 

Garrison  No.  2 

12  230 

±  180  yr  B.P. 

13  480 

±  250  yr  B.P. 

Gastropoda 

shell 

Osteichthyes 

B 

Aves 

B 

Sorex  sp. 

B 

Ochotona  cf.  princeps 

D 

B,  D 

Sylvihigus  idahocnsis 

B 

B 

Thomomys  sp. 

B 

Spermophilus  sp. 

B 

cf.  Peromyscus 

B 

Neotoma  sp. 

D 

B,  D 

Microtus  sp. 

B 

B 

Camelops  cf.  hesternus 

B 

Ovis  or  Odocoileiis 

D 

elements.  This  canyon  bottom  habitat  and  possibly  the  canyon  slopes  could  have  sup- 
ported much  of  the  fauna  (e.g.,  Phenacomys  and  Microtus)  recovered  from  the  three 
caves  and  the  rockshelters.  Ochotona  cf.  princeps  probably  occurred  on  all  suitable 
talus  slopes  throughout  the  unglaciated  portions  of  the  canyon  from  the  creek  level  at 
Streamview  to  Smith  Creek  Cave. 

Smith  Creek  Canyon  empties  into  the  broad  Snake  Valley  to  the  east.  During  part 
of  the  late  Wisconsinan  a  western  arm  of  Lake  Bonneville  filled  this  valley  to  an 
elevation  of  1580  m  (Mifflin  and  Wheat  1979).  This  high  lake  stand  was  only  4  km  from 
the  entrance  of  Smith  Creek  Canyon.  The  late  Pleistocene  vegetation  of  the  Snake 
Valley  in  the  area  above  the  lake  level  is  poorly  known. 

Thirty  km  south  of  Smith  Creek  Canyon  but  still  within  Snake  Valley,  we  re- 
covered packrat  middens  from  a  rocky  outcrop  in  the  center  of  the  valley  (near  the 
town  of  Garrison,  Utah,  10  km  east  of  the  Snake  Range,  the  closest  mountain  mass). 
The  late  Pleistocene  age  fauna-flora  assemblage  from  the  packrat  middens  is  presum- 
ably an  indication  of  the  type  of  habitat  above  the  pluvial  lake  level  but  below  the 
mountain  masses.  The  isolated  rock  outcrop  provided  a  suitable  habitat  for  limber  pine 
and  pika  (Table  5;  Thompson  and  Mead  1982).  Adjacent  areas  appear  to  have  been 
a  shrub  community,  with  some  nearby  areas  of  meadow.  Megafauna  of  the  valley 
consisted  of  at  least  Camelops  cf.  hesternus  and  Ovis  or  Odocoileus  (Table  5).  The 
Rancholabrean  age  fauna  of  Snake  Valley  is  not  well  known.  A  cave  acting  as  a  natural 
trap  has  produced  the  skull  of  a  wolverine  {Gulo  gulo),  though  unfortunately  the  age 
of  the  animal  is  not  known  (Barker  1976). 

The  density  of  the  shrub  communities  below  the  mountain  mass  but  above  the 
lake  level  cannot  be  determined  with  certainty.  The  recovery  of  Sylvilagus  idahoensis 
implies  that  at  least  some  areas  were  fairly  dense  with  tall  stands  of  Artemisia  spp. 
(Hall  1946).  Conversely,  the  recovery  of  Crotaphytus  wisHzeni  and  Phrynosoma  platy- 
rhinos  imply  areas  of  relatively  open  to  sparse  habitat,  possibly  exposed  playa  adjacent 
to  the  lake. 

Most  of  the  fauna  reported  here  (Tables  3  and  4)  come  from  the  inadequately 
dated  Reddish-brown  Silt  Zone  in  Smith  Creek  Cave.  We  have  already  mentioned  that 
the  accuracy  of  the  single  radiocarbon  date  on  bone  fragments  from  the  unit  (ca.  28  000 
yr  B.P.)  is  suspect.  The  minimum  age  this  unit  could  be  is  approximately  12  000  yr 
B.P.,  assuming  the  stratigraphic  associations  described  by  Bryan  (1979fl)  are  correct. 
Equally  plausible  is  that  the  unit  may  date  from  the  late  Wisconsinan  full  glacial  (ca. 
18  000  to  22  000  yr  B.P.)  or  even  middle  Wisconsinan  (^30  000  yr  B.P.).  The  temporal 
depth  of  this  unit  also  is  not  known.  Because  of  these  drawbacks  we  cannot  definitely 
state  when  or  what  the  faunal  associations  were  in  the  late  Pleistocene  of  Smith  Creek 


24 


Canyon.  We  can  state,  however,  that  at  least  these  taxa  were  in  the  canyon  during  the 
late  Wisconsinan.  This  becomes  important  when  considering  the  species  such  as  Och- 
otona,  Phenacomys,  and  other  locally  extirpated  animals. 

Tanner  (1978)  has  stated  that  most  of  the  present  Great  Basin  Desert  reptiles  have 
extended  their  ranges  into  the  Great  Basin  as  post-Pleistocene  introductions  from  a 
Pleistocene  refugium  of  the  southern  deserts  (Chihuahua,  Coahuila,  Sonora;  Ballinger 
and  Tinkle  1972).  The  late  Wisconsinan  records  oi  Crotaphytus  wislizeni,  Phrynosoma 
platyrhinos,  Sceloporus  magister,  Hypsiglena  torquata  and  Rhinocheilus  lecontei  in 
the  Smith  Creek  Canyon  fauna  presented  here  does  not  support  Tanner's  hypothesis. 
If  the  fauna  from  the  Reddish-brown  Silt  Zone  in  Smith  Creek  Cave  is  of  approximately 
12  000  to  22  000  yr  B.P.  or  even  >30  000  yr  B.P.,  then  many  of  the  lizards  and  snakes 
presently  inhabiting  the  eastern  Great  Basin  are  not  post-Pleistocene  invaders.  The 
important  fact  is  that  desert  elements  in  the  modern  fauna  were  already  in  the  Great 
Basin  prior  to  the  end  of  the  late  Wisconsinan.  These  results  call  for  reconsideration 
of  current  biogeographical  hypotheses  on  the  evolution  of  the  Great  Basin  Desert 
fauna. 

At  some  time  in  the  late  Pleistocene  and  early  Holocene,  some  of  the  amphibians, 
reptiles,  and  mammals  found  in  the  Snake  Range  and  Snake  Valley  either  adjusted 
their  distributions  or  became  extinct.  Martin  (1967)  has  previously  expressed  his  model 
that  Early  Man  exterminated  the  megafauna  of  North  America.  The  amphibians,  rep- 
tiles, and  small  mammals  adapted  individually  to  the  climatic  changes,  however  minor 
or  major,  of  the  late  Pleistocene  and  Holocene. 

Brown  ( 1971 ,  1978)  and  Grayson  (1 98 1 ,  in  press  a,b)  have  theorized  that  local  extir- 
pations in  the  Great  Basin  of  an  assemblage  of  small  mammals  on  a  mountain  range 
are  related  to  the  size  of  the  animal  population  and  the  mass  of  the  mountain.  In  the 
case  of  the  Snake  Range  (emphasizing  Smith  Creek  Canyon)  certain  animal  species 
became  extirpated  {Sceloporus  magister,  Phenacomys  cf.  intermedins,  Ochotona  cf. 
princeps,  and  others)  while  additional  species  developed  relictual  populations,  inhab- 
itants of  a  boreal  island  in  a  sea  of  sagebrush  (e.g.,  Lampropeltis  pyromelana  and 
Marmota  flaviventris).  The  smaller  mammals,  at  least  Ochotona  and  Phenacomys, 
lingered  on  into  the  early  Holocene  (Grayson  1981;  this  report).  The  Ochotona  cf. 
princeps  remains  reported  here  indicate  that  suitable  habitat  and  climate  occurred  in 
the  Snake  Valley  at  least  until  ca.  12  000  yr  B.P.  and  in  Smith  Creek  Canyon  until  ca. 
6500  yr  B.P.  (Thompson  and  Mead  1982).  Similar  extralocal  occurrences  of  Och- 
otona are  documented  as  late  as  7000  yr  B.P.  in  eastern  Oregon  (Grayson  1981,  in 
press).  Grayson  has  argued  that  the  timing  of  extinction  of  these  relictual  small  mammal 
(and  possibly  some  amphibian  and  reptile)  populations  was  in  large  part  determined 
by  the  size  and  distribution  of  habitat  islands  and  by  the  size  of  the  species  population 
on  the  given  habitat  island.  It  is  also  possible  that  the  mid-Holocene  period  of  warmer- 
than-present  temperatures,  seen  in  the  elevational  raising  of  upper  treeline  on  the 
Snake  Range  and  elsewhere  in  the  Great  Basin  (LaMarche  1973,  LaMarche  and  Moo- 
ney  1972)  may  have  reduced  the  size  of  montane  habitat  islands  and  accelerated  the 
rate  of  extinction  of  animal  populations. 

The  faunal  account  presented  here  is  the  first  detailed  account  of  amphibians  and 
reptiles  from  late  Pleistocene  and  early  Holocene  age  deposits  in  the  interior  Great 
Basin;  in  addition,  the  assemblage  has  provided  additional  data  on  mammalian  species. 
The  Phenacomys  cf.  intermedins  reported  here  is  the  first  late  Pleistocene  record  of 
the  genus  in  the  Great  Basin.  Because  some  of  the  fauna  (especially  Ochotona)  was 
recovered  from  packrat  middens,  it  is  associated  directly  with  a  reconstruction  of  the 
local  flora  based  on  plant  macrofossils.  To  our  knowledge,  similar  plant  and  animal 
associations  have  not  been  pursued  in  this  detail  for  the  Great  Basin. 

The  research  in  Smith  Creek  Canyon  over  the  last  50  years  has  illustrated  the 
wealth  of  information,  mainly  faunal,  available  in  dry  cave  deposits  of  the  Great  Basin. 
With  the  recent  surge  of  packrat  midden  analyses  in  Smith  Creek  Canyon,  a  whole 
new  aspect  of  late  Pleistocene  and  Holocene  community  reconstruction  has  emerged. 
Faunal  assemblages  can  now  be  found  in  and  associated  temporally  with  the  wealth  of 
flora  data  recoverable  from  packrat  middens. 


25 


Acknowledgments 

Our  sincere  thanks  goes  to  Donald  K.  Grayson  for  his  helpful  consultations  and 
encouragement  in  our  study  of  the  Great  Basin  natural  history.  Gerald  R.  Smith  kindly 
provided  the  identification  of  the  fish  remains  and  his  personal  comments.  Jeffrey  S. 
Green  provided  the  pygmy  rabbit  fecal  pellets  for  our  dung  analysis.  We  also  thank 
Susanne  Miller  and  Alan  Bryan  for  allowing  us  to  study  the  Smith  Creek  Cave  her- 
petofauna  from  previous  excavations,  and  Donald  R.  Tuohy  for  his  field  assistance  and 
guidance.  Radiocarbon  dates  were  provided  by  the  University  of  Arizona,  the  Uni- 
versity of  Waikato,  and  Geochron  Laboratories.  Research  was  funded  by  National 
Science  Foundation  Grant  DEB75- 13944  to  Paul  S.  Martin.  Special  thanks  are  due  to 
Marna  A.  Thompson  and  Eugene  Hattori  for  helping  with  field  collections,  to  Emilee 
Mead  for  the  photography,  and  Deborah  Gaines  for  typing. 


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tocene and  Holocene  mammal  fauna  of  Wyo- 
ming. Pages  78-87  in  Michael  Wilson  (editor). 
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26 


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Jones,  J.  Knox.  Dilford  C.  Carter,  and  Hugh  H. 
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Martin,  Paul  S.  1967.  Prehistoric  overkill.  Pages 
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Mawby,  John  E.  1967.  Fossil  vertebrates  of  the 
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H.  M.  Wormington  and  Dorothy  Ellis  (edi- 
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Nevada  State  Museum,  Anthropological  Pa- 
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Robinson,  Michael  D.,  and  Thomas  R.  Van  De- 
vender.  1973.  Miocene  lizards  from  Wyoming 
and  Nebraska.  Copeia  1973:698-704. 

Smith.  Gerald  R.  1978.  Biogeography  of  inter- 
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.  W.   L.   Stokes,  and   K.   F.  Horn.    1968. 

Some  late  Pleistocene  fishes  of  Lake  Bonne- 
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Stebbins,  Robert  C.  1966.  A  field  guide  to  western 
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Stock,  Chester.  1936.  A  new  mountain  goat  from 
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Bulletin  of  the  Southern  California  Academy 
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Tanner.  Wilmer  W.  1978.  Zoogeography  of  rep- 
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361-380  //;  Donald  R.  Tuohy  and  Doris  L. 
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Eariy  Holocene  and  late  Pleistocene  amphib- 
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middens.  Copeia  1978:464-475. 


TRANSACTIONS 
OF  THE  SAN  DIEGO 
SOCIETY  OF 
NATURAL  HISTORY 


Volume  20  Number  2  pp.  27-55         25  January  1983 


Fossil  decapod  crustaceans  from  the  Lower  Cretaceous,  -  ^    " 

Glen  Rose  Limestone  of  Central  Texas 

Gale  A.  Bishop 

Department  of  Geology  and  Geography,  Georgia  Southern  College. 
Stalesboro,  Georgia  30460  USA 

Abstract.  The  Lower  Albian  carbonate  rocks  of  the  Glen  Rose  Limestone  of  Central  Texas  contain 
a  diverse  decapod  fauna  dominated  by  Protocallianassa  species  with  10  other  taxa  represented  by  fewer 
specimens.  The  fauna  consists  of  Protocallianassa  species,  P.  klofi,  new  species,  Pagurus  handerensis 
Rathbun  1935,  Roemerus  robustus.  new  genus  and  species,  Palaeodromites  naglei.  new  species,  Diaulax 
roddai.  new  species,  Hillius  yoimgi,  new  genus  and  species,  Dioratiopus  scotti,  new  species,  Pseudo- 
necrocarcinus  stenzeli,  new  species,  Prehepatus  hodgesi,  new  species,  and  Torynommal  densus,  new 
species. 

Introduction 

The  remains  of  fossil  decapods  have  been  among  the  most  rarely  reported  fossils 
in  the  Glen  Rose  Limestone  of  Central  Texas.  Rathbun  (1935:35)  described  Pagurus 
handerensis  from  a  single  fragmental  specimen.  Stenzel  (1945:435)  revised  that  descrip- 
tion on  the  basis  of  several  additional  specimens  and  mentioned  that  other  decapods 
were  associated  with  the  P.  handerensis  remains  described  by  him.  "Among  the  other 
chelae  found  by  Mr.  Watkins  at  the  same  locality  there  are  some  which  obviously  can 
have  nothing  to  do  with  Pagurus,  because  they  belong  to  other  well-established  and 
unrelated  genera  such  as  Callianassa""  (Stenzel  1945:437). 

The  fauna  described  here  is  important  because  it  testifies  to  the  presence,  abun- 
dance, and  diversity  of  the  decapod  fauna  of  the  Glen  Rose  Limestone.  This  fauna 
adds  to  our  knowledge  of  the  Early  Cretaceous  decapod  fauna  of  North  America.  A 
diverse  fauna  of  Late  Albian  age  has  been  described  by  Rathbun  (1935). 

The  Glen  Rose  Limestone 

The  Glen  Rose  Limestone  is  composed  of  beds  of  limestone  that  are  resistant  to 
weathering  alternating  with  less  resistant  marls,  giving  rise  to  a  characteristic  stair-step 
topography  (Fig.  1).  The  presence  of  mud  cracks,  bored  bedding  planes,  bedding  planes 
with  encrusting  oysters,  algal  mat  laminations,  ripple  marks,  dinosaur  trailways,  and 
plant  debris  is  evidence  for  supratidal,  intertidal.  or  shallow  subtidal  conditions.  Beds 
of  lime  muds  containing  echinoids,  miliolid  foraminifera,  infaunal  pelecypods,  corals 
and  numerous  gastropods  imply  marine  conditions  prevailed  at  times.  The  interbedding 
of  these  sediments  points  to  a  depositional  system  of  shallow  marine  lagoons  with 
numerous  small  islands  or  rapidly  prograding  supratidal  areas  in  protected  lagoons 
behind  a  reef-like  barrier  (Winter  1962,  Hendricks  and  Wilson  1967:5,  Stricklin  et  al. 
1971,  Young  1972:1). 

The  Glen  Rose  Limestone  of  Central  Texas  contains  a  scant  ammonoid  fauna 
which  allows  correlation  with  the  European  section  (Young  1972:1 1,  1974:179).  The 
Glen  Rose  Limestone  is  divided  into  upper  and  lower  members  by  a  bed  (or  zone  of 
beds)  containing  numerous  steinkems  of  the  bivalve  Corhula  (Stricklin  et  al.  1971:23). 


28 


Figure  1.     Lithostratigraphy  and  biostratigraphy  of  the  Glen  Rose  Limestone  (after  Young  1974). 


The  '"Corbula'''  bed  is  used  in  this  study  as  a  stratigraphic  marker  to  locate  the  strati- 
graphic  position  of  each  collection.  All  of  the  Glen  Rose  Limestone  below  the  Corbula 
bed  and  to  a  level  of  about  41  m  (135  ft)  above  it  are  considered  to  be  Early  Albian 
by  Young  (1974:176).  This  includes  all  of  the  specimens  described  in  this  study. 


Localities 

Most  specimens  of  decapods  have  been  collected  from  3  localities  (Fig.  2).  The 
specimens  are  from  several  collections:  (GAB  =  Gale  A.  Bishop;  SDSNH  =  San  Diego 
Natural  History  Museum;  WSA  =  W.  S.  Adkins;  UT  =  University  of  Texas  [Austin]). 

The  Nagle  Locality  (GAB  27).— This  exposure  was  extensively  collected  by  J.  S. 
Nagle  during  the  early  1960s.  It  is  situated  at  the  junction  of  Highways  290  and  281, 
about  9.7  km  (6  mi)  south  of  Johnson  City,  Blanco  Co.,  Texas  (Fig.  3). 

Decapods  occur  throughout  about  9  m  (30  ft)  of  Glen  Rose  Limestone  but  are 
more  abundant  in  an  interval  just  below  a  Corbula  bed  and  in  a  biomicrite  about  7.6 
m  (25  ft)  above  the  Corbula  bed  (Fig.  4). 

Interesting  assemblages  of  microscopic  claws  and  dactyli  were  recovered  from 
samples  taken  at  points  indicated  in  the  measured  section  (Fig.  4).  The  microscopic 
decapod  material  is  particularly  abundant  in  the  Salenia  texana  marl. 

The  associated  fauna  consists  of  numerous  steinkerns  of  bivalves  and  gastropods. 

Boerne-Sisterdale  Locality  {GAB  25).  — This  exposure  is  a  road  cut  on  Texas  Farm 
Road  1376  at  the  top  of  a  hill  (Fig.  5)  12  km  (7.5  mi)  north  of  Boerne,  Kendall  Co., 
Texas.  Approximately  9  m  (30  ft)  of  Glen  Rose  Limestone  is  exposed  in  the  road  cut 
(Fig.  6).  The  road  cut  is  about  37  m  (120  ft)  above  exposures  of  the  Corbula  bed  at 
the  Hodges  Range  Section  (GAB  26)  1.6  km  (1  mi)  to  the  northwest,  and  probably 


29 


Figure  2.     Index  map  of  major  collecting  localities  in  Central  Texas. 

near  the  top  of  the  fourth  open  shelf  unit  of  Young  (1974:177).  Numerous  remains  of 
decapods  were  collected  from  a  biomicrite  near  the  bottom  of  the  road  cut  (at  arrow 
in  Fig.  6). 

Decapod  remains  occur  throughout  an  interval  3  m  (10  ft)  in  thickness  but  are 
most  abundant  in  a  46  cm  ( 1 8  in)  bed  of  fossiliferous  biomicrite.  The  specimens  weather 
out  of  the  platey  marl  and  accumulate  on  the  ledge  formed  by  the  underlying  resistant 
bed. 

The  entire  fauna  has  not  been  investigated  in  detail  but  numerous  specimens  of 
Homomya  are  present  and  preserved  in  living  position.  Other  elements  of  the  fauna 
include  numerous  pelecypods,  gastropods,  and  masses  of  serpulid  worm  tubes. 


ROAD 


Figure  3.  Topographic  map  of  Nagle  locality  (GAB  21)  9.7  km  (6  mi.)  south  of  Johnson  City  at  junction 
of  U.S.  Highways  290  and  281,  Blanco  Co.,  Texas.  U.S.G.S.  Monument  Hill  Quadrangle,  scale  1:24  000, 
contour  interval  20  ft  (6.1  m). 


30 


30- 


20- 


10- 


L  i  me  s  lo  ne 


~^T-      Nodular    Ls 


Dolomi  te 


"I_~_"       C  I  a  y  s  f  o  n  e 


O  Decapods 


*  Mi  crocrustaceans 


I.  j_  L I  4^-L__l_-lJ C o r b u I g    Bed 

Figure  4.     Measured  section  in  Glen  Rose  Limestone  at  Nagle  locality  (GAB  21). 


(7496 


Figure  5.  Topographic  map  of  Boeme-Sisterdale  (GAB  25)  and  Hodges  Range  (GAB  26)  localities,  12.9 
km  (8  mi)  north  of  Boeme  on  Farm  Road  1376,  Kendall  Co.,  Texas.  U.S.G.S.  Sisterdale  Quadrangle  Scale 
1:24  000,  contour  interval  20  ft  (6.1  m). 


31 


Figure  6.     Photograph  of  Boeme-Sisterdale  outcrop  (GAB  25),  decapod  interval  marked  by  arrow.  Vertical 
scale  of  foreground  approximately  1  cm  =  2  m  (1"  =  17'). 

The  Hodges  Range  Locality  {GAB  26).— The  Hodges  Ranch  Locahty  (Fig.  5)  is 
situated  on  the  third  tributary  of  Wasp  Creek  west  of  Texas  Farm  Road  1376  about 
12.9  km  (8  mi)  north  of  Boerne,  Kendall  Co.,  Texas.  Specimens  were  collected  from 
a  soft  marl  that  contained  Salenia  texana]\xs\  below  a  Corbula  bed  (Fig.  1).  The  locality 
also  produced  numerous  sea  urchins  and  some  crinoids. 

The  outcrop  is  a  semi-barren  gentle  slope  (Fig.  7)  just  above  a  small  creek.  The 
surface  is  covered  by  great  numbers  of  fossils,  especially  the  heart  urchin  Enallaster 
obliquatus.  Other  faunal  elements  include  oysters,  gastropods,  and  bivalves. 

The  decapod  part  of  the  fauna  from  this  locality  is  almost  exclusively  dominated 
by  claws  of  the  hermit  crab,  Pagurus  banderensis.  A  collection  of  gastropod  steinkerns 
was  made  to  examine  the  possibility  of  preserved  pagurids  within  the  lithified  mud  of 
the  steinkerns.  No  evidence  of  decapod  exoskeleton  was  observed  in  the  steinkerns.  In 
fact,  the  steinkerns  were  packed  with  numerous  shells  and  shell  fragments  mixed  with 
carbonate  mud.  This  suggests  a  fair  amount  of  washing  by  currents  or  bioturbation 
after  the  shells  were  last  occupied  (by  hermit  crabs?)  which  would  destroy  evidence  of 
any  such  occupation. 

Other  localities  from  which  Glen  Rose  decapods  have  been  collected  are  listed 
below: 

1.  Bandera-Pipe  Creek  Road  (Texas  Highway  16)  1.6  km  (1  mi)  east  of  Bandera, 
Bandera  Co.,  Texas;  Salenia  texana  zone.  Rathbun  1935:39. 


Figure  7.     Photograph  of  Hodges  Range  locality  (GAB  26).  Collecting  bag  in  center  38  cm  (1 5  m.)  high). 


32 


B 


H 


Figure  8.  Line  drawings  of  decapods  from  the  Glen  Rose  Limestone,  Central  Texas.  A,  Palaeodromites 
naglei  Bishop;  B,  Pseudonecrocarcinus  stenzeli  Bishop;  C,  Hillius  youngi  Bishop;  D,  Dioratiopus  scotti  Bishop; 
E,  Diaulax  roddai  Bishop;  F,  Prehepatus  hodgesi  Bishop;  G,  Roemems  wbustus  Bishop;  H,  Pagurus  ban- 
derensis  Rathbun  1935;  \,  Protocallianassa  klofi  Bishop;  J,  Torynomma?  densus  Bishop. 


2.  Bandera-Pipe  Creek  Road  (Texas  Highway  16)  3.2  km  (2  mi)  east  of  Bandera; 
Salenia  texana  zone.  Stenzel  1945:437. 

3.  "0.15  mile  [0.24  km]  south  of  state  Highway  No.  29  (Burnet-Austin  road)  and 
0.07  mile  [0.1 1  km]  east  of  the  Southern  Pacific  Railroad  track  1.42  miles  [2.27 


33 


Table  1.    Tabulation  of  decapod  collected  from  the  Glen  Rose  Limestone. 


GAB-2 1 

GAB-25 

GAB-26 

Other 

Total 

Protocallianassa  sp." 

52 

750 

0 

Yes" 

802+ 

P.  klofi 

1 

14 

0 

0 

15 

Pagurus  banderensi^ 

1 

12 

10 

7 

30 

Roemerus  robustus 

0 

2 

1 

1 

4 

Palacodromites  naglei 

1 

8 

0 

0 

9 

Diaulax  roddai 

0 

3 

0 

0 

3 

Hillius  youngi 

0 

1 

0 

0 

1 

Dioratiopus  scotti 

0 

0 

0 

1 

1 

Pseudonecrocarcinus  stenzeli 

0 

6 

0 

0 

6 

Prehepatus  hodgesi 

1 

27 

2 

0 

30 

Torynomma?  densus 

0 

2 

0 

0 

2 

"  Nearly  whole  palms. 

''  Numerous  other  localities  yield  Protocallianassa. 

"  Does  not  include  movable  fingers. 


8. 


km]  airline  distance  east  of  the  courthouse  in  Burnet,  Burnet  Co.,  Texas."  Stenzel 

1945:437. 

Junction  of  Highways  16  and  689,  2.6  km  (1.6  mi)  east  of  Bandera;  Salenia 

texana  zone;  collected  by  Mrs.  Henry  W.  Sebesta,  sent  to  Keith  Young  by 

George  His. 

Blackman  Ranch,  Little  Blanco  River,  Blanco  Co.,  Texas;  WSA  16215  collected 

by  A.  W.  Cleaves. 

Bluff  of  Blanco  River,  4.8  km  (3  mi)  west  of  Blanco,  Blanco  Co.,  Texas;  Salenia 

texana  zone  1.5  m  (5  ft)  below  Corbida  Bed;  UT  45488  (2  specimens)  collected 

by  N.  B.  Waechter. 

Shingle  Hills  Section,  Travis  Co.,  Texas;  Salenia  texana  zone;  collected  by  G. 

L.  Dawe. 

From  a  limestone  ledge  about  1.5  m  (5  ft)  above  water  level  at  Jacob's  Well,  a 

spring  on  Cypress  Creek,  1.5  km  (3.2  mi)  northwest  of  Wimberley,  Hays  Co., 

Texas,  about  55  m  ( 1 80  ft)  below  the  Corbula  Bed,  Lower  Glen  Rose  Limestone 

(calculated  from  Young  1974). 


Fauna 

The  decapod  fauna  (Fig.  8)  of  the  Glen  Rose  Limestone  consists  of  1 1  taxa  (Table 
1).  Protocallianassa  sp.  is  the  most  abundant  taxon,  and  Pagurus  and  Prehepatus  are 
second  most  abundant.  The  decapod  fauna  at  each  locality  forms  only  a  part  of  a  much 
larger  fauna  dominated  by  molluscs. 

Preservation.— ThQ  fossil  decapods  of  the  Glen  Rose  Limestone  are  found  as  dis- 
articulated fragments.  Claws  are  most  abundant  but  a  few  carapaces  are  present.  The 
condition  of  the  remains  at  the  time  of  final  entombment  can  not  be  determined.  The 
presence  of  numerous  decapod  remains  is  probably  due  to  the  similarity  of  the  min- 
eralized exoskeleton  and  surrounding  carbonate  rocks.  The  mineralized  exoskeleton  is 
often  only  represented  by  a  chalky  carbonate  material  that  weathers  readily.  This  gives 
rise  to  many  steinkerns  (internal  molds).  Steinkerns  of  brachyurans  reflect  surface 
morphology  but  certainly  differ  significantly  from  specimens  with  exoskeletons.  The 
finger  tips  of  claws  are  often  filled  with  sparry  calcite  signifying  that  they  were  not 
completely  filled  with  mud  at  the  time  of  burial. 

Decapods.— The  Glen  Rose  decapod  fauna  (Pis.  1-3)  gives  a  crude  measure  of  the 
diversity  of  decapods  in  North  America  in  the  Early  Albian.  Because  of  the  small  size 
of  most  of  these  decapods,  they  probably  are  seldom  collected  and,  hence,  the  antici- 
pated total  decapod  fauna  certainly  exceeds  that  described  here.  This  fauna  is  the  earliest 
Cretaceous  decapod  fauna  described  from  North  America. 


34 


The  presence  and  abundance  of  hermit  crabs  (Pagnnis)  in  these  collections  is  due 
to  their  size,  original  abundance  in  the  living  fauna,  and  their  heavy  mineralization. 
The  postulated  shallow-lagoonal  environment  is  very  compatable  with  these  shallow- 
water  decapods.  One  specimen  o^  Pagurus  (PI.  2,  fig.  17)  has  an  oyster  attached  to  its 
claw.  The  lack  of  hermit  crab  fossils,  except  claws  of  pagurids,  is  due  to  *the  thin 
exoskeleton  everywhere  but  on  the  heavily  mineralized  claws  that  function  as  an  oper- 
culum to  close  the  ''borrowed"  gastropod  shells  in  which  hermit  crabs  live.  Roemerus 
robustus  was  also  probably  a  hermit  crab. 

Because  this  is  a  large,  new  fauna  there  are  several  extensions  of  geographic  range 
of  the  particularly  well-known  taxa  from  the  Albian  of  England  (Wright  and  Collins 
1972).  Palaeodromites  nag/ei,  Diaulax  roddai,  and  Pseudonecrocarcinus  stenzeli  all 
extend  the  geographic  ranges  of  the  genera  from  Europe  into  North  America.  Tory- 
nomma?  densiis  extends  the  range  of  that  genus  from  Australia  to  North  America. 
Dioratiopus  scotti  is  the  second  described  species  of  this  taxon  in  North  America  (the 
other,  Dioratiopus  dawsonensis  (Bishop  1973)  is  from  the  Maestrichian  of  Montana). 
A  third  species  is  present  on  the  north  flank  of  the  Black  Hills,  low  in  the  Pierre  Shale. 

Hilliiis  youngi  is  so  far  known  from  1  steinkern.  It  is  hoped  that  additional  material 
will  substantiate  the  basic  diagnostic  features  of  this  poorly  represented  taxon. 

Prehepatus  hodgesi  joins  other  members  of  the  genus,  P.  cretaceous  and  P.  paw- 
pawensis  from  the  Early  Cretaceous  of  Texas,  and  P.  dilksi  from  the  Late  Cretaceous 
Merchantville  Fm.  of  Maple  Shade,  New  Jersey.  The  pattern  and  variability  of  orna- 
menting tubercles  on  these  little  claws  is  extremely  interesting.  Possibly  the  variation 
may  be  due  to  sexual  dimorphism  as  in  the  fiddler  crabs  and  perhaps  may  even  have 
had  some  ritualistic  or  actual  function  in  this  crab's  behavior. 

Callianassids.  — The  most  abundant  decapod  remains  found  in  this  collection  and 
throughout  the  Glen  Rose  Limestone  are  thalassinids  belonging  to  Protocallianassa, 
Callianassa,  Axius,  Jaxea,  or  some  other  closely  related  taxon.  Only  the  chelae  are 
represented  in  the  collections  from  the  Glen  Rose  Limestone. 

The  thalassinids  are  differentiated  on  the  basis  of  carapace  morphology,  and  che- 
lation of  walking  legs.  Within  a  given  taxon  (such  as  Callianassa)  sexual  dimorphism, 
differing  morphology  of  right  and  left  claws,  intraspecific  variation,  and  changes  in 
morphology  in  different  instars  are  all  probable  (Rathbun  1935:29).  These  factors  seem 
to  have  combined  to  yield  a  baffling  spectrum  of  chelae  morphology.  The  claws  found 
in  the  Glen  Rose  exhibit  a  wide  variation.  Morphotypes  can  be  established  easily  but 
as  the  numerous  specimens  are  examined  the  morphotypes  become  impossible  to 
maintain.  The  claws  will  be  analyzed  again  when  a  graphic  analyzing  computer  system 
becomes  available. 

Microcrustaceans.  — The  presence  of  crustacean  micro  fossils  in  the  Glen  Rose 
Limestone  was  pointed  out  to  me  by  Peter  Rodda  while  curating  the  collection  of  the 
Texas  Bureau  of  Economic  Geology.  Subsequently,  microfossil  residues  were  prepared 
from  the  marls  at  each  locality  and  numerous  other  Glen  Rose  localities.  They  dem- 
onstrated a  diverse  and  abundant  crustacean  microfauna  consisting  of  claws  and  dactyli 
(PI.  3,  fig.  44). 


Plate  I 

Figures  1-2.  Diaulax  roddai,  1,  Dorsal  view  of  carapace,  xl.O;  2,  Dorsal  view  stereo,  x2.0  SDSNH  23640 
(orig.  GAB  25,  specimen  2).  Figures  3-5.  Pseudonecrocarcinus  stenzeli.  Dorsal  view  of  latex  cast  of  carapace; 
3,  xl.O;  4,  stereo,  X2.0;  5,  view  of  impression  of  carapace  in  limestone,  Xl.O,  SDSNH  23641  (orig.  GAB 
25,  specimen  8).  Figures  6-7.  Dioratiopus  scotti,  6,  Dorsal  view  of  carapace,  Xl.O;  7,  Dorsal,  stereo,  X2.0, 
SDSNH  23642  (orig.  GAB  27,  specimen  1).  Figure  8-1  1.  Hillius  voungi,  8,  Carapace,  Xl.O;  9,  Carapace, 
stereo,  X2.0;  10,  Anterior  of  Carapace  X2.0;  1  1,  Right  side  of  carapace,  X2.0,  SDSNH  23643  (orig.  GAB 
25,  specimen  3).  Figures  12-17.  Palaeodromites  naglei,  2-13,  Carapace,  Holotype  SDSNH  23644  (orig. 
GAB  25,  specimen  1);  12,  Xl.O;  13,  stereo,  X2.0;  14-17,  Carapace  features  of  specimen  SDSNH  23645  (orig. 
GAB  21,  specimen  21)  collected  by  Nagle;  14,  Dorsal,  Xl.O;  15,  Dorsal  of  carapace,  x2.0. 


36 


Table  2.    Measurements  in  millimeters  of  the  major  claw  of  Protocallianassa  klofi. 


Propodal 

Right  or 

length 

Palm  length 

Palm  height 

Specimen* 

left 

(mm) 

(mm) 

(mm) 

25-67 

R 

13.7 

9.8 

7.7 

25-160 

R 

8.0 

7.0 

25-153 

L 

9.2 

8.6 

SDNHM  23665  (=25-152) 

R 

9.2 

8.5 

25-87 

L 

10.7 

9.8 

25-116 

R 

15.6 

11.0 

9.2 

25-133 

L 

10.8 

9.3 

25-124 

L 

10.9 

7.0 

6.0 

25-123 

R 

9.8 

6.6 

25-144 

R 

25-127 

R 

8.2 

5.8 

25-164 

R 

8.1 

6.8 

5.4 

25-136 

L 

8.5 

6.7 

5.8 

*  Specimens  in  the  tables  are  listed  by  locality  number  followed  by  the  specimen  number  (e.g.  25-67  is 
locality  GAB25,  specimen  67). 


Systematic  Paleontology 

Order  Decapoda  Latreille  1 803 

Suborder  Pleocyemata  Burkenroad  1963 

Infraorder  Anomura  H.  Milne-Edwards  1832 

Superfamily  Thalassinoidea  Latreille  1831 

Family  Callianassidae  Dana  1852 

Subfamily  Protocallianassinae  Beurlen  1930 

Genus  Protocallianassa  Beurlen  1930 

Type  species.  — Callianassa  archiaci  A.  Milne-Edwards  1860  by  original  designa- 
tion. 

Z)/a^«05/5.— "Carapace  with  linea  thalassinica\  first  pereiopods  with  well  developed 
chelae,  hetereochelous;  abdomen  with  pleura  developed  on  second  to  sixth  somites; 
uropods  without  diaeresis  .  .  .  (Single  chelae  are  hardly  distinguishable  from  those  of 
Protaxius  or  Callianassa).'"'  (Glaessner  1969:478). 

Protocallianassa  klofi  new  species 
PI.  3,  figs.  41-43;  Fig.  81;  Tab  2 

Type.—T\yQ  holotype,  a  right  major  propodus  (orig.  GAB  25,  specimen  152)  is 
deposited  in  the  San  Diego  Natural  History  Museum  (SDNHM  23665). 


Plate  2 

Figures  1-16.  Right  chelae  of  Pagurusbanderensis.  1-5,  Specimen  SDSNH  23646  (orig.  GAB  25,  specimen 
7).  1,  Outer  face;  2,  Inner  face,  3,  Bottom,  and  4,  Top;  x2.0.  5,  Outer  face,  Xl.O.  6-8,  Specimen  SDSNH 

23647  (orig.  GAB  2 1 ,  specimen  1 7).  6.  Outer  face  and  7,  Inner  face;  X2.0;  8,  Outer  face,  X 1 .0.  9-12,  Specimen 
UT  45473  (Univ.  Texas,  Austin).  9,  Outer;  1 0,  Inner;  1 1 ,  Lower;  and  1 2  Top,  X 1 .0.  1 3-1 5,  Specimen  SDSNH 

23648  (orig,  GAB  26,  specimen  5).  13,  Outer;  14,  Inner;  and  15,  TopX2.0.  16,  Specimen  WSA  16215  (Univ. 
Texas,  Austin),  Outer  face  with  walking  leg,  Xl.O.  Figures  17-25.  Left  chelae  of  Pagnrus  banderensis.  17, 
Specimen  SDSNH  23649  (orig.  GAB  26,  specimen  3),  Inner  face  of  left  claw  with  attached  oyster,  X2.0.  18, 
Specimen  SHSNH  23650  (orig.  GAB  26,  specimen  4).  Outer  face  of  propodus,  Xl.O.  19,  Specimen  UT 
45488.  Outer  face  of  claw,  Xl.O.  20-21,  Specimen  SDSNH  23651  (orig.  GAB  25,  specimen  1  1),  Outer  (20), 
Inner  (21),  faces  of  nearly  complete  minor  claw,  Xl.O.  22-25,  Specimen  SDSNH  23652  (orig.  GAB  26, 
specimen  1),  Outer  (22),  Inner  (23),  Bottom  (24),  and  Top  (25)  views,  X2.0.  Figures  26-31.  Dactyli  of 
Pagurus  banderensis.  26-29,  Specimen  SDSNH  23653  (orig.  GAB  26,  specimen  12),  Dactylus  of  walking 
leg,  X2.0.  30-31,  Specimen  SDSNH  23654  (orig.  GAB  26,  specimen  1 1),  Dactylus  of  major  claw,  30,  Side 
Xl.O  and  31,  Occlusional  surface,  X2.0. 


38 


Table  3.    Height  (H),  length  (L),  thickness  (T)  in  millimeters,  and  height/thickness  (H/T)  ratios  of  claws  of 
Pagurus  banderensis. 


Height 

Length 

Thickness 

(mm) 

(mm) 

(mm) 

H/T* 

Right  Claw 

25-22 

3.7 

5.5 

1.9 

1.90 

25-23 

4.2 

7.3 

2.5 

1.70 

SDNHM  23647  (= 

=21- 

-17) 

4.9 

8.6 

2.3 

2.09 

25-25 

5.4 

9.8 

3.6 

1.52 

25-24 

5.6 

7.5* 

3.4 

1.65 

SDNHM  23646  (= 

=25- 

-7) 

6.0 

9.74+ 

4.0 

1.51 

25-26 

10.2 

15.7* 

6.3 

1.62 

25-28 

10.6 

19.2 

26-14 

10.8 

7.2 

1.49 

25-27 

11.9 

18.7 

5.7 

2.09 

SDNHM  23648  (= 

=26- 

-5) 

13.8 

23.8 

9.0 

1.54 

Stenzel 

14.2 

26.7 

9.0 

1.57 

UT  45473 

16.3 

32.4 

9.3 

1.75 

26-16 

17.0 

11.1 

1.53 

WSA  16215 

17.9 

30.2* 

11.4 

1.57 

26-6 

19.7 

11.8 

1.67 

26-19 

20.0 

11.8 

1.69 

Stenzel** 

22.8 

10.8 

2.11 

Left  Claw 

SDNHM  23651  (= 

=25- 

-11) 

7.1 

14.3 

4.47 

1.56 

Stenzel 

7.4 

16.2 

5.1 

1.45 

25-31 

7.5 

10.8* 

4.94 

1.51 

25-30 

7.5* 

5.12 

1.46 

25-29 

7.6 

15.2* 

26-15 

10.1 

6.27 

1.61 

26-7 

10.3 

16.8+ 

7.34 

1.40 

SDNHM  23652  (= 

=26- 

-1) 

10.9 

17.9 

6.50 

1.67 

SDNHM  23649  (= 

=26- 

-3) 

11.4 

7.8 

1.46 

26-2 

11.5 

7.4 

1.55 

UT  45488 

11.8 

20.5* 

8.86 

1.33 

UT  45488 

13.9 

19.8* 

8.90 

1.56 

*  Calculated  before  rounding  thickness  measurements  from  hundreths  of  a  mm. 
**  Stenzel's  holotype  of  P.  travisensis. 


Plate  3 

Figures  1-19.  Prehepatus  hodgesi.  1-5,  Holotype,  Specimen  SDSNH  23655  (orig.  GAB  25,  specimen  14); 
Outer  surface  of  propodus,  x  1 .0,  2-5,  Outer,  Inner,  Top,  and  Distal  views  of  propodus,  X2.0.  6-11,  Specimen 
SDSNH  23656  (orig.  GAB  26,  specimen  9);  6,  Outer  face  of  propodus,  xl.O;  7-1 1,  Outer,  Inner,  Top,  Bottom, 
and  Distal  views,  x2.0.  12-13,  Specimen  SDSNH  23657  (orig.  GAB  25,  specimen  16);  12,  Inner  face  and 
13,  Outer  face  of  propodus,  X2.0.  14-16,  Specimen  SDSNH  23658  (orig.  GAB  25,  specimen  137),  articulated 
carpus  and  propodus  in  oblique  (14),  Front,  (15),  and  Top  view  (16),  X2.0.  17,  Specimen  SDSNH  23659 
(orig.  GAB  2 1 ,  specimen  22),  complete  propodus  and  disarticulated  dactylus  in  front  view,  X2.0.  1 8,  Specimen 
SDSNH  23660  (orig.  GAB  25,  specimen  18),  left  propodus  in  front  view,  X2.0.  19,  Specimen  SDSNH  23661 
(orig.  GAB  25,  specimen  15),  left  propodus  in  front  view,  X2.0.  Figures  20-31.  Roemems  robustus.  20- 
25,  Specimen  SDSNH  23662  (orig.  GAB  26,  specimen  8).  20,  Outer  view,  Xl.O.  21-25,  Views  of  outer  face, 
inner  face,  bottom,  distal  end,  and  top  of  left  propodus.  26-31,  Holotype,  Specimen  UT  45704.  31,  Outer 
view,  Xl.O.  26-30,  Outer,  inner,  top  and  bottom  views,  X2.0.  Figures  32-40.  Torynommal  densus.  32- 
35,  Paratype  Specimen  SDSNH  23663  (orig,  GAB  25,  specimen  10),  32,  Outer  view,  Xl.O.  33-35,  Outer, 
inner  and  distal  views  X2.0.  36^0,  Holotype,  right  propodus.  Specimen  SDSNH  23664  (orig.  GAB  25, 
specimen  5),  36,  outer,  Xl.O.  37-40,  Outer,  inner,  bottom,  and  top  views,  X2.0.  Figures  41- 
43.  Protocallianassa  klofi.  Holotype,  Right  major  propodus.  Specimen  SDSNH  23665  (orig.  GAB  25, 
specimen  152).  41-^3,  Outer,  distal,  and  inner  views,  X2.0.  Figure  44.  Slide  of  microcrustacean  appendage 
elements,  X3.25. 


40 


20 


10 


/ 


V 


^* 


/ 


/ 


.-"^^'^ 


\ 


\ 


0) 


•-0  ■ 

15 


■^5 


1  0  <- 


10- 
5-^ 


1  5 
0-* 


.0. 


B.  Thiclcnes  s 


10 


.,% 


.°^ 


0) 


4  0-*- 


1  o  — 

30-*- 


->-20 


^  30 

20  -« 1  O 


->•  40 
0-* 


A.    Length 


Figure  9.  Graph  of  A.  height  (mm)  vs.  length  (mm)  (bottom)  and  B.  height  (mm),  vs.  thickness  (mm)  of 
Pagurus  banderensis  Rathbun  1935.  Right  claws  plotted  as  dots  (read  thickness  or  length  on  upper  scale 
increasing  from  left  to  right)  and  left  claws  as  stars  (read  thickness  of  length  on  lower  scale  increasing  from 
right  to  left). 


Occurrence,  sample  size,  and  preservation.  — Fourteen  specimens  of  this  taxon  were 
collected  at  GAB  25  and  1  at  GAB  21.  Most  are  preserved  as  single  isolated  propodi 
with  chalky  exoskeleton  over  a  firm  micrite  filling. 

Etymo/og}^.  — Named  in  honor  of  L.  R.  KJof,  Texas  sedimentologist,  who  often 
exhibited  nocturnal  and  fossorial  habits. 

Description.  — Fropodus  flat,  broad,  nearly  twice  as  long  as  high.  Palm  rectangular, 
slightly  longer  than  high,  thin.  Upper  margin  bowed  slightly  into  convex  arch.  Proximal 
(carpal)  edge  nearly  vertical.  Distal  margin  slants  slightly  outward  to  top  of  short  fixed 
finger.  Two  sinuses  present  on  back  of  hand  along  margin;  uppermost  about  Vi  the 
distance  to  top  of  the  fixed  finger,  the  second  lies  immediately  above  fixed  finger.  Lower 
margin  convex  proximally  and  concave  beneath  base  of  fixed  finger.  Lower  proximal 
corner  produced  into  a  rounded  projection.  Propodus  convex  on  outer  face  (back  of 
hand),  nearly  flat  on  inner  face  (palm).  Convexity  of  outer  face  continues  onto  rounded 
upper  margin  which  overhangs  inner  side  (palm)  forming  a  shallow  depression  along 
top  of  inner  side  (palm)  parallel  to  upper  margin.  Another  shallow  depression  parallels 
the  wedge-shaped  lower  margin  on  palm.  Lower  margin  of  outer  side  produced  into  a 
narrow  keel  from  proximal  edge  almost  to  base  of  fixed  finger. 

Fixed  finger  short,  nearly  horizontal  and  turned  inward.  Most  large  specimens  with 
auxiliary  ridge  along  outer  edge  of  occlusional  surface  of  fixed  finger,  terminating  in 
tooth-like  projection.  An  oblique  ridge  runs  ofl' propodus  onto  fixed  finger  on  back  of 
hand  (outer  side  and  on  palm  [inner  side]). 


41 


Convex  outer  face  ornamented  by  4  or  6  large  hair  pits  along  ridge  running  onto 
finger.  Four  hair  pits  arranged  in  a  horizontal  row  just  above  level  of  sinus  immediately 
above  base  of  fixed  finger.  One  hair  pit  situated  just  above  uppermost  sinus.  Three  hair 
pits  form  broad-based  isosceles  triangle  just  below  proximal  edge  of  finger  ridge.  Four 
to  8  hair  slits  slant  upward  and  distally  along  lower  margin  just  above  fine  keel. 

Palm  (inner  face)  has  approximately  10  downward  and  distally  slanting  hair  slits 
along  upper  margin  just  beneath  overturned  angulaled  edge  where  flat  palm  meets  the 
convex  outer  face.  Approximately  15  hair  slits  slanting  upward  and  distally  situated 
along  lower  edge  of  palm. 

Comparison.— Protocallianassa  k/ofi  is  similar  to  P.  praecepta  Roberts  1962  but 
differs  from  it  by  having  a  relatively  shorter  palm,  a  rounded  lower  proximal  corner, 
and  lacking  the  ridge  at  the  base  of  the  fixed  finger  on  the  inner  face. 

Remarks.— The  minor  chela  of  this  taxon  is  not  as  yet  known.  No  pairs  of  chelae 
were  found  preserved  together  to  directly  tie  the  major  and  minor  chelae  to  one  another. 

Protocallianassa  sp. 

The  majority  of  the  specimens  referrable  to  Protocallianassa  comprise  a  highly 
variable  series  of  chelae.  Attempts  to  differentiate  morphotypes  failed  except  in  the 
case  of  P.  klofi  because  gradations  were  found  between  all  other  morphotypes  I  attempted 
to  establish. 

The  claws  vary  from  proximally  expanded,  through  rectangular,  to  nearly  oval  in 
shape.  The  cross-sectional  shape  varies  from  biconvex,  through  convex  on  the  other 
face,  to  spatulate.  The  fixed  fingers  are  usually  curved  slightly  inward.  Ornamentation 
by  hair  pits  is  extremely  variable. 

Superfamily  Paguroidea  Latreille  1803 

Family  Paguridae  Latreille  1802 

Subfamily  Pagurinae  Latreille  1802 

Genus  Pagurus  Fabricius  1775 

Type  species.  — '''Cancer  bernhardus  Linne  1758"  (on  official  list,  ICZN);  subse- 
quent designation  Latreille  1810  =Eupagurus  Brandt  1851  (type.  Cancer  bernhardus 
Linne'  1758;  subsequent  designation  Stimpson  1858)  (obj.).''  (Glaessner  1969:R479). 

Diagnosis.  — ""CheWped^  usually  dissimilar  and  unequal,  right  being  much  larger 
than  left,  very  rarely  subequal;  4th  periopods  subchelate."  (Glaessner  1969:R479). 

Pagurus  banderensis  Rathbun  1935 
PL  2,  figs.  1-31;  Figs.  8H,  9;  Tab.  3 

Pagurus  banderensis  Rathbun  1935,  p.  39,  PI.  9,  figs.  7,  8. 

Pagurus  banderensis  (Rathbun);  Stenzel  1945,  p.  435,  PI.  45,  figs.  7-15. 

Palaeopagurus  banderensis  {Ralhhun);  Roberts  1962,  p.  175. 

Occurrence. —Specimens  of  Pagurus  banderensis  have  been  collected  at  many  local- 
ities including  GAB  21,  GAB  25,  GAB  26,  and  Localities  1-7. 

Previous  descriptions.— See  Rathbun  1935:30  and  Stenzel  1945:435-437. 

Remarks.  — The  collections  made  at  GAB  25  and  GAB  26  give  the  first  suites  of 
specimens  of  P.  banderensis.  Height,  length,  and  thickness  data  were  gathered  (Table 
3)  and  are  presented  graphically  in  Fig.  9. 

Size  variation  is  much  greater  in  the  right  claw  than  in  the  left  claw.  This  gener- 
alization also  carries  over  to  their  morphology  and  ornamentation;  the  right  claws  are 
highly  variable  in  shape,  cross  section,  and  degree  of  granulation,  whereas  the  left  claws 
are  more  consistent  in  shape  and  ornamentation.  Small  right  (major)  chelae  tend  to 
have  a  straighter  lower  margin  which  becomes  gently  convex  as  size  increases.  There 
seems  to  be  a  tendency  for  a  single  row  of  upper  margin  granules  in  small  sizes  and  2 
distant  rows  (surrounded  by  many  smaller  granules)  in  larger  specimens. 

Two  specimens  (GAB  21.  specimen  17  and  GAB  25,  specimen  27)  are  thinner 


42 


Table  4.    Measurements  in  millimeters  of  claws  of  Roemerus  robustus. 


Total  length 

Height 

Palm  length 

Thickness 

(mm) 

(mm) 

(mm) 

(mm) 

Claw 

UT  45704 

10.5 

7.75 

6.75 

4.5 

keft 

SDNHM  23662  (=26-8) 

10.4 

8.9 

7.8 

5.1 

Left 

25-162 

6.7 

5.0 

4.7 

2.6 

Right 

25-163 

6.8 

5.1 

4.6 

3.2 

Left 

25-165 

6.3 

5.8 

8.3 

3.9 

Right 

than  the  others  (H/T-1.62)  and  might  have  been  called  P.  travisensis  (H/T-2.11)  by 
Rathbun.  I  consider  these  thinner  specimens  to  be  P.  banderensis  until  such  time  as 
sufficient  material  becomes  available  to  clearly  separate  them  as  P.  travisensis  upon 
the  basis  of  shape,  or  until  the  left  (minor)  claw  of  P.  travisensis  can  be  demonstrated 
to  exist. 

One  other  specimen  (GAB  25,  specimen  26)  is  different  enough  to  merit  special 
consideration.  The  upper  margin  is  narrow  and  the  claw  pear-shaped  in  cross  section. 
In  front  view  the  lower  margin  is  straight  and  the  upper  and  lower  margins  highly 
divergent. 

A  walking  leg  is  preserved  with  a  large  major  chela  (PI.  2,  fig.  16).  A  fragment  of 
a  dactylus  of  a  walking  leg  is  also  figured  (PI.  2,  figs.  26-29).  A  fairly  common  element 
in  the  collections  is  the  movable  finger  belonging  to  this  taxon  (PI.  2,  figs.  30-31). 

Family  Paguridae  Latreille  1802 
Subfamily  Uncertain 
Roemerus  new  genus 

Type  species.—  Roemerus  robustus  new  species. 

Diagnosis.— ChtXdiQ  elongate,  similar,  with  rectangular  palm,  outer  face  transversely 
convex,  inner  flat.  Carpal  and  dactyl  articulations  perpendicular  to  lower  margin  which 
is  straight  except  for  convexity  below  base  of  fixed  finger.  Fingers  short  and  turned 
slightly  inward.  Fixed  finger  has  at  least  1  proximal  tooth.  Tip  of  movable  finger  overlaps 
fixed  finger  in  smaller  claws  and  closes  onto  outer  edge  of  tip  in  large  claws.  Surface 
sparsely  covered  with  large,  low  granules  which  become  numerous  and  prominent  on 
lower  edge.  The  upper  surface  surmounted  by  a  low,  oblique  ridge  accentuated  on  the 
proximal  inner  face  by  a  few  low  granules  and  becoming  less  conspicuous  as  it  runs 
toward  the  top  center  of  the  distal  margin  of  the  claw. 

Etymology.  — \n  honor  of  Ferdinand  Roemer,  pioneer  geologist  and  paleontologist 
of  Texas  and  Mexico. 

Comparison.— This  claw  is  similar  to  Palaeopagurus  Van  Straelen  1925  but  differs 
in  being  more  rectangular,  having  a  more  vertical  distal  margin,  and  having  a  slightly 
upturned  fixed  finger  instead  of  slightly  downtumed.  Roemerus  robustus  is  easily  dis- 
tinguished from  Pagurus  banderensis  Rathbun  1935  by  its  rectangular  shape. 

Roemerus  robustus  new  species 
PI.  3,  figs.  20-31;  Fig.  8G;  Tab.  4 

Types.— ThQ  holotype,  a  left  propodus  and  dactylus  (UT  45704),  was  collected  by 
G.  L.  Dawe  from  the  Salenia  texana  marl  at  Shingle  Hills,  Travis  Co.,  Texas.  The 
paratype,  a  left  propodus  (orig.  GAB  26,  specimen  8),  was  collected  at  GAB  26.  The 
holotype  is  deposited  at  the  Univ.  of  Texas  (Austin)  as  UT  45704.  The  paratype  is 
deposited  in  the  San  Diego  Natural  History  Museum  (SDSNH  23662). 

Occurrence,  sample  size,  and  preservation.  — ¥'\\q  specimens  oi  Roemerus  robustus 
were  available  for  description;  1  from  Shingle  Hills  (UT  45704),  1  from  (GAB  26), 
and  3  from  (GAB  25). 

Etymology.  — The  name  is  taken  from  the  robust  nature  of  the  claws  of  this  taxon. 

Description.  — Claws  robust,  similar,  rectangular,  and  thick.  Palm  slightly  longer 


43 


Ui 


41 


1  n 

1U 

t 

• 

• 

E 

D 

» 

5mm  10  15 

Width 

Figure  10.     Graph  of  carapace  width  (mm)  vs.  length  (mm)  of  Palaeodromites  hodgesi  Bishop. 


than  high  and  fingers  short,  slightly  incurved,  movable  finger  overlapping  tip  of  fixed 
finger. 

Lower  edge  straight  except  for  slight  convexity  below  base  of  fixed  finger.  Proximal 
edge  perpendicular  to  lower  margin.  Distal  edge,  above  the  fixed  finger  vertical.  Upper 
margin  convex,  especially  proximally,  where  it  curves  down  onto  prominent  carpal 
articulator. 

Outer  face  transversely  convex  and  ornamented  by  broad  ridges  along  proximal 
and  distal  edges  formed  by  narrow  furrows  on  inside.  Surface  covered  by  sparse,  large, 
subdued  granules  except  for  lower  edge  which  has  numerous,  large,  prominent  granules. 

Inner  face  relatively  flat  and  ornamented  by  a  distal  ridge  and  a  bend  in  the 
exoskeleton  near  proximal  margin  forming  a  groove. 

Upper  margin  surmounted  by  an  oblique  ridge  which  runs  from  outer-distal  corner 
to  inner-proximal  corner  and  is  progressively  more  pronounced  proximally  until  it 
forms  a  noticeable  low  ridge  on  upper  edge  of  inner  face.  A  few  large  granules  may 
accentuate  ridge. 

At  least  1  tooth  can  be  seen  situated  on  fixed  finger  near  its  base. 

Comparison.— Roemerus  robust  us  differs  from  most  other  pagurids  by  having  a 
pronounced  rectangular  shape.  Only  Palaeopagurus  Van  Straelen  1925  and  Petrochirus 
Stimpson  1859  even  approach  this  shape. 

Infraorder  Brachyura  Latreille  1 803 

Section  Dromiacea  deHaan  1833 

Superfamily  Domiodea  deHaan  1833 

Family  Dynomenidae  Ortmann  1892 

Genus  Palaeodromites  A.  Milne-Edwards  1865 

Type  species.  — By  monoXyxiy',  Palaeodromites  octodentat us  A.  Milne-Edwards  1865, 
p.  345,  pi.  5;  Hauterivian  of  France. 

Diagnosis.— Carapace  broader  than  long,  rounded  pentagonal  or  hexagonal,  widest 
two-thirds  from  front,  gently  arched  transversely  and  longitudinally.  Front  square  to 
trapezoidal,  turned  strongly  downward;  orbits  large,  oval,  widely  spaced.  Anterolateral 


44 


Table  5.    Measurements  in  millimeters  of  the  carapace  of  Palaeodromites  naglei. 

Length  Width 

Specimen                                                                          (mm)  (mm) 

SDNHM  23644  (=25-1)                                                 8.5  10.7 

25^                                                                                    9.3+  11.5+ 

25-52                                                                              6.0?  8.6? 

25-53                                                                              7.8+  10.2 

25-54                                                                                  4.4  5.1 

25-57                                                                                  9.6+  11.5 

SDNHM  23645  (=21-21)                                              12.1  15.6 


borders  convex  with  tooth-like  spines  or  lobes,  posterolateral  borders  short,  straight, 
or  concave  without  tooth-like  lobes,  hind  margin  short,  straight,  or  concave.  Cervical 
furrow  clearly  defined,  sinuous  to  straight;  branchiocardiac  furrows  weakly  defined. 
(After  Wright  and  Collins  1972:49). 

Palaeodromites  naglei  new  species 
PI.  1,  figs.  12-17;  Figs.  8 A,  10;  Tab.  5 

Types.— The  holotype  SDSNH  23644  (orig.  GAB  25,  specimen  1)  and  paratype 
SDSNH  23645  (originally  GAB  21,  specimen  21)  of  Palaeodromites  naglei  are  both 
carapaces  and  are  deposited  in  the  San  Diego  Natural  History  Museum. 

Occurrence,  sample  size,  and  preservation.  —  Palaeodromites  naglei  has  been  found 
at  GAB  21  and  GAB  25. 

Etymology.  — In  honor  of  J.  Stuart  Nagle,  who  discovered  the  first  specimen  of  this 
taxon. 

Description.— Carapace  hexagonal,  1.2  times  wide  as  long,  very  convex  longitu- 
dinally, less  convex  transversely.  Carapace  furrows  poorly  developed,  a  faint  cervical 
furrow  and  3  branchial  furrows  present.  Rostrum  rounded,  nearly  vertical.  Orbits  poorly 
defined  in  dorsal  view  arching  upward.  Width  between  outer  angles  of  orbits  57% 
carapace  width.  Mesogastric  region  barely  set  off  from  rest  of  cephalic  arch.  Three  faint 
grooves  on  branchial  region  parallel  cervical  furrow.  Metagastric  and  protogastric  regions 
set  off  from  the  cardiac  and  branchial  regions  by  faint  muscle  attachment  scars.  Hind 
margin  paralleled  by  a  marginal  groove. 

Branchial  regions  split  into  4  fields  by  very  faint  grooves;  anteriormost  parallels 
the  cervical  furrow  to  a  point  behind  outer  angle  of  orbit  then  banks  backward  and 
outward  to  back  of  first  lateral  spine.  Second  groove  beginning  at  anterior  end  of  muscle 
attachment  field,  trends  outward  to  first  lateral  spine.  Third  furrow  extremely  faint, 
lying  just  inside  a  ridge  paralleling  posterolateral  margin. 

Anterolateral  margins  each  composed  of  4  forward  facing  broad  spines  which  get 
progressively  larger  posteriorly;  the  first  little  more  than  a  broadening  of  the  carapace 
edge  just  behind  outer  angle  of  orbit,  the  second  asymmetrical,  bent  sightly  forward, 
the  third  the  shape  of  an  equilateral  triangle,  and  the  fourth  broadly  rounded,  forming 
the  widest  part  of  carapace. 

Posterolateral  margin  serrated  by  a  series  of  3  or  4  spines  which  decrease  rapidly 
in  size  to  the  last  which  is  little  more  than  a  granule,  anteriormost  a  small  spine  on 
the  dorsal  shield  edge  about  the  size  of  second  anterolateral  spine. 

The  ventral  side  and  appendages  presently  unknown  because  none  of  these  parts 
definitely  attached  to  a  carapace. 

Comparison.  —Palaeodromites  naglei  is  much  smoother  and  less  ornamented  than 
other  species  of  this  genus.  Palaeodromites  naglei  is  additionally  distinguished  from  P. 
sinusosulcatus  Wright  and  Collins  1972  by  a  straight  cervical  furrow  and  lower  con- 
vexity, from  P.  incertiis  (Bell  1863)  by  the  lack  of  coarse  granulation,  and  from  P. 
transiens  Wright  and  Collins  1972  by  the  lack  of  posterolateral  ornamentation. 


45 


Table  6.    Carapace  measurements  in  millimeters  of  Diaulax  roddai. 


Width 
(mm) 

Length  from 

front  orbit  to 

hind  margin 

(mm) 

Orbital  wi 
(mm) 

dth 

Rostral 

Specimen 

Width 
(mm) 

Length 
(mm) 

SDNHM  23640  (=  25-2) 

25-58 

25-161 

10.6 
5.8 
6.6 

10.7 
6.1  + 

6.8 
4.0 
4.2 

1.6 

1.0+ 

Family  Diaulacidae  Wright  and  Collins  1972 
Genus  Diaulax  Bell  1863 

Type  species.  — Diaulax  carteriana  Bell  1863,  by  original  designation. 

Diagnosis.  — ^"Ihe  carapace  is  more  or  less  hexagonal,  widest  just  in  front  of  or 
just  behind  the  ends  of  the  cervical  furrow,  in  longitudinal  section  curved  more  or  less 
steeply  down  anteriorly  but  flat  posteriorly,  in  transverse  section  flat.  The  front  is 
generally  downturned,  pointed  or  squared;  it  may  be  sulcate  with  the  edges  turned  up 
into  prominent  lobes  or  nearly  flat.  The  antero-  and  posterolateral  margins  are  very 
sharp,  not  lobed  and  with  only  a  few  small,  sharp  spines  directed  forwards.  The  cervical 
and  branchiocardiac  furrows  are  weak  and  tend  to  be  straight  and  transverse.  The 
regions  are  poorly  defined.  The  surface  is  very  finely  granulate."  (Wright  and  Collins 
1972:56). 

Diaulax  roddai  new  species 
PI.  1,  figs.  1-2;  Fig.  8E;  Tab.  6 

Type.—T\iQ  holotype,  a  partial  carapace  (orig.  GAB  25,  specimen  2)  is  deposited 
in  the  San  Diego  Natural  History  Museum,  SDSNH  23640. 

Occurrence,  sample  size,  and  preservation.— Tht  holotype  is  1  of  3  specimens  of 
this  taxon  thus  far  collected.  It  is  an  almost  complete  carapace  missing  only  the  rostrum 
and  left  rear  corner  of  the  carapace.  The  exoskeleton  is  preserved  as  a  chalky  limestone 
and  does  not  show  surface  ornamentation  very  well.  One  specimen  (GAB  25,  specimen 
58)  has  a  partly  preserved  rostrum  and  the  third  specimen  (GAB  25,  specimen  161)  is 
a  poorly  preserved,  crushed  carapace. 

Etymology.  — ¥ov  Peter  U.  Rodda,  Curator  of  Geology,  California  Academy  of 
Sciences,  whose  encouragement  led  to  the  completion  of  this  study. 

Description.  — C2LY?LX)2LCt  kite-shaped  with  truncated  anterior  and  posterior  slightly 
longer  than  wide,  widest  about  '/3  distance  from  front.  Large  specimens  flat  with  raised 
anterolateral  margins  turned  slightly  under  the  dorsal  shield  on  pterygostomial  regions; 
smaller  specimens  with  higher  relief. 

Cervical  groove  indistinct  except  for  notch  where  it  meets  edge  of  the  dorsal  shield 
near  widest  part  of  carapace.  Epimeral  muscle  scars  present  but  not  deeply  incised. 

Regions  poorly  differentiated;  rostrum  fairly  broad,  long.  Cephalic  arch  dominated 
by  spines  at  outer  edges  of  orbits  and  raised  anterolateral  margins  running  from  orbital 
spines  to  cervical  notch.  Gastric  areas  slightly  higher  than  rest  of  cephalic  arch  with 
slight  protogastric  bosses.  Distance  between  outer  edges  of  orbits  -A  the  carapace  width. 
Scapular  arch  flat,  undiflerentiated  except  for  epimeral  muscle  scars  and  slight  bosses 
distal  to  epimeral  peninsulas,  forming  ridges  that  continue  almost  to  hind  margin. 

Anterolateral  margins  dominated  by  large  upward  and  forward  pointing  spines  at 
outer  edge  of  upward  turned  orbits  that  have  a  single  fissure  on  lower  edge,  raised 
margin  concave  to  cervical  notch.  A  large  upward,  forward  pointing  spine  and  a  second, 
smaller  spine  lie  on  margin  behind  cervical  notch  (second  spine  about  same  distance 
behind  notch  as  orbital  spine  is  ahead  of  it).  Dorsal  shield  margin  convex  to  first 
scapular  spine  and  runs  almost  in  a  straight  line  to  concave  V-shaped  posterior  margin. 
Hind  margin  bordered  by  furrow. 


46 


Comparison.— Diaiilax  roddai  is  distinguished  from  D.  oweni  (Bell  1850)  and  D. 
carteriana  Bell  1863  by  being  relatively  longer  and  more  flat,  especially  longitudinally. 
Diaulax  roddai  most  resembles  D.  feliceps  Wright  and  Collins  1972  but  is  relatively 
longer,  has  its  maximum  width  further  forward,  and  has  a  longer  rostrum. 

Remarks.  SrmW  specimens  of  this  taxon  appear  to  have  a  better  differentiated 
carapace  than  larger  specimens. 

Superfamily  Dorippoidea  de  Haan  1841 

Family  Dorippidae  de  Haan  1841 

Subfamily  Dorippinae  de  Haan  1841 

Hillius  new  genus 

Type  species.— Hillius  youngi  new  species 

Diagnosis.— CdiYapsLCQ  pentagonal,  fairly  flat,  slightly  wider  than  long,  widest  half 
the  distance  from  front.  Grooves  broad  and  indistinct.  Rostrum  broad.  Orbital  width 
50%  carapace  width,  orbits  small,  upturned,  notched  near  inner  comer,  with  raised 
rim.  Epibranchial  areas  wide,  giving  specimens  a  wing-like  or  ray-like  appearance. 

Etymology.  — ¥or  Robert  Thomas  Hill,  pioneer  Texas  geologist,  stratigrapher,  and 
paleontologist. 

Comparison.— This  taxon  has  a  striking  resemblance  in  carapace  shape  to  Dorippe 
Weber  1795,  Goniochele  Bell  1858,  and  Orthopsis  Carter  1872.  It  differs  from  Dorippe 
by  having  a  wider  front  with  convex  anterolateral  margins.  Hillius  differs  from  Gon- 
iochele by  having  the  widest  part  of  the  carapace  relatively  more  forward  and  formed 
by  the  epibranchial  lobes.  Hillius  is  most  similar  to  Orthopsis  but  differs  from  it  by  its 
having  less  relief,  lack  of  anterolateral  spination,  and  single  orbital  lobe. 

Hillius  voungi  new  species 
PI.  1,  figs.  8-11;  Fig.  8C 

Type.— The  Holotype,  a  carapace  steinkern  (orig.  GAB  25,  specimen  3),  is  depos- 
ited in  the  San  Diego  Natural  History  Museum  (SDSNH  23643). 

Occurrence,  sample  size,  and  preservation.— The  Holotype  is  the  only  specimen  of 
this  taxon's  carapace  that  is  nearly  complete.  The  specimen  was  collected  at  GAB  25. 
As  with  any  description  from  decortiated  specimens  only  major  ornamentation  features 
are  likely  to  be  decipherable,  and  a  specimen  with  exoskeleton  will  be  needed  to 
completely  define  this  taxon. 

Etymology.  — In  honor  of  Keith  Young,  Texas  Cretaceous  stratigrapher  and  pale- 
ontologist. 

Description.— Carapace  pentagonal,  probably  slightly  wider  than  long  (partial  length 
11.5  mm),  widest  about  half  the  distance  from  the  front.  Carapace  slightly  convex 
longitudinally;  cephalic  arch  moderately  convex  transversely,  scapular  arch  fairly  flat 
transversely,  except  at  the  edges. 

Cervical  furrow  broad  and  indistinct  except  where  it  crosses  dorsal  shield  margin 
in  a  pronounced  notch  continuing  on  subhepatic  region  as  a  well-defined  groove  to 
base  of  orbit.  An  indistinct  groove  on  subhepatic  region  lies  above  and  is  parallel  to 
cervical  furrow.  Faint  but  distinct  mesogastric  grooves  present.  Hepatic  grooves  broad 
and  poorly  defined.  The  most  prominent  grooves  on  carapace  separate  gastrocardiac 
region  from  branchial  regions.  Broad  grooves  separate  urogastric,  cardiac,  and  intestinal 
regions,  two  faint  grooves  separate  branchial  regions  into  3  parts,  the  anterior  runs 
over  carapace  edge  just  behind  lateral  spine  then  swings  rapidly  forward  toward  cervical 
furrow.  Rostrum  about  'A  carapace  width.  Orbits  small,  upturned,  with  notch  near  inner 
corner,  with  raised  rim.  Distance  between  outer  edges  of  orbits  about  50%  carapace 
width.  Cephalic  arch  differentiated  into  small  mesogastric  area;  large  protogastric  area 
surmounted  by  large,  low  circular  bosses;  and  an  upturned  hepatic  region  with  at  least 
2  small  marginal  spines  immediately  ahead  of  cervical  notch.  Scapular  arch  well  dif- 
ferentiated into  a  segmented  medial  ridge  (consisting  of  urogastric,  cardiac,  and  intes- 
tinal regions)  and  branchial  regions  (divided  into  epibranchial,  mesobranchial,  and 


47 


metabranchial  regions).  Urogastric  regions  have  a  gentle  forward  slope  and  a  steep 
posterior  slope  giving  rise  to  2  transverse  crescentic  ridges  with  small  medial  spine 
where  they  meet.  Cardiac  region  diamond-shaped  with  2  tubercles  symmetrically  placed 
across  medial  axis.  Intestinal  region  poorly  defined  and  partly  missing,  a  single  medial 
intestinal  tubercle  near  hind  margin.  Short  longitudinal  ridges  lie  in  each  gastrobranchial 
groove,  perhaps  the  ridges  formed  within  the  epimeral  muscle  scars.  Epibranchial  region 
small  but  forming  the  prominent  lateral  wing  of  this  taxon.  A  small  marginal  epigastric 
spine  lies  in  posterior  part  of  cervical  notch.  Two  small  spines  are  situated  near  lateral 
margin  of  epibranchial  wing;  the  anterior  smaller  and  the  posterior  larger.  Mesobran- 
chial  and  metabranchial  regions  with  fairly  continuous  broad  longitudinal  ridges  from 
near  the  epibranchial  groove  to  hind  margin.  Posterior  of  mesobranchial  region  with 
small  boss  on  this  ridge,  surmounted  by  several  granules.  A  small  marginal  spine 
situated  at  anterior  of  metabranchial  region.  Hind  margin  missing. 

Comparison.— Hillius  youngi  differs  from  Orthopsis  bonneyi  Carter  1872  by  its 
lack  of  anterolateral  spines,  single  instead  of  double  orbital  lobe,  more  subdued  carapace 
relief,  and  different  carapace  outline  due  to  the  widest  point  of  the  carapace  being 
farther  forward. 

Remarks.  — T\\Q  line  drawing  of  the  carapace  of  this  taxon  is  based  on  a  single 
steinkem.  When  further  material  becomes  available  the  description  should  be  amended 
to  include  surface  ornamentation. 

Family  Torynommidae  Glaessner  1980 
Genus  Dioratiopus  Woods  1953 

Dioratiopus  Woods  1953,  p.  52;  Wright  and  Collins  1972,  p.  33,  34,  42. 
Doratiopus  Woods,  Glaessner  1969,  p.  492  (erroneous  spelling). 
Glaessneria  Wright  and  Collins  1972  {non  Takeda  and  Miyake  1964),  p.  34  ff. 
Glaessnerella  Wright  and  Collins  1975,  p.  441. 

Type  species.— Homolopsis  spinosa  (Van  Straelen  1936),  p.  33;  Albian  of  Valcourt 
France. 

Diagnosis.  — ''C2iV2ipa.cQ  more  or  less  pentagonal  with  parallel  sides,  strongly  pro- 
jected frontal  area,  long  rostrum  with  lateral  spines  and  large  shallow  indistinct  orbits 
complete  above;  the  sides  are  vertical  and  there  are  traces  of  a  lateral  margin  anteriorly, 
but  it  is  normally  not  sharp  or  fully  developed;  the  cervical  and  branchiocardiac  furrows 
are  strongly  marked;  a  short  oblique  furrow  runs  forward  from  the  outer  end  of  the 
branchiocardiac  and  may  extend  as  far  as  the  cervical,  delimiting  an  epibranchial  lobe; 
there  is  a  strong  postorbital  spine  at  or  just  behind  the  anterolateral  angle."  (Wright 
and  Collins  1972:34). 

Dioratiopus  scotti  new  species 
PI.  1,  figs.  6-7;  Fig.  8D 

rv'pc  — The  holotype,  a  partial  carapace  steinkern  (orig.  GAB  27,  specimen  1), 
collected  at  Jacob's  Well,  Hays  Co.,  Texas,  is  deposited  in  the  San  Diego  Natural 
History  Museum  (SDSNH  23642). 

Occurrence,  sample  size,  and  preservation.— The  holotype  is  the  only  specimen  of 
this  taxon  so  far  collected.  It  is  decortiated  carapace  steinkern  preserving  most  of  the 
dorsal  shield. 

Etymology.  — ¥ov  Alan  J.  Scott,  Texas  Cretaceous  and  Holocene  paleontologist  and 
stratigrapher. 

Description.— CsiVdiTp^ce  rectangular,  longer  (partial  length  7.9  than  wide  (partial 
width  6.8  mm).  Cephalic  arch  moderately  convex  transversely;  scapular  arch  fairly  flat 
transversely;  carapace  relatively  level  longitudinally. 

Cervical  furrow  narrow,  deep,  and  prominent;  dorsally  parallel  to  anterior  dorsal 
shield  edge,  turning  inward  and  backward,  then  backward  for  a  short  distance  cutting 
across  medial  ridge.  Branchiocardiac  furrow  crosses  carapace  just  behind  cervical  fur- 


48 

row.  Furrows  parallel  to  point  where  cervical  furrow  bends  inward  to  cross  medial 
ridge,  from  where  branchiocardiac  furrow  continues  backward  joining  epimeral  muscle 
scars  and  spliting  at  point  near  posterior  epimeral  muscle  scar,  1  part  swinging  inward 
crossing  medial  ridge  as  a  broad,  poorly  defined  groove  and  other  continuing  as  epimeral 
muscle  scar.  Outer  arm  of  epimeral  muscle  scar  loops  back  inward  forming  small,  flat, 
oval  area.  Well-defined  branchial  furrow  splits  off'  epimeral  muscle  scar  near  where  it 
begins  the  loop,  proceeds  outward  and  forward  to  dorsal  shield  edge. 

Rostrum  probably  triangular,  occupying  40%  carapace  width.  Orbits  apparently 
small;  with  sharply  upturned  rims,  occupying  55%  carapace  width.  Mesogastric  area 
separated  from  broad,  swollen  protogastric  regions  by  shallow,  distinct  groove,  narrow 
anteriorly  but  rapidly  widening  at  posterior  half  Protogastric  region  with  small  circular 
boss  situated  at  center.  A  row  of  12  small  granules  begins  on  raised  orbital  rim  and 
forms  an  incomplete  circle  to  mesogastric  grooves  around  each  protogastric  boss.  The 
region  lying  between  2  transverse  grooves,  crescent-shaped,  concave  side  anterior,  with 
short  transverse  base  posterior  to  middle,  giving  rise  to  2  transverse  ridges.  Medial 
area  behind,  where  the  second  groove  crosses,  a  raised  region  separated  by  a  shallow 
medial  groove  at  its  summit  into  2  longitudinal  ridges.  Rear  portion  of  dorsal  shield 
missing.  Branchial  regions  divided  into  2  parts  by  branchial  cardiac  furrows  running 
forward  and  outward;  anterior  region  has  single  longitudinally  expanded  granule  near 
outer  edge,  posterior  branchial  region  has  2  granules  on  outer  margin  directly  behind 
1  on  the  anterior  branchial  lobe.  The  anterior  of  these  2  is  longitudinally  expanded 
and  large;  the  posterior  round  and  small. 

Margins  of  dorsal  shield  poorly  preserved.  The  photograph,  taken  before  prepa- 
ration was  finished,  gives  the  impression  of  a  straight  lateral  margin  (left  side)  nearly 
to  a  point  on  line  with  the  rear  of  the  orbits.  The  right  side  appears  to  be  gently  convex. 

Comparison.— Dioratiopus scotti  is  most  similar  to  D.  spinosa  (Van  Straelen  1 936) 
in  the  size  and  shape  of  carapace  regions.  It  differs  from  D.  spinosa  by  having  a  wider 
urogastric  region  with  2  transverse  ridges,  a  longitudinally  bilobate  cardiac  region, 
apparently  no  furrow  delimiting  an  epigastric  region,  distal  spines  on  the  branchial 
regions,  and  probably  a  smoother  carapace.  The  differences  in  carapace  size  and  shape, 
size,  shape,  and  arrangement  of  carapace  regions,  and  ornamentation  is  even  greater 
between  D.  scotti  and  other  congeners. 

Remarks.— The  placement  of  this  single  specimen  into  generic  level  taxon  is  strongly 
hampered  by  its  mode  of  preservation  as  a  steinkem  and  by  the  obscure  nature  of  the 
lateral  margins  of  the  carapace  fragment.  The  discontinuous  nature  of  the  lateral  margins 
may  point  to  the  lack  of  dorsal  pleural  sutures,  linea  homolica,  in  which  case  this 
specimen  does  not  belong  in  Homolopsis.  This  lack  of  a  straight  break  and  the  similarity 
of  carapace  morphology  to  Dioratiopus  suggests  a  close  alliance  with  this  genus,  and 
the  specimen  is  therefore  assigned  to  Dioratiopus  until  more  complete  material  becomes 
available. 

Section  Oxystomata  H.  Milne-Edwards  1834 
Superfamily  Calappoidea  de  Haan  1833 

Family  Calappidae  de  Haan  1833 
Subfamily  Necrocarcininae  Forster  1968 
Genus  Pseudonecrocarcinus  Forster  1968 

Type  species.  — By  monotypy;  Necrocarcinus  quadriscissus  Noetling  1881,  p.  368, 
pi.  20,  fig.  4);  Maastrichtian,  Limbourg,  Holland. 

Diagnosis.— CdiVdiPdiQe  wider  than  long,  frontal-orbital  margin  about  Vi  carapace 
width.  Medial  regions  poorly  differentiated.  Mesogastric  and  protogastric  regions  com- 
bined into  wing-like  swellings.  Inner  side  of  epibranchial  region  with  ridge,  outer  side 
with  tubercle  groups;  metabranchial  region  with  weak  longitudinal  ridges.  Sulcus  of 
rostrum  with  2  or  4  pits.  Deeply  incised  angular  grooves  form  the  lateral  boundaries 
of  the  urogastric  region. 


49 


Pseudonecrocarcinus  stenzeli  new  species 
PI.  1,  figs.  3-5;  Fig.  8B 

Types.— The  Holotypc,  an  impression  of  a  carapace,  (orig.  GAB  25,  specimen  8) 
is  deposited  in  the  San  Diego  Natural  History  Museum  (SDSNH  23641). 

Occurrence,  sample  size,  and  preservation.— T\\q  Holotype,  an  impression,  and  5 
partial  carapaces  were  collected  at  GAB  25. 

Zinv;;t)/o^v.— Named  in  honor  of  Dr.  Henryk  B.  Stenzel,  a  leader  in  the  study  of 
Cretaceous  and  Tertiary  Texas  decapods. 

Description.— C^iVSipsiCQ  slightly  wider  than  long;  widest  about  V3  distance  from 
front.  Length  from  broken  tip  of  rostrum  to  hind  margin  1 1 .8  mm  and  width  (computed 
as  2  times  the  width  of  half  the  crab)  12.5  mm. 

Carapace  fairly  flat  transversely  and  longitudinally,  anterolateral  margins  lie  lower 
than  gastric  arch.  Cervical  furrow  broad,  faint  distally  but  narrower  and  more  distinct 
on  central  part  of  carapace  near  where  it  ends  in  a  pair  of  gastric  pits;  branching  at 
distal  end,  1  branch  continues  laterally  to  dorsal  shield  edge  and  other  swings  anteriorly 
forming  a  broad,  shallow  depression  which  borders  anterior  side  of  hepatic  region, 
arched  orbital  region,  and  arched  protogastric  region.  Epimeral  muscle  scars  deep, 
forming  the  most  noticeable  grooves.  Inner  side  of  V's  thus  formed  continue  forward 
as  shallow,  broad  depressions  connecting  with  cervical  furrow;  very  faint  grooves  sep- 
arate mesogastric  and  protogastric  regions. 

Carapace  regions  poorly  differentiated  by  shallow  grooves;  separation  into  bosses 
or  areoles  similarly  subdued.  Rostrum  broad,  nearly  25%  carapace  width.  Orbits  large, 
arched  upward,  2  fissures  on  rear  margins,  width  between  outer  edges  of  orbits  about 
50%  carapace  width.  Mesogastric  and  protogastric  regions  barely  discernable.  Two  pairs 
of  pits  lie  behind  rostrum  in  mesogastric-protogastric  grooves  on  line  with  back  of 
supraorbital  fissures;  outermost  pair  larger,  situated  slightly  more  forward  than  smaller, 
better-defined  pair.  Protogastric  regions  with  broad,  poorly  defined  bosses.  Hepatic 
region  with  raised  boss  directly  above  anterolateral  spine.  Cardiac  regions  sharply  set 
off'  from  branchial  regions  by  epimeral  muscle  scar  but  barely  separated  anteriorly 
except  for  shallow  cervical  groove,  continuing  posteriorly  with  intestinal  regions  to 
hind  margin.  Branchial  regions  broadly  arched  near  epimeral  muscle  scars,  with  a  raised 
ridge  along  carapace  edge  from  just  behind  cervical  furrow  to  a  point  on  line  with 
middle  part  of  epimeral  muscle  scar. 

Anterolateral  margin  slightly  concave  for  a  short  distance  as  it  leaves  protruding 
outer  edge  of  orbit,  dropping  in  elevation  until  it  begins  to  become  convex,  arching 
slightly  upward  then  downward  just  in  front  of  hepatic  spines.  Indentations  in  carapace 
margin  in  front  of  hepatic  spine  and  behind  it  where  cervical  furrow  meets  carapace 
edge.  Posterolateral  margin  slightly  convex  from  cervical  furrow  to  hind  margin.  Hind 
margin  concave,  bordered  by  shallow  but  distinct  groove. 

Carapace  ornamentation  consists  of  the  few  broad  bosses,  anterodistal  branchial 
ridges,  deep  epimeral  muscle  scar,  shallow  grooves,  and  a  very  fine  granulation  (slightly 
coarser  on  medial  part)  over  carapace. 

Comparison.  — ThQ  genus  Pseudonecrocarcinus  contains  P.  quadriscissus  (Noetling 
1881).  P.  biscissus  Wright  and  Collins  1972,  and  P.  stenzeli  Bishop  new  species.  All  3 
species  have  post-rostral  pits  in  common.  This  character  separates  this  genus  from  the 
other  genera  of  the  Necrocarcininae.  In  Pseudonecrocarcinus  stenzeli  and  P.  quadris- 
cissus the  pits  are  elongated  into  slits.  The  carapace  of  P.  stenzeli  is  much  smoother 
than  that  of  P.  quadriscissus. 

Remarks.— The  presence  of  pits  at  the  base  of  the  rostrum  unite  P.  quadriscissus, 
P.  biscissus,  and  P.  stenzeli  into  a  distinct  group.  I  believe  it  is  best  to  maintain  their 
distinction  as  a  separate  genus  until  such  time  as  we  have  more  data  with  which  to 
judge  the  phylogenetic  affinities  of  the  necrocarcinids. 


50 


Figure  1 1.     Tubercle  placement  on  claws  of  Prehepat us  hodgesi  Bishop. 


Superfamily  Calappoidea  de  Haan  1833 

Family  Calappidae  de  Haan  1833 

Subfamily  Matutinae  McLeay  1838 

Genus  Prehepatus  Rathbun  1935 

Type  species.  — ^y  original  designation:  Prehepatus  cretaceous  Rathbun  1935,  p. 
47,  PI.  11,  figs.  29-30. 

Diagnosis.  — ChtX^Q  small,  increasing  in  height  to  distal  end  of  palm;  fixed  finger 
short,  movable  finger  stout;  transversely  flat  to  concave  on  inner  face,  convex  on  outer 
face  with  upper  margin  broadly  rounded  to  flat  and  forming  an  oblique  keel  which 
overhangs  the  inner  face;  surface  ornamented  by  strong  tubercles. 

Prehepatus  hodgesi  new  species 
PI.  3,  figs.  1-19;  Figs.  8F,  11,  12;  Tab.  7 

Types.— ThQ  holotype  o^ Prehepatus  hodgesi  is  a  partial  right  propodus  (orig.  GAB 
25,  specimen  14)  deposited  in  the  San  Diego  Natural  History  Museum  (SDSNH  23655). 
Six  paratypes  (SDSNH  23656  to  23661)  are  also  deposited  in  the  San  Diego  Natural 
History  Museum. 

Occurrence,  sample  size,  and  preservation.— Twenty  claws  of  this  taxon  are  known 
from  localities  GAB  21,  GAB  25,  and  GAB  26. 

Etymo/ogy.  —  Named  in  honor  of  Floyd  Hodges  whose  fortuitous  spilling  of  coffee 
on  his  lap  caused  the  discovery  of  the  Boeme-Sisterdale  locality  (GAB  25)  which  yielded 
so  many  fine  specimens  of  this  taxon. 

Description.— Carpus  with  large  tubercle  on  rear  distal  corner,  smaller  tubercle  on 
front  distal  corner,  1  on  center  of  dorsal  face  and  1  midway  on  rear  dorsal  margin. 
Margin  of  proximal,  dorsal  side  with  row  of  5  granules. 

Right  propodus  triangular,  about  twice  as  long  as  high  (Table  7),  palms  slightly 
longer  than  high,  highest  at  distal  end  of  palm.  Level  upper  margin  about  half  the 
length  of  convex  lower  margin,  flat,  horizontally  overhanging  inner  face  at  proximal 
edge.  Carinate  proximal  part  of  crest  gives  way  to  4  broad,  thin  blade-like  spines  which 
become  progressively  more  vertical  distally  to  base  of  dactylus.  Lower  margin  trans- 
versely convex.  Outer  face  convex  longitudinally,  very  convex  transversely.  Inner  face 
fairly  flat.  Two  broad,  shallow  concavities  present,  1  below  upper  margin  and  1  at 
proximal  end  just  above  lower  margin.  Propodus  ornamented  by  numerous  tubercles 
(situated  in  3  lines).  Upper  sinuous  row  of  6  tubercles  (9,  7,  8,  10,  11)  beginning  at 
lower  carpal  articulator  follows  the  carpal  margin,  swinging  forward  to  follow  the  upper 
margin  to  distal  edge  above  dactyl  articulator.  Second  row  begins  at  same  point  as  first 
(near  lower  carpal  articulator),  proceeds  in  sinuous  path  to  base  of  fixed  finger  through 
4  large  tubercles  (counting  the  first  one  again)  (9,  3,  2,  1)  and  1  minor  tubercle  (18) 
between  distal  2  tubercles  (1  and  2).  The  distal  3  tubercles  of  this  row  are  the  most 


51 


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Figure  12.  Graph  of  palm  height  (mm)  vs.  palm  length  (mm)  for  Prehepatus  hodgesi  Bishop.  Right  claws 
plotted  as  dots  (read  length  on  top  scale  increasing  left  to  right)  and  left  claws  as  stars  (read  length  on  lower 
scale  increasing  from  right  to  left). 


consistent  of  the  tubercles  in  presence  and  relative  size.  A  third  row  of  tubercles  (5, 
12,  13,  14)  runs  in  a  straight  line  from  a  medium  tubercle  (5)  above  midpoint  of  lower 
margin  almost  to  lower  edge  of  the  fixed  finger.  In  addition  to  these  "rows"  a  small 
node  is  sometimes  found  in  angle  formed  by  the  upper  rows  (16),  a  medium  tubercle 
(6)  below  middle  of  upper  row,  a  small  one  (15)  below  and  behind  tubercle  2,  a  small 
one  (17)  below  tubercle  5,  and  a  small  one  (19)  above  and  between  tubercles  10  and 
1 1 .  A  spot  above  tubercle  8  sometimes  is  roughened  or  tuberculate.  Inner  face  orna- 
mented by  a  single  tubercle  midway  along  carpal  margin.  Fixed  finger  turned  downward 
and  slightly  inward,  with  faint  furrow  on  lower  edge  of  outer  face.  At  least  2  "teeth" 
are  present  on  prehensile  edge;  proximal  tooth's  surface  divided  into  3  lobes.  Movable 
finger  strongly  curved,  with  narrow  angular  ridge  on  outer  side  from  the  articulator  to 
midway  along  dactyl,  where  it  broadens  and  rounds  out  to  tip.  Fingers  finely  granulate 
and  weathered  differently  than  rest  of  claw,  appearing  more  stable  or  resistant. 

Left  propodus  same  shape  as  right,  probably  also  same  size  (Table  7),  outer  surface 
with  fewer  tubercles  than  right.  Tubercle  size  highly  variable,  positions  more  variable 
than  those  on  right  claw.  A  row  of  4  tubercles  (3,  4,  5,  10)  runs  in  an  arc  along  carpal 
margin  from  lower  articular  onto  palm.  Most  other  tubercles  lie  on  2  lines  that  run 
between  major  tubercle  (1)  at  base  of  fixed  finger  to  a  large  spine  (2)  distal  of  lower 
carpal  articulator,  upper  row  convex  upwards,  consisting  of  4  tubercles  (1,  8,  6,  and 
2),  lower  row  convex  downward  and  consisting  of  4  tubercles  (1,7,9  and  2).  Two  small 
tubercles  (1 1  and  12)  may  be  situated  on  lower  edge  of  palm  below  tubercles  1  and  7. 
Three  small  tubercles  (13,  14,  and  15)  may  be  present  on  flattened  crest. 

Comparison.  — Prehepatus  hodgesi  has  a  consistent  (though  variable)  pattern  of 
tuberculation  different  than  P.  cretaceus  Rathbun  1935;  P.  pawpawensis  Rathbun  1935; 
and  P.  dilksi  Roberts  1962.  Prehepatus  hodgesi  is  not  as  rectangular  as  P.  cretaceus 
Rathbun  1935,  lacks  the  great  number  of  spines  on  the  upper  margin,  does  not  possess 
the  large  tubercle  near  the  upper  dactyl  articulator,  and  does  not  have  a  tuberculate 
dactylus. 

Prehepatus  hodgesi  does  not  have  the  numerous  fine  granules  of  P.  pawpawensis. 


53 


Table  8.    Measurements  in  millimeters  of  claws  of  Torvnomma?  densus. 


Specimen 

Palm  length 
(mm) 

Height 

(mm) 

Claw  length 
(mm) 

Claw  thickness 
(mm) 

SDNHM  23664  (=25-5) 
SDNHM  23663  (=25-10) 

7.5 
7.2 

7.0 
5.8 

12.3 

3.8 

Prehepatiis  hodgesi  differs  from  P.  dilksi  Roberts  1962  in  its  ornamentation,  lack  of 
granulate  dactylus,  and  lack  of  the  vertical  furrow  and  rim  on  the  propodus  along  the 
distal  end  of  the  outer  surface. 

Remarks.  — Tho:  taxon  shows  a  surprisingly  consistent  pattern  of  ornamentation  in 
the  arrangement  of  tubercles  and  an  equally  surprising  amount  of  variation  in  size  (or 
presence)  of  the  tubercles.  If  only  a  few  specimens  had  been  found,  it  is  quite  probable 
I  would  have  been  tempted  to  place  them  in  different  species  level  taxa. 

I  feel  fairly  confident  that  there  are  at  least  2  recognizable  instars.  The  smaller 
specimens  are  relatively  thinner  and  nearly  smooth.  A  second  instar  shows  tremendous 
variation  in  tubercle  size.  A  third  possible  instar  may  be  present  and  contains  the 
largest,  most-ornamented  specimens. 

The  data  on  cheliped  ornamentation  are  included  as  I  believe  there  may  be  a 
behavioral  analogy  with  the  claws  of  fiddler  crabs  (Crane  1975)  and  the  data  may  be 
useful  to  subsequent  decapod  workers. 

Section  Brachyrhyncha  Borradaile  1907 
Superfamily  Dorippoidea  de  Haan  1841 

Family  Dorippidae  de  Haan  1841 

Genus  Torvnomma  V^oods  1953 

Type  species.  — Torvnomma  qiiadrata  by  original  designation. 
Z)/a^A205Z5.— "Carapace  subquadrate,  widest  anteriorly,  orbital  grooves  large,  ros- 
trum narrow,  oviduct  opening  on  coxa  of  3rd  periopods."  (Glaessner  1969:493.) 


Torvnomma?  densus  new  species 
PI.  3,  f^g.  32-40;  Fig.  8J;  Tab.  8 

Types.  — l^YiQ  Holotype,  a  right  propodus  (GAB  25,  specimen  5)  and  paratype  (GAB 
25,  specimen  10)  are  deposited  in  the  San  Diego  Natural  History  Museum  (SDSNH 
23664  and  SDSNH  23663  respectively). 

Occurrence,  sample  size,  and  preservation.  — Tory nomma?  densus  is  represented 
by  2  right  propodi  from  GAB  25.  The  holotype  is  nearly  complete  except  for  some 
dissolution  of  exoskeleton.  The  paratype  is  partly  crushed  on  the  inner  side  and  has 
the  fixed  finger  broken  off  Neither  specimen  preserves  the  dactylus.  The  band  of  dense 
exoskeleton  along  the  distal  margin  and  on  the  fixed  finger  is  excellently  preserved  on 
each  specimen. 

Etymology.  — ¥ov  dense  exoskeleton  along  distal  edge  of  claw. 

Description.  — Claw  nearly  twice  as  long  as  high,  palm  slightly  longer  than  high. 
Lower  margin  slightly  convex,  tightly  rounded  at  lower  proximal  corner  slanting  forward 
along  carpal  articulation,  then  broadly  rounded  to  convex  upper  margin.  Distal  edge 
nearly  vertical  concave  to  base  of  fixed  finger.  Fixed  finger  long,  narrow,  curving  inward; 
decreasing  in  size  distally  by  2  steps  before  it  reaches  pointed  tip.  Outer  face  of  claw 
convex,  inner  face  slightly  convex.  A  band  of  dense  exoskeleton  present  along  distal 
edge  of  claw  and  on  fixed  finger. 

Shallow  depression  on  outer  face  on  dense  band  at  base  of  upper  margin  of  fixed 
finger,  giving  way  to  shallow  groove  that  parallels  lower  margin  of  finger  to  its  tip.  Field 
of  small  granules  near  the  upper  part  of  dense  band  and  below  fixed  finger  groove.  A 
smooth  lineation  lies  on  the  lower  edge  of  the  claw  giving  rise  to  an  apparent  groove. 
Knob-shaped  upper  carpal  articulator  granulate. 


54 


Inner  face  fairly  flat,  shallow  depression  below  upper  margin  forming  slight  over- 
hanging ridge,  coarsely  granulate  on  dense  band.  A  shallow,  smooth  groove  running 
along  middle  of  finger  joins  second  step-down.  Dense  band  abundantly  granulate  below 
groove  and  sparsely  granulate  above  it.  A  field  of  numerous  granules  lies  on  the  dense 
band  even  with  upper  margin  of  fixed  finger.  Occlusional  surface  formed  by  a  broad 
ridge  with  2  "teeth"  formed  where  the  step-downs  in  size  occur. 

Comparison.  — Toryfwmma?  densus  has  a  shape  similar  to  Torynomma  quadrata 
Woods  1953  but  differs  by  a  more  convex  lower  margin,  a  stouter,  stepped-down  fixed 
finger,  and  the  band  of  very  dense  exoskeleton  on  the  fixed  finger  and  along  the  distal 
edge  of  the  palm. 

Remarks.— ThQ  assignment  of  this  claw  to  a  taxon  was  most  difficult  because  most 
taxa  that  have  a  preserved  carapace  do  not  have  an  adequate  description  of  the  chelae. 
The  dense  exoskeleton  on  the  finger  and  distal  edge  of  the  claw  suggests  that  this  may 
be  a  xanthid-like  crab.  It  was  not  assigned  to  the  Xanthidae  because  on  xanthids  usually 
only  the  fingers  have  dense  exoskeleton,  the  fixed  finger  in  xanthids  normally  has 
"teeth,"  and  this  taxon  would  have  extended  the  range  of  xanthids  from  Upper  Cre- 
taceous into  the  Lower  Cretaceous. 

It  was  deemed  better  in  this  case  to  name  a  new  taxon  (knowing  it  will  most  likely 
be  synonomized  when  the  claw  is  matched  to  a  carapace)  than  to  have  yet  another 
nameless  taxon  to  refer  to  in  the  literature. 

Acknowledgments 

Many  persons  were  directly  or  indirectly  involved  in  this  study.  Those  who  aided 
by  helping  collect  specimens  were  Nelda  Bishop,  Susan  Deutsch  Conger,  Arthur  Cleaves, 
Lyman  Dawe,  Tom  Grimshaw,  Floyd  Hodges,  Don  Lentzen,  John  Newcomb,  Mary 
Beth  Bowers  Schwartz,  and  Keith  Young.  Discussions  with  Keith  Young,  H.  B.  Roberts, 
Alan  Scott,  and  particularly  Peter  Rodda  resulted  in  a  stimulus  to  complete  this  study. 
The  manuscript  was  reviewed  by  Rodney  Feldmann  and  Karl  Waage.  Jacque  Causey 
typed  the  final  manuscript  at  Georgia  Southern  College.  A  Faculty  Research  Grant 
from  the  Faculty  Research  Committee,  Georgia  Southern  College,  expedited  the  com- 
pletion of  this  work,  and  an  NSF  grant  (DEB  80115  70)  provided  time  for  major  revision 
and  publication  costs. 

Literature  cited 


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Bell,  T.  1863.  A  monograph  of  the  fossil  mala- 
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Crane,  J.  1975.  Fiddlercrabsofthe  world.  Prince- 
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Hendricks,  L.,  and  W.  F.  Wilson,  1967.  Co- 
manchean  (Lower  Cretaceous)  stratigraphy  and 
paleontology  of  Texas.  Permian  Basin  Section 
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Noetling,  F.  1881.  Ueber  einige  Brachyuren  aus 
dem  Senon  von  Maestricht  und  dem  Tertiar 
Narddeutschlands.  Deutsch.  Geol.  Gesell. 
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Rathbun,  M.  J.  1935.  Fossil  Crustacea  of  the 
Atlantic  and  Gulf  Coastal  Plain.  Geol.  Soc.  Am. 
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Roberts,  H.  B.  1962.  Crustacea.  Pages  163-191, 
in  H.  G.  Richards  et  al.  (editors).  The  Creta- 
ceous fossils  of  New  Jersey:  New  Jersey  Bur. 
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ton, N.J.,  USA. 

Stenzel,  H.  B.  1945.  Decapod  Crustacea  from  the 
Cretaceous  of  Texas.  Texas  Univ.,  Publ.  No. 
4401:401^76. 

Stimpson,  W.  1859.  Notes  on  North  American 
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1971.  Stratigraphy  of  Lower  Cretaceous  Trin- 
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Geol.  Rept.  Invest.  71:1-63. 


55 


Van  Straelen,  V.  1936a.  Contribution  a  I'etude 
des  crustaces  decapodes  de  la  periode  juras- 
sique.  Mem.  Acac.  r.  Belg.  CI.  Sci.  7:1^62. 

.     1936^.    Crustaces  decapodes  nouveaux 

ou  peu  connus  de  Tepoque  cretacique.  Bull. 
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strata  (Subsurface  and  Lower  Cretaceous) 
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Cretaceous  Crabs.  Palaeontographical  Soc. 
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TRANSACTIONS 
OF  THE  SAN  DIEGO 
SOCIETY  OF 
NATURAL  HISTORY 


Volume  20  Number  3  pp.  57-68  24  June  1983 


A  new  subspecies  of  Euphyes  vestris  (Boisduval)  from 
southern  California  (Lepidoptera:  Hesperiidae) 

John  W.  Brown  '-'PRAft'Y 

Enloniolog}'  Department.  San  Diego  Natural  History  Museum. 
San  Diego^  California  92112  USA 

r 

William  VV.  McGuire  ' 

Saint  Francis  Hospital.  Colorado  Springs.  Colorado  80903  USA 

Abstract.  Euphyes  vestris  harbisoni.  from  southern  California,  is  described  and  illustrated.  This 
subspecies  is  phenotypicalh  distinct  and  geographically  isolated  from  ail  other  populations  oi  E.  vestris 
(Boisduval).  Its  range  is  restricted  in  part  by  the  distribution  of  the  larval  host  plant,  Carex  spissa  Bailey 
(Cyperaceae).  Life  history  data  are  presented  including  descriptions  of  the  immature  stages.  Additionally, 
E.  ruricola  (Boisduval)  is  removed  from  the  synonymy  of  £".  vestris. 

Introduction 

Euphyes  vestris  (Boisduval  1852)  has  long  been  recognized  as  a  polytypic  species, 
with  subjective  separation  of  an  eastern  population,  Euphyes  vestris  Dietacoiuet  (Harris 
1862).  from  the  nominate  western  population.  Additionally,  it  has  been  appreciated 
by  some  that  a  phenotypically  distinct  population  of  £".  vestris  QxisXs  in  extreme  southern 
California  (Emmel  and  Emmel  1973).  Data  available  indicate  the  presence  of  this  insect 
in  scattered  populations  from  Orange  County  (Orsak  1977)  through  western  San  Diego 
County,  with  speculative  distribution  to  include  the  northern  mountain  ranges  of  Baja 
California  Norte.  Mexico  (MacNeill  1962).  However,  because  sampling  and  study  of 
the  southern  California  population  has  been  extremely  limited,  definitive  analysis  had 
not  been  previously  accomplished  and  its  status  remained  uncertain.  Based  on  the  data 
herein  presented,  it  is  now  obvious  that  the  population  of  E.  vestris  centered  in  San 
Diego  County.  California,  represents  a  morphologically  distinct  form  that  is  geograph- 
ically isolated  from  the  nearest  E.  vestris  populations  both  to  the  north  and  east. 
Accordingly,  we  propose  a  name  for  this  southwestern  population,  and  report  previously 
unknown  life  history  data. 

Depositories  abbreviated  in  the  text  are  as  follows:  RB.  Richard  Breedlove.  San 
Diego.  California:  GB.  Guy  Bruyea.  Poway.  California:  CS.  Chuck  Sekerman.  North 
Hollywood.  California:  LG.  Lee  Guidry.  Point  Loma.  California;  LACM.  Los  Angeles 
County  Museum  of  Natural  History:  SDNHM.  San  Diego  Natural  History  Museum; 
CIS.  California  Insect  Survey.  University  of  California.  Berkeley;  and  UCI,  University 
of  California.  Irvine. 

Systematics 

Considerable  controversy  and  confusion  exist  regarding  the  correct  specific  name 
of  Euphyes  vestris  (Boisdu\al).  The  2  nominal  taxa  ^  Ilesperia'!  vestri.s"  [sic]  and  '"Hes- 
peria  ruricola'  were  described  by  Boisduval  (1852)  from  specimens  collected  in  Cali- 
fornia by  P.  Lorquin.  The  2  have  become  confused  largely  owing  to  the  loss  of  the  type 
of  ruricola  (fide  Oberthiir.  Evans  1955);  the  type  of  vestris  is  reportedh  in  the  British 


58 


Museum  (Evans  1955).  Although  ruricola  has  page  priority,  the  absence  of  type  material 
and  the  inconclusive  nature  of  the  original  description  have  traditionally  relegated  this 
taxon  to  junior  synonym  ol^  vcstris.  However,  Evans  ( 1955)  indicates  that  the  description 
of  ruricola  in  "no  way"  agrees  with  vcstris.  Furthermore,  he  maintains  that  the  two 
taxa  are  actually  in  different  genera;  ruricola  belonging  to  the  genus  Hesperia  f"abricius, 
and  vcstris  correctly  placed  in  Euphyes  Scudder.  This  is  supported,  in  part,  by  the 
question  mark  placed  after  the  genus  ^' Hesperia"'  in  Boisduval's  original  description  of 
vcstris.  The  fact  that  Boisduval  questioned  the  congeneric  nature  of  vcstris  and  ruricola 
clearly  indicates  that  these  two  taxa  do  not  represent  the  same  species.  Therefore,  Evans 
was  correct  in  removing  ruricola  from  the  synonymy  of  vcstris.  F.  M.  Brown  (1957) 
states  "Holland  [(1905)]  was  responsible  for  the  confusion  when  he,  contrary  to  most 
of  the  evidence  and  the  original  description,  applied  [the  name]  ruricola  to  the  Dun 
Skipper  [E.  vcstris]  ....'' 

Miller  and  Brown  (1981)  recently  revived  E.  ruricola  "on  the  basis  of  a  specimen 
that  might  be  the  type  that  is  labelled  as  ^ruricola"  in  Boisduval's  hand  .  .  ."  which  had 
been  located  in  the  collection  of  the  Carnegie  Museum.  This  specimen  was  designated 
as  the  lectotype  in  1976  by  Brown,  Miller,  and  Clench,  according  to  the  lectotype  label 
(Carnegie  Museum  lectotype  no.  733).  This  type  designation,  however,  was  never 
published,  and  the  senior  author's  personal  examination  of  the  data  accompanying  the 
specimen  revealed  a  single  label  that  could  possibly  have  been  written  by  Boisduval  — 
"Californie  (Lorquin)."  Contrary  to  Miller  and  Brown  (1981),  there  is  no  evidence  to 
indicate  that  the  specimen  in  question  represents  Boisduval's  concept  of  ruricola.  Since 
a  legitimate  holotype  of  vcstris  exists  in  the  British  Museum  (Evans  1955),  and  the 
description  of  ruricola  is  obviously  not  consistent  with  vcstris,  we  propose  that  ruricola 
be  removed  from  the  synonymy  of  vcstris. 

In  agreeing  with  Stanford  (1981)  regarding  the  validity  of  the  subspecific  taxon  E. 
vestris  kiowah  (Reakirt  1866).  we  recognize  the  following  subspecies  of  £".  vcstris: 

Euphyes  vestris  (Boisduval  1852) 
E.  V.  v^'^ms  (Boisduval  1852) 

i=Pai}iphila  osceola  Lintner  1878) 

{^Painphila  califoruica  Mabille  1883) 
E.  V.  nictaconu't  (Harris  1862) 

{^Paiuphila  rurca  W.  H.  Edwards  1862) 

{^Hesperia  osyka  W.  H.  Edwards  1867) 
E.  V.  kiowah  (Reakirt  1866) 

In  addition  to  the  above,  the  distinctive  southern  California  population  of  E.  vcstris 
deserves  formal  recognition.  Extensive  review  of  the  synonymy  has  failed  to  uncover 
a  previous  description  that  could  be  unequivocally  attributed  to  this  southern  California 
population,  hence  we  propose  a  new  name. 

Euphyes  vcstris  harbisoni  new  subspecies 
Figures  1-4 

Type  material.  — \\o\o\ypQ  male  (SDNHM  type  L-49),  13.3  km  east  of  Dulzura, 
north  slope  of  Tecate  Peak,  elevation  500  m,  San  Diego  County,  California,  9  June 
1981  (J.  Brown):  allotype  (SDNHM  type  L-50),  13.3  km  east  of  Dulzura,  north  slope 
of  Tecate  Peak,  elevation  500  m,  San  Diego  County,  California,  14  June  1981  (J. 
Brown).  Paratypes:  18  6<5  and  16  99,  same  locality  as  holotype,  but  dates  as  follows:  1 
3,  1  9,  28  June  1980:  2  53,  2  99,  30  June  1980:  3  <5<5,  1  9,  12  July  1980:  4  99,  9  June 
1981:  4  33,  2  99,  14  June  1981:  2  33,  2  99,  21  June  1981:  1  9,  27  June  1981:  1  9,  ex- 
pupa,  2  June  1982:  1  9,  ex-pupa,  3  June  1982  (all  J.  Brown):  6  33,  19,  12  June  1982 
(L.  Guidry). 

Disposition  c^/O'/^^'-s.  — Holotype  male  and  allotype  are  deposited  in  the  collection 
of  the  San  Diego  Natural  History  Museum:  paratypes  distributed  among  collections  of 
the  following  institutions:  Los  Angeles  County  Museum  of  Natural  History:  California 
Academy  of  Sciences,  San  Francisco:  United  States  National  Museum  of  Natural  His- 


59 


Figure  1 .     Euphyes  vestris  harbisoni,  new  subspecies.  6.  dorsal.  1 3.3  km  east  of  Dulzura,  San  Diego  County, 

California.  12  June  1980. 

Fr.i  RE  2.     Kiiphvcs  vestris  harbisoni.  new  subspecies,  3,  ventral.  13.3  km  east  of  Dulzura.  San  Diego  Countv. 

California,  12  June  1980. 

FiCiURE  3.     Euphyes  vestris  harbisoni,  new  subspecies,  2,  dorsal,  1  3.3  km  east  of  Dulzura.  San  Diego  Countv, 

California,  6  June  1981. 

Fkure  4.     Euphyes  vestris  harbisoni.  new  subspecies,  2,  ventral.  13.3  km  east  of  Dulzura.  San  Diego  County, 
California.  6  June  1981. 


tory.  Washington.  D.  C;  Alhn  Museum  of  EntomologN .  Sarasota.  Florida:  Carnegie 
Museum  of  Natural  Histor>.  Pittsburgh.  Pennsyhania.  All  remaining  paratxpcs  to  be 
retained  by  the  San  Diego  Natural  History  Museum. 

Additionahualerial examined. -CALIFORNIA:  SAN  DIEGO  COUNTY:  No  fur- 
ther locality.  1  5.  5  June  1936.  1  2,  12  June  1936  (J.  Creelman,  RB).  1  2.  17  June  1936 
(J.  Creelman.  LACM).  1  3.  5  June  1936.  1  2.  7  June  1938  (J.  Creelman.  SDNHM).  .San 
Diego.  1  6.  4  July  1920  (O.  E.  Sette.  LACM).  San  Diego  City.  1  <3,  1  2.  14  June  1936 
(J.  Creelman,  LACM).  Adobe  Falls.  2  SS.  1  2.  13  June  1936  (F.  Thorne.  SDNHM). 
Flinn  Springs.  1  2.  1  June  1939.  1  c?.  1  5  June  1939.  6  (53.  1  2.  16  June  1939.  2  22.  18 
June  1939  (all  W.  P.  Medlar.  SDNHM).  1  3.  18  June  1939  (W.  P.  Medlar.  UCI),  8  <35, 
5  22,  18  June  1939  (W.  P.  Medlar.  LACM).  2  33.  10  June  1980.  1  3.  18  June  1981  (all 
J.  Brown,  SDNHM).  Avocado  orchard.  Flinn  Springs.  1  3.  ex-pupa.  15  June  1982  (J. 
Brown.  SDNHM).  Blossom  Valley,  8  km  WNW  Alpine.  2  66.  3  22.  17  June  1940.  2 
66.  29  June  1942  (all  F.  Thorne.  SDNHM).  Hellhole  Canyon.  1  3.  12  July  1954  (M. 
Kenney.  CIS).  Lower  Hellhole  Creek,  1  3.  1  3  June  1981  (Brown  and  Brown.  SDNHM). 
El  Cajon,  2  66,  29  June  1963  (O.  Shields.  LACM).  El  Monte  Oaks.  I  3.  30  May  1965 
(R.  Breedlove.  RB).  1.6  km  W  Tccate  turn-ofl.  1  3.  30  June  1980(D.  Faulkner.  SDNHM). 

5  km  NW  Fallbrook.  1  3.  23  May  1981.  1  3.  e.\-pupa.  1  June  1981.  1  2.  e.x-larva.  12 
June  1981  (all  Brown  and  Brown.  SDNHM).  Old  Viejas  Grade.  Poser  Mountain.  4  66. 
2  22.  12  June  1981.  2  22.  17  June  1981.  2  33,  26  June  1981  (all  J.  Brown.  SDNHM). 
Behind  San  Pasqual  Academy.  1  3.  30  May  1981  (D.  Faulkner.  SDNHM).  1  2,  ex- 
larva,  30  May  1982  (L.  Guidry.  LG).  Poway  Green  Valley  Truck  Trail.  1  3.  ex-larva. 

6  June  1982  (L.  Guidry.  LG).  Poway.  1  3,  ex-pupa.  15  May  1982  (C.  Sekerman.  CS). 
Lake  Poway.  1  2,  ex-pupa,  20  June  1982,  1  6.  ex-pupa,  27  June  1982,  I  6,  28  June 


60 


5 


6 


Figure  5. 

Figure  6. 
1958. 


Male  genitalia  of  Euphyes  vestris  harbisoni.  new  subspecies. 

Male  genitalia  of  nominate  Euphyes  vestris.  Plantation.  Sonoma  County,  California.  16  May 


1982,  1  9,  ex-pupa.  4  July  1982  (all  G.  Bruyea,  GB).  ORANGE  COUNTY:  Silverado 
Canyon,  1  <5.  29  June  1972  (C.  Sexton,  UCI).  2  <5(5,  8  June  1982  (J.  Brown,  SDNHM), 
2  <55,  1  9,  5  June  1982  (C.  Sekerman,  CS). 

Distribution.  — Currenlly  known  only  from  San  Diego  and  Orange  Counties.  Cal- 
ifornia. 

Diagnosis.  — Male:  forewing  length  x=  15.2  mm  (range  15.0-16.1  mm).  Head, 
thorax  and  abdomen  dark  brown,  covered  with  thin  light-brown  hairs,  much  lighter 
beneath.  Palpi  whitish  with  orange-brown  terminal  segment.  Antennae  gold-bi"own, 
ventral  surface  of  club  gold,  apiculus  brown.  Forewings  chocolate-brown  above,  with 
lustrous  orange  over-scaling  near  a  bold  black  stigma.  Stigma  composed  of  two  oval 
patches  forming  an  inconspicuously  broken  black  dash.  Outer  margin  of  forewing  with 
a  very  fine  light  tan  fringe.  Forewings  dull  brown  below,  with  basal  and  distal  blackening 
representing  the  undersurface  of  the  stigma.  Hindwings  same  color  as  forewings  above, 


61 


7 


50 


-S         40J 

TO 

3 

■o 

c 

Z         30 


(0 

a> 

E 

3 


20 


10 


j^t 


Males 
Females 


May 


June 


July 


August 


Figure  7.     Flight  period  graph  of  Eiiphycs  veslris  harbisoni.  new  subspecies,  based  on  wild  caught  adults, 
using  all  specimens  examined. 


but  with  less  lustrous  orange  over-scaling;  no  additional  markings.  Hindwings  dull 
brown  below,  with  no  markings. 

Male  genitalia:  as  illustrated  (Fig.  5).  The  structures  of  the  male  genitalia  of /f. 
vestris  harbisoni  are  very  similar  to  those  of  both  the  Arizona  populations  of  i:'.  vcstris 
kiowah  (not  illustrated)  and  the  northern  California  populations  of  E.  vcstris  vcstris 
(Fig.  6).  Compared  to  nominate  vcstris.  the  genitalic  capsule  of  harbisoni  is  propor- 
tionately longer  and  slightK  less  robust,  with  the  vahae  more  broadh  rounded,  and 
lacking  the  ventral  caudal  indentation  commonly  present  in  nominate  vcstris.  The  two 
genitalia  figured  represent  the  extremes  to  more  clearh  illustrate  these  differences. 

Female:  forewing  length  x  =  16.0  mm  (range  14.7-17.0  mm).  As  in  male  except 
for  primary  and  secondary  sex  characters.  Forewings  dark  brown  abo\c,  with  lustrous 
orange  over-scaling.  Two  moderately  well-defincd  h\aline  post-median  spots  just  below 
the  discal  cell.  Females  generally  with  more  rounded  outer  margin  of  forewing.  Fore- 
wings  dull  brown  below,  with  blackening  in  basal  areas  slighth  more  diffuse  than  in 
male:  the  two  h\aline  spots  less  well-dcfmed.  Hindwings  abo\e  as  in  male:  dull  brown 
below,  occasionally  with  traces  of  light  spots  in  the  discal  band  area. 

Discussion.  — The  southern  California  population  of  Euphycs  vcstris  is  single- 
brooded,  the  flight  period  extending  from  late  May  through  mid-July  (Fig.  7).  The 
most  distinguishing  phenotypic  character  of  the  southern  California  population  is  its 


62 


8 


NO.  ADULTS     MEAN  FOREWING     STANDARD 
POPULATION  STUDIED       EXAMINED  LENGTH  (mm)  DEVIATION      "P"  VALUE 


Oregon  cT 
San  Diego  Ck). 

d" 

25 

40 

13.524 
15.155 

0  5659 
0  5296 

<0  001 

Oregon    § 
San  Diego  Co. 

9 

7 
22 

14.000 
15.9773 

07024 
0.6962 

<  0.001 

Arizona    (^ 
San  Diego  Co. 

d' 

30 
40 

14.137 
15.155 

0.5648 
0.5296 

<  0.001 

Arizona  O 
San  Diego  Co. 

9 

19 
22 

14.411 
15.9773 

0.4736 
0.6962 

<  0.001 

Arizona   (^ 
Oregon    (^ 

30 
25 

14.137 
13.524 

0.5648 
0.5659 

<  O.OIO(NS) 

Arizona    ^ 
Oregon    § 

19 

7 

14.411 
14.000 

0.4736 
0.7024 

<  O.IOO(NS) 

Figure  8.  Statistical  analysis  of  forewing  lengths  of  various  Euphyes  vcstris  populations  using  Student's  T- 
test  comparing  two  sample  means.  "P"  value  represents  the  probability  that  the  two  samples  are  homogeneous. 
"NS"  =  not  statistically  significant. 


larger  size  relative  lo  populations  from  northern  California.  Oregon  and  Washington. 
Its  larger  size  and  extensive  orange  over-scaling  clearly  distinguish  it  from  the  Rocky 
Mountain  E.  vestris  populations,  which  have  been  referred  to  as  E.  vestris  kiowah  by 
some  authors  (Stanford  1981),  and  the  eastern  E.  vestris  nietaconiet.  There  are  some 
populations  of  E.  vestris  in  Arizona  that  may  be  referable  to  E.  vestris  kiowah.  but  at 
this  time  have  not  been  thoroughly  studied.  Again,  these  differ  from  the  southern 
California  population  in  both  size  and  maculation.  An  analysis  of  the  male  and  female 
forewing  measurements  between  various  populations  sampled  (Fig.  8)  demonstrates 
the  statistically  significant  difference  between  the  San  Diego  County  population  and 
others  based  on  this  parameter. 

The  habitat  occupied  by  the  southern  California  population  is  remarkably  dissim- 
ilar to  that  of  all  other  known  populations  of  £■.  vestris.  The  southern  California  pop- 
ulation is  extremely  local,  generally  occurring  in  chaparral  or  riparian  communities  in 
narrow  canyons  where  there  is  a  seep  or  spring  providing  perennial  water  (Figs.  9- 
12):  populations  found  to  date  have  been  at  elevations  below  650  m.  The  o\iposition 
substrate  and  lar\  al  hostplant  is  Carex  spissa  Bailey  (C\  peraceae),  with  which  the  adults 
are  commonly  associated  (Fig.  13).  As  indicated  by  figure  14.  C.  spissa  has  a  limited 
and  spotty  distribution  from  San  Luis  Obispo  County.  California,  to  Baja  California 
Norte.  Mexico  (Munz  1974).  The  range  of  the  host  plant  is  obviously  a  factor  limiting 
the  distribution  of  the  butterflN.  as  supported  by  the  fact  that  all  known  populations 
of  E.  vestris  liarlusoni  (Fig.  15)  fall  within  the  range  of  C.  spissa.  Howc\er.  factors 
other  than  host  plant  availability  must  contribute  to  restrict  the  insect's  range,  as 
indicated  by  the  fact  that  the  butterfly  occurs  within  only  a  small  portion  of  the  host 
plant's  distribution. 

Initial  observations  and  efforts  to  study  this  insect  centered  on  examination  of 
previously  captured  material  and  old  collection  records.  Howcxer,  there  were  few 
previous  records  for  the  southern  California  population  of  Ii.  vestris.  and  the  last 
specimen  collected  prior  to  this  study  was  in  1 969  (Stanford  ///  litt.).  Many  prior  records 
came  from  the  Adobe  Falls  area  near  San  Diego  State  University,  but  current  devel- 
opment in  this  area  has  altered  the  habitat  to  the  point  where  it  no  longer  supports  the 


63 


Fku.re  9.  Typical  chaparral  habitat  near  Tccatc  Peak.  San  Diego  County.  C'alilbrnia. 

Fk.iki    10.  Habitat  near  Dul/ura.  Caiirornia. 

Fu.i  HI    11.  Seep  area  at  Dul/ura  k)cality. 

FicL  HI   12.  Habitat  near  Dul/ura.  California. 

Fk.iki   13.  Larval  host  C'(/'c\  s/j/sv^/  HailcN. 

insect.  The  only  fccent  coUcclions  outside  of  San  Diego  C"ouni\   were  made  \n  the 
Silverado  Canyon  area  of  Orange  County  (Orsak  1977). 

Using  prior  collection  records  as  a  guide,  we  examined  iikeh  habitats  in  1980 
and  1981  in  San  Diego  County  and  I3aja  California  Norte.  .Adult  specimens  were  found 
first  near  Flinn  Springs  Count\  Park,  and  additional  specimens  later  found  in  other 
localities  noted  abo\  e.  Fourth  instar  lar\  ae  were  subsequentK  observed  1  3.3  km  east 


64 


14 


San  Francisco 


FiC.URE   14. 

Mexico. 


Approximate  distribution  of  Carex  spissa  in  southern  California  and  adjacent  Baja  California, 


of  Dulzura  in  January  198  1.  All  larvae  were  found  in  typical  hibernacula  formed  from 
C.  spissa  (Fig.  16).  These  hibernacula  were  formed  by  the  attachment  of  two.  three  or 
four  leaves  creating  a  silk-lined,  tube-like  chamber  7-12  cm  long  and  open  at  the 
superior  end.  The  chamber  itself  is  usually  located  in  the  superior  one-third  of  the 
Carc.x  leaves.  Each  leaf  or  blade  is  characteristically  bent,  generally  at  less  than  a  100° 
angle,  allowing  them  to  be  pulled  together  and  joined  with  silk  by  the  larva.  This  type 
of  hibernaculum  is  fairly  characteristic  and  consistent  with  previous  observations  made 
on  other  members  of  this  genus,  including  /:.  a/aha  mac  {Lindsey  1923).  /:.  macguirei 
(Freeman  1975).  and  /:.  ciukcsi  (Lindsey  1923)  (William  McGuire  personal  observa- 
tion). 

Jjynio/ogy.— The  subspecies  /:.  vestris  harhisoni  is  named  in  honor  of  Charles 
Harbison,  Curator  Emeritus.  San  Diego  Natural  History  Museum.  Mr.  Harbison  has 
been  a  patient  instructor,  and  enthusiastic  field  entomologist,  and  a  pioneer  in  inves- 
tigating the  entomofauna  of  Baja  Califorinia,  Mexico. 


65 


BFallbrook 


mvEnsiDE  CO. 

SAIM    OIEGO   CO. 


■  Escondido 


HFIinn  Springs 


••    Tecate 


FiCii'RF  15.     Distribution  of  Euphyes  vcstris  harhisoiii.  new  subspecies. 


Life  History 


Early  stages.  — The  early  stages  of  E.  vestris  harbisoui  closely  resemble  those  de- 
scribed by  Heitzman  (1964)  for  the  eastern  E.  vestris  metacomet.  The  larvae  of  E. 
vestris  harbisoni,  however,  are  much  larger,  and  the  pupae  dark  brown  rather  than 
green. 

Egg.  — (Fig.  17).  1.5  mm.  hemispherical,  pale  yellow-green  with  an  irreg- 
ular red  splotch  on  the  apex.  An  irregular  red  band  encircling  the  entire  egg  midway 
between  the  base  and  the  apex. 

Eirst  inslar.  — Length  7.0  mm.  Head:  shiny  brown.  Body:  ground  color  translucent 
green,  abdominal  segments  9  and  10  translucent  yellow-tan.  Several  long,  fme  hairs 
projecting  upwards  from  the  last  body  segment:  thin  black  collar  just  behind  the  head 
indicating  the  prothoracic  shield. 

Second  inslar.  —  LengXh  14.0  mm.  Head:  yellow-orange  with  a  conspicuous  cara- 
mel-colored band  encircling  the  head  along  its  outer  margins  except  above  the  man- 
dibles. Mandibles  dark  brown.  An  oblong  dark  brown  spot  set  vertically  in  the  upper 
center  of  the  face.  Body:  ground  color  translucent  green,  abdominal  segments  9  and  10 
translucent,  revealing  yellow-tan  internal  organs.  Body  co\ered  with  numerous  white 
setae,  several  longer  thin  hairs  at  end  of  the  last  segment.  Each  spiracle  indicated  by  a 
fine  black  dot. 

Third  instar.  —  Lcnglh  21.0  mm.  Head:  cream-colored  with  a  well-defined  dark 
caramel-colored  longitudinal  stripe  on  each  side  of  the  face,  and  a  light  brown  band 
extending  from  the  dark  spot  to  the  mandibles.  Body:  ground  color  translucent  green, 
abdominal  segments  9  and  10  more  translucent;  covered  with  fine  white  hairs.  Spiracles 
indicated  bv  a  fine  black  dot. 


66 


Figure  16.     Typical  hibernaculum  formed  from  Carex  spissa.  Arrow  indicates  position  of  the  larva. 


Fourth  />75/(3r  —  Length  28.0  mm.  Head:  caramel-brown  with  an  oblong  black  spot 
in  the  upper  center  of  the  face;  a  narrow  cream-colored  longitudinal  band  running 
parallel  to,  and  on  each  side  of  the  black  spot;  a  cream-colored  band  around  the  outer 
edge  of  the  jaws.  Body:  ground  color  green,  with  numerous  thin  white  dashes  of  in- 
consistent length  (previously  visible  only  under  microscopy). 

Final  instar.  — (Fig.  18).  Length  36.0  mm.  Head:  caramel-brown  with  two  cream- 
colored  vertical  bands;  a  black  oval  spot  in  the  upper  center  of  the  face.  Body:  green 
with  extremely  thin,  wavy  longitudinal  white  dashes.  A  single,  subtle,  darker  green 
longitudinal  stripe  along  the  center  of  the  dorsal  surface.  Minute  black  dots  sprinkled 
over  entire  body.  Terminal  abdominal  segment  translucent  pale  green.  Prothorax  white 
with  prothoracic  shield  indicated  by  a  thin  black  line  running  into  the  enlarged  first 
spiracle  dot,  forming  a  thin  black  wedge.  Each  spiracle  indicated  by  a  black  dot,  larger 
for  the  first  and  anal  spiracles.  Ventral  surface  of  abdominal  segments  8  and  9  covered 
with  a  fine  layer  of  silver-white  cottony  material. 

Pupa.  — (Fig.  19).  Length  25.0  mm.  Dark  brown.  Abdomen  slightly  dusted  with 
white  cottony  material.  Most  of  body  covered  with  short,  fine  bristles;  wing  cases  bare. 
Bristles  on  head  and  end  of  abdomen  longer.  Tongue  case  long,  extending  beyond  the 
fourth  abdominal  segment,  lying  along  the  mid-dorsal  surface  of  the  pupa,  detached 
above  the  abdominal  segments. 

Behavior.  — Eggs  are  laid  singly,  usually  on  the  undersurface  of  the  leaf  blades  near 
the  base  of  the  plant.  During  the  first  instar  the  young  larvae  are  generally  found  along 
the  mid-ridge  crease  on  the  upper  surface  of  the  leaves,  near  the  base  of  the  plant.  The 
second  and  third  instar  larvae,  when  not  feeding,  are  usually  found  in  silk-lined  tubes 
constructed  by  the  larvae  by  attaching  two  to  four  leaves  together  longitudinally.  It  is 
during  the  fourth,  or  occasionally  the  third  instar,  that  the  larvae  construct  the  over- 
wintering hibernacula  previously  described,  and  enter  a  short  diapause.  The  head  is 
always  aimed  towards  the  superior  end  of  the  hibernacula. 

During  the  last  instar  the  larva  consumes  the  portion  of  the  leaf  blades  extending 
beyond  the  superior  end  of  this  hibernaculum.  This  tube  is  then  abandoned,  and  the 
larva  moves  to  a  dilTcrent  location  on  the  plant,  or  to  a  different  plant  and  constructs 
a  new  chamber.  The  upper  portion  of  these  leaf  blades  are  also  usually  eaten  by  the 


67 


18 


FiGi'RE  17.     Egg  of  Euphyes  vestrls  harbisoni.  new  subspecies. 

Ficii  RE  18.     Last  instar  larva  of  E  up  fives  vestris  harbisoni.  new  subspecies. 

Fita  RE  19.     Pupa  of  Euphyes  vestris  harbisoni,  new  subspecies. 


larva.  When  a  larva  is  ready  to  pupate,  it  fills  the  upper  end  of  the  tube  with  a  flocky. 
cottony  plug  8-10  mm  thick.  The  larva  is  generally  situated  in  the  tube  with  the  head 
towards  the  superior  end:  the  pupa  is  always  situated  in  this  manner.  Pupation  lasts 
18  to  21  days. 

Adults  emerge  from  late  May  through  June,  the  flight  period  extending  into  early 
July.  At  one  locality  a  last  instar  larva,  a  pupa,  and  a  single  adult  were  collected  on  the 
same  day  (23  May  1981). 

Males  commonly  ''patrol"  the  canyons,  never  venturing  far  from  the  vicinit\  of 
the  host  plant.  Both  sexes  are  avidly  attracted  to  nectar  sources  which  include  morning 
glory  {Calystegia  macrostegia  tcnuifolia  (Abrams)  Brummitt),  red  thistle  (Clrsiuni  oc- 
cidentale  (Null.)  Jepson).  loosestrife  (Lythrum  californicwn  T.  &  G.),  and  rarely,  golden 
yarrow  {Eriophyllum  confcrlifloruni  (D.  C.)  Gray)  and  black  mustard  (Brussica  nigra 
(L.)  Kock.).  Commonly  females  are  observed  perching  on  the  C'arc.x  and  basking  in 
the  late  morning  and  early  afternoon  sun. 

Conclusion 

Euphyes  vestris  harbisoni  is  isolated  by  geograph\  and  by  the  availability  of  a  select 
host  plant.  While  several  of  the  populations  found  appear  to  be  stable,  all  are  extraor- 
dinarily small.  Some  of  the  prior  known  colonies  have  been  extirpated  by  construction 


68 


projects  and  other  habitat  modifications.  A  search  for  new  populations  and  extension 
of  range  should  appropriately  be  directed  into  northern  Baja  California  and  northward 
up  to  and  including  San  Luis  Obispo  County,  thus  encompassing  the  known  range  of 
the  larval  host  plant.  Carex  spissa.  This  search  in  the  projected  northern  range  of  the 
insect  is  important  when  considering  potential  intergradations  with  the  nofthern  Cal- 
ifornia E.  vestris  vestris.  E.  vesths  harbisoni  appears  to  be  morphologically  and  bio- 
logically distinct,  with  significant  geographic  separation  from  all  other  known  E.  vestris 
populations.  This  restricted  range,  coupled  with  the  continued  threat  of  habitat  deg- 
radation as  has  been  demonstrated  in  the  immediate  San  Diego  city  area,  makes  it 
imperative  that  careful  monitoring  be  continued. 

Acknowledgments 

We  are  most  grateful  to  the  following  for  providing  assistance  and  encouragement 
during  the  course  of  this  study:  David  Faulkner,  Dr.  Reid  Moran.  Dr.  Ray  Stanford. 
Dr.  John  Emmel.  and  Dr.  Amadeo  Rea.  Special  thanks  go  to  Poody  Brown.  William 
Smithey,  and  John  Johnson,  all  of  whom  provided  photographic  assistance,  and  to 
Marie  HoflT  Steinauer.  and  Shirley  Latislaw  for  providing  the  maps  and  graphs. 

We  would  also  like  to  thank  the  following  for  providing  specimens  and/or  collecting 
records:  Richard  Breedlove.  San  Diego.  California;  Fred  Thorne.  Curator  Emeritus, 
San  Diego  Natural  History  Museum;  Julian  Donahue,  Entomology  Section,  Los  Angeles 
County  Museum;  Guy  Bruyea.  Poway.  California;  Lee  Guidry.  Point  Loma,  California; 
Chuck  Sekerman.  North  Hollywood,  California;  and  Gordon  Marsh.  University  of 
California,  Irvine.  Additionally,  we  would  like  to  thank  Dr.  Ginter  Ekis.  Carnegie 
Museum,  for  the  loan  of  the  unpublished  supposed  lectotype  of  Euphycs  rwicola. 

Finally,  we  would  like  to  acknowledge  the  Xerces  Society  for  their  financial  support 
of  this  project,  and  their  recognition  of  the  sensitive  status  of  this  insect.  Their  support 
allowed  one  of  us  (John  W.  Brown)  to  undertake  the  extensive  field  work  which  made 
this  study  possible. 


Literature  Cited 


Boisduval,  J.  A.  1852.  Lepidopteres  de  la  Cali- 
fornie.  Ann.  Soc.  Ent.  France,  ser.  2,  10:315, 
317. 

Brown.  F.  M.  1957.  Colorado  butterflies,  part  5. 
Proc.  Denver  Mus.  Nat.  Hist.  7:313. 

Emmel.  T.  C.  and  J.  F.  Emmel.  1973.  Butterflies 
of  southern  California.  Nat.  Hist.  Mus.  Los 
Angeles  Co..  Sci.  Ser.  26:1-148. 

Evans.  W.  H.  1955.  A  Catalogue  of  the  American 
Hesperiidae  in  the  British  Museum  (Natural 
History),  part  4.  London,  Trustees  British  Mu- 
seum. 499  pp. 

Heitzman.  R.  J.  1964.  The  early  stages  of  Ew/?/?;^^^ 
vestris.  J.  Res.  Lepid.  3(3):  15 1-1 53. 

MacNeill,  C.  D.  1962.  A  preliminary  report  on 
the  Hesperiidae  of  Baja  California.  Proc.  Calif. 
Acad.  Sci.  30(5):91-1 16. 


Miller.  L.D.  and  F.  M.Brown.  1981.  A  catalogue/ 
checklist  of  the  butterflies  of  America  north  of 
Mexico.  Lepid.  Soc.  Mem.  2:1-280. 

Munz,  P.  A.  1974.  A  Flora  of  Southern  California. 
Univ.  Calif  Press,  Berkeley.  Los  Angeles,  Lon- 
don. 

Orsak,  L.  J.  1977.  Butterflies  of  Orange  County. 
Univ.  Calif  Irvine.  Mus.  Syst.  Biol.  Res. 
ser.  4. 

Stanford.  R.  E.  1981.  //;  Ferris,  C.  D..  and  F.  M. 
Brown  (editors).  Butterflies  of  the  Rocky 
Mountain  States.  Univ.  Oklahoma  Press,  Nor- 
man. 


TRANSACTIONS 
OF  THE  SAN  DIEGO 
SOCIETY  OF 
NATURAL  HISTORY 


Volume  20  Number  4  pp.  69-79  24  June  1983 


Two  new  idoteid  isopods  from  Baja  California  and  the  Gulf  of 
California  (Mexico)  and  an  analysis  of  the  evolutionary  history 
the  genus  Colidotea  (Crustacea:  Isopoda:  Idoteidae)  fARY 

Richard  C.  Brusca 

Department  of  Marine  Invertebrates,  San  Diego  Natural  History  Museum. 
San  Diego,  California  92112  iSA 

Abstract.  Two  new  species  of  idoteine  isopod  crustaceans  are  described  from  Baja  California, 
Colidotea  wallersteini  new  species  and  Synidoteafrancesac  new  species  (Crustacea:  Val\ifera:  Idoteidae: 
Idoteinae).  The  genus  Colidotea  is  rediagnosed.  a  key  to  the  known  species  presented,  and  its  evolution 
and  historical  biogeography  described.  Both  trans-oceanic  dispersal  and  vicariance  phenomona  appear 
to  have  played  roles  in  the  establishment  of  modern  distributional  patterns  of  Colidotea. 

Introduction 

The  idoteid  isopod  fauna  of  the  northeast  Pacific  has.  in  the  past  10  years,  come 
to  be  quite  well  known  (see  Brusca  and  Wallerstein  1979b  for  an  introduction  to  the 
literature).  The  present  study  adds  2  new  species  to  this  fauna,  based  on  material 
collected  by  Dr.  D.  G.  Lindquist  (University  of  North  Carolina)  and  some  previously 
unsorted  samples  of  microcrustaceans  at  the  Allan  Hancock  Foundation  (University 
of  Southern  California). 

The  terminology  used  in  this  paper  is  standard  for  the  idoteid  isopod  literature 
(see  Menzies  1950  for  a  summary).  The  following  abbrev  iations  arc  used:  AHF.  Allan 
Hancock  Foundation;  USNM,  National  Museum  of  Natural  History;  SDNHM.  San 
Diego  Natural  History  Museum. 

Systematics 

Order  Isopoda 

Suborder  Valvifera 

Family  Idoteidae 

Subfamily  Idoteinae 

Genus  Synidotca  Harger,  1878 

The  systematics  and  biogeography  of  the  genus  Synidotca  were  reviewed  by  Men- 
zies and  Miller  (1972).  who  recognized  36  valid  species  and  subspecies.  8  of  which 
occur  in  California.  Iverson  (1972)  described  a  ninth  species  from  California  {S.  Dwdia). 
Brusca  and  Wallerstein  (1979a)  presented  new  distributional  records  for  S.  harfordl 
Benedict,  1897,  which  at  that  time  was  believed  to  be  the  southernmost  ranging  species 
of  the  genus  in  the  northeastern  Pacific.  They  noted  its  disjunct  occurrence:  central 
California  to  Magdalena  Bay  (SW  Baja  California),  Mexico,  reappearing  again  m  the 
central  Gulf  of  California.  Mexico  (2  known  records).  Brusca  and  Wallerstein  (1979b) 
reviewed  the  distributions  of  the  7  known  northeastern  Pacific  shallow-water  Synidotca. 
noting  that  all  but  S.  harfordi  were  entirely  restricted  to  the  cold-temperate  waters  of 
the  Arctic.  Aleutian  and  Oregonian  Provinces.  Brusca  (in  press)  discusses  phylogenetic 


70 


Figure  1.     Synidotea  francesae  new  species.  Holotype,  AHF  736.  Female. 

relationships,  evolutionary  history  and  zoogeography  of  Synidotea.  as  well  as  the  21 
other  genera  of  Idoteinae. 


Synidotea  francesae  new  species 
Figures  1,  2.  3 

Types.  — HoloXype:  female.  AHF  736;  Allotype:  male.  AHF  736a.  Paratypes:  1 
female,  SDNHM;  1  female,  USNM. 

Locality.  — AW  specimens  from  single  collection:  Mexico,  Gulf  of  California,  So- 
nora.  El  Golfo  dc  Santa  Clara  (about  4  miles  SE  of  town);  found  on  sandy  beach, 
"scavenging"  on  cast  up  tubes  of  Chaetopterus  species  (Polychaeta);  air  temperature 
24°C;  water  temperature  (surO  19°C;   17  April   1973;  collected  by  D.  G.  Lindquist. 

Diagnosis.  — CQp\\3.\or\  without  horns  or  other  projections;  frontal  margin  convex; 
eyes  bulge  outward;  body  without  tubercles  or  rugae;  plcotclson  wider  than  long;  per- 


71 


FidiRF  2.  Synlciotca  francesac  ncv,'  species.  Holotype.  AHF  736.  Female,  a.  antenna  1.  b.  antenna  2.  c,  left 
mandible,  d,  lacinia  of  right  mandible,  e,  maxilla  1.  f.  maxilla  2.  g.  maxilliped.  h.  pereopod  I.  i.  pereiopod 
IV.  J.  pereopod  VII.  k.  uropod. 


72 


Figure  3.     Synidotea  francesae  new  species,  a,  pleopod  I  (holotype).  b,  pleopod  2  (holotype).  c,  appendix 
masculinum  of  pleopod  2.  male  (allotype),  d,  pleopod  5  (holotype). 


eonites  V-VII  manifestly  shorter  than  I-IV;  median  dorsal  pattern  rounded,  as  in  S. 
laticauda;  dorsum  with  heavy  pigmentation. 

Description.  — hQxxgXh  to  at  least  6  mm;  body  ovate  and  darkly  pigmented:  dorsum 
smooth. 

Cephalon:  Lacking  horns  or  other  projections;  frontal  margin  convex,  although 
anterolateral  angles  somewhat  expanded;  eyes  elevated,  on  distinct  bulges.  Antenna  1 
with  tri-articulate  peduncle  and  uniarticulate  flagellum;  distal  third  of  terminal  article 
with  simple  setae  and  esthetascs.  Antenna  2  with  5-articulate  peduncle  and  8-articulate 
flagellum;  all  articles  with  setae;  articles  4  and  5  of  peduncle  each  with  distinct  distal 
patch  of  chromatophores;  flagellum  quite  short,  extended  only  to  pereonitc  II.  Max- 
illiped  typical  for  the  genus,  with  single  coupling  hook.  Maxilla  1  typical  for  the  genus; 
inner  lobe  with  2  stout  plumose  setae;  outer  lobe  with  6  long  simple  setae  and  4  short 
pectinate  setae.  Maxilla  2  typical  for  the  genus,  with  plumose,  simple  and  comb  setae 
as  figured.  Mandible  with  4-toothed  incisor  and  large  molar  process,  the  latter  smooth 
on  outer  margin  and  toothed  on  inner  margin;  lacinia  mobilis  of  left  mandible  3- 
toothcd;  lacinia  of  right  mandible  3-toothed  but  with  additional  large  serrate  spinelike 
process. 

Pereon;  Manifestly  elevated  along  middorsal  line;  entire  dorsal  surface  with  dark 
pigmentation,  except  along  lateral  margins.  Pereonites  I-IV  large,  long  and  without 
dorsal  coxal  plates;  pereonites  V-VII  small,  short,  and  with  large  dorsal  coxal  plates, 
visible  in  dorsal  aspect  by  presence  of  faint  suture  lines  just  median  of  edge  of  chro- 
matophorc  pattern;  all  pereopods  form  ventral  coxal  plates  that  meet  in  midline  of 
sterna.  Median  dorsal  pattern  of  pereonites  II-IV  {scusu  Menzies  and  Miller  1972)  with 
rounded  posterior  margins.  Pereonites  I  V-VII  slightly  produced  along  posteromedial 
margin.  Pereopod  I  with  dactyl  nearly  as  long  as  propodus. 

Picon;  Posterior  border  with  faint  indentation,  otherwise  evenly  convex.  Pleon 
(=plcotelson)  width  equal  to  length,  or  up  to  1.5  times  wider  than  long.  Plcopods  1-3 


73 


Table  1.     Principal  attributes  distinguishing  Colidotca  and  its  sister-group.  Synisoma.  from  each  other  and 
from  all  other  genera  of  Idoteinae.* 


Colidotea 

Synisoma 

Flagella  of  second  an- 

multiarticulate 

multiarticulate 

tennae 

Number  of  articles  in 
maxillipedal  palp 

Appendix  masculina  (of 
male) 

4 

long,  arising  from  base  of  endo- 
pod  (of  second  pleopod) 

4 

long,  arising  from  base  of  endopod 
(of  second  pleopod) 

Lacinia  mobilis 

very  large;  as  large  or  nearly  as 
large  as  incisor  process 

very  large;  as  large  or  nearly  as  large 
as  incisor  process 

Pleon 

relatively  short:  much  less  than 
'A  total  body  length 

relatively  long;  'A  or  more  total 
body  length 

Pleomere  fusion 

all  pleomeres  fused;  one  pair  lat- 

all pleomeres  fused;  no  lateral  inci- 

eral incisions  present 

sions  remain 

Eyes 

very  small  (except  in  C.  findleyi); 
set  on  lateral  margins  of  cepha- 
lon 

very  small;  set  on  lateral  margins  of 
cephalon 

For  comparable  data  on  the  other  20  genera  of  Idoteinae  see  Brusca  (in  press). 


with  plumose  marginal  setae,  decreasing  posteriorward;  pleopods  4-5  without  plumose 
marginal  setae.  Uropod  with  3  stout  plumose  setae  at  junction  of  protopod  and  lamella. 
Appendix  masculinum  of  male  simple,  with  weakly  grooved  inner  margin  and  a  few 
distal  setae;  penes  entirely  fused  into  large  tongue-like  flap. 

Ety}}iology\  —  Synidotea  fraucesae  is  named  in  honor  of  Frances  Runyan:  biological 
illustrator,  horticulturist,  and  friend. 

Discussion.— Synidoicajrancesae  is  similar  to  S.  laticaucia  Benedict,  1897,  and  S. 
harfordi  Benedict.  1897  (both  known  from  California  shoi"es),  in  having  a  smooth 
dorsum  with  a  characteristic  medial  dorsal  pattern,  evenly  convex  lateral  margins  on 
the  body  somites,  elevated  (bulging)  eyes,  and  notch  at  the  apex  of  an  otherwise  evenly 
convex  pleotelson.  It  differs  most  strikingly  from  these  species  in  having  a  pleon  as 
wide  or  wider  than  long,  a  barely  perceptable  notch  at  the  pleotelson  apex  (rather  than 
a  deep  notch),  pereonites  V-VII  manifestly  reduced  in  size,  and  a  distinct  dorsal  chro- 
matophore  pattern.  It  differs  further  from  5".  laticauda  in  having  an  evenly  convex 
tVontal  margin,  a  4-toothed  incisor  on  the  mandible,  and  shorter  antennae.  It  differs 
further  from  S.  harfordi  in  having  rounded  posterior  margins  on  the  medial  dorsal 
pattern  of  pereonites  II-IV,  shorter  antennae,  and  a  coupling  hook  on  the  maxillipedal 
endite. 

Of  the  18  species  o^  Synidotea  now  known  from  the  eastern  Pacific,  only  S.  fran- 
ccsac  is  a  regular  resident  of  warm  waters  (i.e.,  the  Eastern  Pacific  Zoogeographic 
Region);  all  others  are  temperate  or  polar  species.  The  genus  is  worldwide  in  distribution 
but  notably  absent  from  the  New  World  tropics.  Brusca  and  Wallerstein  (1979b)  and 
Wallerstein  and  Brusca  (1982)  hypothesized  that  this  absence  is  due  to  the  inability  of 
Synidotea  species  to  develop  certain  morphological  (size;  body  spination)  and  life 
history  (early  reproduction;  predator  avoidance  behaviors)  adaptations  that  have  al- 
lowed successful  radiation  in  this  region  by  other  genera  (e.g.,  Erichsoiwlla.  Eusyni- 
nicriis,  FarasyniDicrus).  and  that  any  species  of  Idoteidae  that  may  be  found  successfully 
inhabiting  the  warm  coastal  waters  of  the  tropical  eastern  Pacific  would  have  had  to 
evolve  some  of  these  adaptations.  In  the  subtropical  waters  of  the  Gulf  of  California. 
S.francesae  has  evolved  at  least  1  (small  size)  if  not  more  (e.g.,  early  reproduction)  of 
these  attributes  which  presumably  ha\e  played  a  role  in  its  ability  to  survive  the 
predation-intense  summers  of  this  region.  Menzies  and  Miller  (1972)  state  that  the 
mean  length  of  the  8  Arctic  species  ol\Synidolca  is  18.8  ±  2.6  mm,  while  the  mean 
length  of  the  5  tropical  species  then  known  (all  from  the  Old  World)  was  8.0  ±  2.2 
mm.  Temperate  species  fall  between  these  two  extremes.  By  comparison,  the  largest 
specimen  oWSynidoteafrancesae  examined  is  onh  6  mm  in  length. 


74 


Genus  Colidotea  Richardson,  1905 

The  genus  Colidotea  was  briefly  reviewed  by  Brusca  and  Wallerstein  (1979b).  The 
genus  now  contains  4  species:  C.  wstrata  (Benedict),  southern  California;  C.  findleyi 
Brusca  and  Wallerstein.  Baja  California;  C.  edniondsoni  Miller.  Hawaii;  arfid  C.  wal- 
Icrsteifii  new  species.  Baja  California  (Fig.  7).  Brusca  (in  press)  discusses  the  phylogeny, 
evolutionar\  history  and  zoogeography  of  Colidotea  and  its  sister-group.  Synisonia 
Leach.  Both  genera  are  restricted  to  warm-temperate  and  subtropical  waters  of  the  New 
World  (Colidotea)  and  Old  World  {Synisoma).  Table  1  summarizes  the  principal  fea- 
tures that  distinguish  these  2  genera  from  one  another  and  from  all  other  genera  of 
Idoteinac. 

A  reexamination  of  the  4  known  species  of  Colidotea  warrants  an  expanded  di- 
agnosis of  the  genus  and  a  key  to  the  species,  as  follows. 

Diag}iosis.  —  \doXQ'maQ  with  body  compact,  pereon  merging  smoothly  with  pleon. 
Cephalon  with  anterolateral  angles  produced;  with  or  without  a  medial  cephalic  process 
or  spine;  eyes  lateral.  Second  antennae  with  flagella  comprised  of  a  few.  or  a  dozen  or 
so  articles,  the  number  increasing  somewhat  with  age  (length).  Maxillipedal  palp  of  4 
free  articles,  apical  article  large  and  ovate;  epipod  present;  endite  with  or  without 
coupling  hooks.  Mandible  with  large  lacinia  mobilis.  nearly  as  large,  or  as  large  as 
incisor  process.  Pereon  with  coxal  plates  present  on  segments  II-VII;  coxae  may  or 
may  not  be  visible  in  dorsal  aspect,  or  may  be  visible  only  on  posteriormost  perconites. 
Pleon  comprised  of  single  piece,  with  one  pair  of  lateral  incisions  (=suture  lines);  pleon 
length  less  than  '/i  total  body  length;  uropods  uniramous.  Appendix  masculinum  of 
male  arising  from  base  of  pleopod  2  endopod.  Uropods  uniramous. 

Key  to  the  Species  of  Colidotea 

1.  Cephalon  with  medial  spine  or  process,  which  may  or  may  not  project  over 
supra-antennal  line 2 

-  Cephalon  without  medial  spine  or  process    3 

2.  Cephalic  process  bifurcate  (2-pronged);  frontal  process  subacute  (convex);  pleo- 
telson  with  posterior  margin  angulate;  bases  of  pereopods  with  dark  chromato- 
phore  band  C  edmondsoni 

-  Cephalic  process  simple  (single  apex);  frontal  process  notched  (concave);  pleo- 
telson  with  posterior  margin  evenly  rounded;  bases  of  pereopods  without  dark 
chromatophore  band    C.  rostrata 

3.  Frontal  process  notched  (concave);  coxal  plates  hidden  in  dorsal  aspect;  pleo- 
telson  posterior  margin  subacute;  eyes  large;  maxilla  2  trilobate   C  findleyi 

-  Frontal  process  not  notched,  but  convex;  coxal  plates  visible  on  pereonites  IV- 
VII.  in  dorsal  aspect;  pleotelson  posterior  margin  evenly  rounded;  eyes  small; 
maxilla  2  bilobate C  wallersteini  new  species 

Colidotea  wallersteini  new  species 
Figures  4,  5 

7y/7C'^.  — Holotype:  male,  AHF452.  Paratypes:  AHF4925;  SDNHM  (1  specimen); 
USNM  (1  specimen). 

Locality.  — Known  from  only  two  localities.  Holotype:  Mexico.  Baja  California 
Norte  (Pacific  coast),  Punta  Clara,  south  Rio  Santo  Tomas,  1 8  December  1 945.  collected 
by  C.  Hubbs  (H45-217a).  Paratypes:  Mexico.  Guadalupe  Island,  off' N  coast  of  Baja 
California.  Melpomene  Cove,  intertidal  collections.  18  December  1949.  collected  by 
R.  J.  Menzies  and  D.  Reish,  I  'elero  III  (Allan  Hancock  Foundation),  Station  No.  1915- 
49. 

Diagnosis.  —  Body  straight-sided;  coxal  plates  visible  in  dorsal  aspect  on  perconites 
IV-VII.  Cephalon  with  supra-antennal  line  forming  3  points.  1  medial  and  2  submedial; 
frontal  process  distally  convex;  antenna  1  flagellum  a  single  article;  antenna  2  flagcllum 
with  about  9  articles;  lacinia  mobili  present  on  both  right  and  left  mandible;  molar 


75 


K-:-- 


\-/ .;■  ■  . 


FiGi'RF  4.     Coliciotca  wallcrsietiii  new  species.  H(>loi\pc.  AHF  452.  Male. 


process  simple,  greatly  reduced  (smaller  than  lacinia);  maxilliped  with  2  coupling  hooks 
on  right.  1  on  left:  maxilla  2  bilobate.  smaller  lobe  with  2-jointed  apical  process  ter- 
minating in  single  large  seta;  posterior  margin  of  pleotelson  evenly  convex. 

/)«c'/7/J//V'//.  — Bod\  smooth,  straight-sided,  with  coxal  plates  on  pereonites  II-VII 


76 


FiciURE  5.     Colidotca  wallcrsieini  new  species.  Holotypc.  AHF  452.  Male.  a.  right  maxilliped.  b.  maxilla 
c,  maxilla  2.  d,  right  mandible,  e,  uropod.  f,  pleopod  1.  g,  pleopod  2.  h,  pcrcopod  IV.  i.  pcrcopod  I. 


SYNISOMA 


RO ST RATA 


FINDLEYI 


77 


C.  WALLERSTEINI 


yr 


rzj  12 


ZJ  11 


Figure  6.  Cladistic  relationships  of  the  4  known  species  of  Colidotea  and  their  sister-group.  Synisoma.  See 
Brusca  (in  press)  for  character  polarity  analysis  and  discussion  of  relationships  outside  this  clade.  Apomorphies 
indicated  are  as  follow:  1,  Reduction  from  a  5-jointed  to  a  4-jointed  maxillipedal  palp.  2.  Lacinia  mobilis 
of  mandible  enlarged,  subequal  to  incisor  process.  3.  Pleon  reduced  to  a  single  piece,  with  a  pair  of  lateral 
incisions  indicating  partial  fusion  of  1  pleomere.  4,  Pleon  reduced  to  a  single  piece,  with  no  remaining  lateral 
incisions.  5.  Pleon  greatly  elongated.  '/?  or  more  total  body  length.  6.  Cephalon  with  an  anteromedial  process. 
7.  Anteromedial  process  of  cephalon  bifid.  8.  Adoption  of  a  symbiotic  relationship  with  sea  urchins  of  the 
genus  Strongylocenlrotiis.  and  associated  morphological  adaptations  (e.g..  elliptical  body;  purple  pigmenta- 
tion). 9.  Eyes  enlarged.  10,  Posterior  margin  of  pleon  acute.  11.  Second  maxillae  with  only  2  lobes.  12, 
Typical  "spines"  of  first  maxilla  outer  lobe  reduced  to  3  slender,  stout,  simple  setae. 


(visible  in  dorsal  aspect  only  on  IV-VII).  Length  to  at  least  16.5  mm;  length  4-5  times 
width. 

Cephalon:  Without  tubercles  or  elevations  of  any  kind;  broadly  immersed  in  per- 
eonite  I;  frontal  process  distally  convex.  Eyes  very  small,  set  on  extreme  lateral  margins. 
Antenna  1  comprised  of  4  articles,  fourth  being  the  single  flagellar  article,  with  terminal 
esthetascs.  Antenna  2  with  9-articulate  flagellum.  Mandible  with  4-toothed  incisor; 
lacinia  with  3  teeth  and  3  stout  setae;  molar  process  greatly  reduced,  simple.  Maxilla 

1  outer  lobe  with  about  9  small  apical  setae  (some  of  which  arc  comb  setae)  and  3  stout 
simple  setae;  inner  lobe  ending  in  single  stout  setose  spine.  Maxilla  2  bilobatc;  larger 
lobe  with  2  very  large  plumose  setae  and  about  7  smaller  setae,  some  ctenose;  smaller 
lobe  with  2-jointed  apical  process  terminating  in  single  large  seta.  Maxilliped  typical 
for  genus;  enditc  with  several  large  apical  setae;  2  coupling  hooks  on  right.  1  on  left. 

Pereon;  Smooth,  straight-sided,  forming  continuous  line  with  pleon;  pereonite  I 
with  anterolateral  angles  produced  to  level  of  eyes,  engulhng  cephalon.  Pereonites 
gradually  increasing  in  width  posteriorly;  II-VI  subequal  in  length;  I  and  VII  somewhat 
shorter  than  others.  Well-developed  coxal  plates  present  on  II-VII.  visible  in  dorsal 
aspect  only  on  IV-VII.  Pcrcopods  I-VII  slender  and  ambulatory,  terminating  in  re- 
flexed,  biungulate  dactyl;  sctation  simple;  ischium  of  percopod  1  groo\  cd  to  receive 
basis. 

Pleon;  Posterior  margin  of  pleotelson  evenly  convex.  Uropods  simple;  with  single 
large  plumose  seta  at  junction  of  protopod  and  lamella.  Pleopods  typical  of  genus;  1- 

2  with  abundant  plumose  marginal  setae;  3-5  reduced,  somewhat  fleshy,  and  cither 


78 


35° 


Fici  RE  7.  Distributions  (locality  records)  of  the  known  species  of  Colidotea.  Open  circle  =  C.  wallcislcim. 
Closed  circle  =  C.  findleyi.  Triangles  =  C  rostrata.  C.  edmondsoni  is  endemic  to  the  Hawaiian  Islands  (not 
on  map). 


naked  or  at  most  with  a  few  scattered  simple  setae  (as  in  C.  findleyi).  Appendix  mas- 
culina  large,  extended  beyond  apex  of  pleopodal  lamellae,  with  short  apical  spines;  base 
of  pleopod  2  endopod  "■muscularized"  just  below  junction  of  appendix  maxulinum. 

Etymology.  — Colidotea  wallersteini  is  named  in  honor  of  Dr.  Barry  Wallerstein, 
in  acknowledgment  of  his  work  on  the  systematics  and  ecology  of  the  Idoteidae  and 


79 


in  recognition  of  his  long-standing  friendship:  furthermore,  the  resemblance  in  form 
is  striking. 

Discussion.— Oi' the  4  known  species  of  Colidolea.  2  {C.  roslrata  and  C.  edmond- 
soni)  share  a  unique  derived  character,  the  large  cephalic  tubercle.  In  addition  to  this 
synapomorphy,  they  have  more  general  characters  in  common  with  one  another  than 
either  shares  with  C.  findlcyi  or  C.  nul/crstcini.  No  synapomorphies  are  recognizable 
to  unite  the  latter  2  taxa  as  sister-species,  so  their  cladistic  relationship  must  be  viewed 
as  a  trichotomy  at  this  time.  The  character  relationships  of  the  known  species  of 
Colidotea  are  expressed  in  the  cladogram  in  Fig.  6. 

Because  Hawaii  arose  as  an  oceanic  island.  C.  cdniondsoni  most  likely  evolved 
there  in  isolation  subsequent  to  a  dispersal  event  from  western  North  America.  Its 
immediate  ancestor  was  probably  a  free-living  species  that  was  also  the  parent  taxon 
to  the  commensal  C.  rostraia.  Colidotea  findleyi  and  C.  wallersteini  can  be  derived 
from  this  same  ancestral  lineage.  The  latter  is  strictly  a  warm-temperate  species,  known 
only  from  northwestern  Baja  California  and  Guadalupe  Island  (Fig.  7).  Colidotea  fin- 
dlcyi is  also  a  warm-temperate  species,  but  exhibits  considerable  eurythermy  in  its 
ability  to  survive  the  warm  summer  months  typical  of  the  northern  Gulf  of  California. 
The  disjunct  occurrence  o{  C.  findleyi  on  both  the  western  coast  of  Baja  California  (and 
Guadalupe  Island),  as  well  as  the  restricted  northern  Gulf  of  California  population, 
suggests  that  it  may  have  entered  the  Gulf  during  a  period  of  lowered  oceanic  paleo- 
isotherms  (i.e..  Pleistocene  glacial  periods).  Brusca  and  Wallerstein  (1979b)  and  Wal- 
lerstein  and  Brusca  (1982)  have  discussed  the  probability  of  such  events,  suggesting 
that  the  Pleistocene  glacials  and  interglacials  (and  the  associated  latitudinal  shifts  in 
coastal  isotherms)  resulted  in  a  series  of  vicariant  events  such  that  populations  of 
temperate  species  were  repeatedly  trapped  in  the  northern  Gulf  of  California  as  lati- 
tudinal isotherms  shifted  from  north  to  south  and  back.  These  events  were  probably 
responsible  for  the  evolution  of  the  various  northern  Gulf  endemic  species  that  have 
warm-temperate  (Californian)  sister-taxa.  If  this  theory  is  correct,  C  findleyi  can  most 
simply  be  envisioned  as  a  sister-species  of  C  wallersteini,  the  2  taxa  being  vicariant 
products  of  one  of  the  earlier  Pleistocene  glacial  episodes.  Sympatry  was  later  effected 
when  a  subsequent  glacial  event  facilitated  the  escape  of  C  findleyi  from  the  Gulf.  The 
present  interglacial  period  and  relatively  high  coastal  isotherms  maintain  the  current 
disjunct  distribution  of  CJindleyi. 

Acknowledgments 

This  study  was  funded  by  a  grant  from  the  National  Science  Foundation  (DEB 
80-17835).  Appreciation  is  extended  to  Bill  Cooke,  George  Schultz.  and  Tom  Bowman 
for  the  loan  of  specimens  of  Colidotea  cdniondsoni:  to  the  British  Museum  (Natural 
History)  for  the  loan  of  Synisoina  specimens:  and  to  Fred  Schram  and  Greg  Pregill  for 
reviewing  the  manuscript.  Frances  Runyan  executed  the  two  dorsal  views. 

Literature  Cited 


Brusca.  R.  C.  In  press.  Phylogeny,  evolution  and 
biogeography  of  the  marine  isopod  subfamily 
Idoleinae  (Crustacea:  Isopoda:  Idoteidac).  San 
Diego  Soc.  Nat.  Hist.  Trans. 

.  and  B.  R.  Wallerstein.    1979a.   The  marine 

isopod  crustaceans  of  the  Gulf  of  California. 
II.  Idoteidae.  New  genus,  new  species,  new  rec- 
ords and  comments  on  the  morphology,  tax- 
onomy and  evolution  within  the  family.  Proc. 
Biol.  Soc.  Wash.  92(2):253-271. 

1979b.   Zoogeographic  patterns  of  idoteid 


isopods  in  the  northeast  Pacihc.  with  a  review 
of  shallow-water  zoogeography  for  the  region. 
Bull.  Biol.  Soc.  Wash.  3:67-105. 
Iverson,  E.  W.     1972.    A  new  subtidal  Synidolea 


from  central  California  (Crustacea:  Isopoda). 
Proc.  Biol.  Soc.  Wash.  85(47):54 1-548. 

Menzies.  R.  J.  1950.  The  ta.xonomy.  ecology  and 
distribution  of  northern  California  isopods  of 
the  genus  Idothca  with  the  description  of  a  new 
species.  Wasmann  .1.  Biol.  8(2):  1  55-195. 

,  and  M.  A.  Miller.    1972.    Systematics  and 

zoogeography  of  the  genus  Synic/owu  (Crus- 
tacea: Isopoda)  with  an  account  of  Californian 
species.  Smithson.  Contrib.  Zool.,  No.  102:1- 
33. 

Wallerstein.  B.  R..  and  R.  C.  Brusca.  1982.  Fish 
predation:  a  preliminary  study  of  its  role  in  the 
zoogeography  and  evolution  of  shallow-water 
idoteid  isopods  (Crustacea:  Isopoda:  Idotei- 
dae). J.  Biogeogr.  9:135-150. 


TRANSACTIONS 
OF  THE  SAN  DIEGO 
SOCIETY  OF 
NATURAL  HISTORY 


Volume  20  Number  5  pp.  81-94         18  January  1984 


Descriptions  of  five  new  muricacean  gastropods  and  comments  -  - 

on  two  additional  species,  in  the  Families  Muricidae  LIBRARY 

and  Coralliophilidae:  (Mollusca)  rpf^  "1  4  ',Gp/j 

Anthony  D'Attilio  and  Barbara  W.  Myers  HARVARD 

Department  of  Marine  Invertebrates.  San  Diego  Natural  History  Museum,  San  Diego,  CA  92112  t/S'/^TY 

Abstract.  Five  new  species  of  muricacean  (superfamily  Muricacea)  gastropods,  one  muricid  and 
four  coraliiophilids,  are  described  as  follows:  from  the  Persian  Gulf  Hexaplex  rileyi  (Muricidae:  Murici- 
nae);  from  Isla  Isabella  (= Albemarle  Island),  Galapagos  Islands,  Babelomurex  deroyorum  (Corallio- 
philidae); from  Cebu  Island,  Philippine  Islands,  Babelomurex  jeanneae,  Coralliophila  armeniaca  and 
Coralliophila  caroleae  (Coralliophilidae).  Additional  comments  and  illustrations  are  provided  on  pro- 
toconch  and  opercular  characters  of  Coralliophila  neritoidea  (Lamarck  1816),  the  type  of  Coralliophila. 
and  for  Coralliophila  erosa  (Roding  1798).  The  latter  two  species  occur  intertidally  and  subtidally  on 
coral  reefs  in  the  tropical  Indo-Pacific  Ocean. 

Introduction 

Within  the  Muricidae,  the  genus  Hexaplex  Perry,  1811,  includes  those  species  with 
a  solid  and  globose  shell  bearing  five  to  eight  spinose  more  or  less  foliaceus  varices. 
The  type  species,  Hexaplex  cichoreum  (Gmelin  1791)  occurs  in  the  Philippine  Islands. 
Specimens  from  the  Persian  Gulf,  heretofore  unknown,  are  referable  to  this  genus  as 
a  new  species,  which  we  describe  in  this  report.  We  compare  this  new  taxa  to  related 
species  from  the  Mediterranean  Sea,  Indian  Ocean  and  western  Atlantic  Ocean. 

Four  new  species  of  Coralliophilidae  are  also  described  here.  The  Coralliophilidae 
is  separable  from  all  other  muricacean  families  in  lacking  a  radula  (Thiele  1929,  Rob- 
ertson 1970).  The  members  of  this  family  feed  suctorially  and  most  are  recognized  as 
ectoparasites  or  predators  on  various  species  of  Cnidaria.  These  mollusks  move  from 
host  to  host,  though  they  are  not  known  to  be  highly  destructive  to  their  prey  (Ward 
1 965,  Robertson  1 970,  1 980).  They  have  a  thaid  type  operculum  with  a  lateral  nucleus. 
Unfortunately  shell  characters  for  this  family  are  unstable  taxonomic  criteria  and  the 
relationships  within  the  family  are  complicated  further  by  the  lack  of  a  radula.  The 
radula  is  often  a  distinctive  character  in  the  classification  of  other  muricacean  taxa  at 
the  generic  level.  Twenty-seven  genera  of  Coralliophilidae  were  enumerated  by  D'At- 
tilio  (1978)  and  four  more  have  since  been  added  (D'Attilio  1979,  Kosuge  1979), 
bringing  the  total  to  31.  The  two  genera  considered  herein  are  Coralliophila  H.  and  A. 
Adams,  1853,  and  Babelomurex  Coen,  1922.  In  Coralliophila  a  spinose  spiral  keel  is 
wanting  and  there  is  little  sign  of  episodic  growth.  Rather  growth  takes  place  in  con- 
tinuous increments.  Species  assigned  to  Babelomurex  possess  a  spinose  shoulder  keel 
and  may  have  additional  spinose  keels  on  the  body  whorl.  The  leading  edge  of  the 
spines  usually  indicates  growth  in  an  episodic  manner  similar  to  varices  in  the  Murici- 
dae. 

In  addition  to  the  five  new  species,  two  others  are  discussed:  Coralliophila  erosa 
(Roding  1798)  and  C.  neritoidea  (Lamarck  1816).  These  two  species  are  widespread 
in  the  Indo-Pacific.  where  they  inhabit  intcrtidal  and  subtidal  coral  reefs.  Illustrations 
of  certain  morphological  details  for  these  two  species  have  not  been  published  previ- 
ously. 


82 


Figures  1-4.     Hexaplex  riteyi  D'Attilio  &  Myers.  Dorsal  (1)  and  apertural  (2)  views  of  holotype,  SDNHM 
81618.  Dorsal  (3)  and  apertural  (4)  views  of  paratype,  SDNHM  81617a. 


The  following  abbreviations  are  used:  SDNHM  (San  Diego  Natural  History  Mu- 
seum); AMNH  (American  Museum  of  Natural  History,  New  York);  and  USNM  (Na- 
tional Museum  of  Natural  History,  Washington,  D.C.). 


Systematic  Account 

Phylum  MoUusca 

Superfamily  Muricacea 

Family  Muricidae  Rafinesque,  1815 

Subfamily  Muricinae  Rafinesque,  1815 

Genus  Hexaplex  Perry,  1811 

Type  species.  — Hexaplex  fo/iacea  Perry,  1811  by  subsequent  designation  Iredale 
1915  {=Mwex  cichoreum  Gmelin,  1791). 

Hexaplex  rileyi  new  species 
Figures  1-6 

[Note:  Biggs  ( 1 973:pl.  5,  figs.  8,  9)  figured  a  specimen  of  Hexap/ex  rileyi,  but  erroneously 
referred  it  to  Murex  kUsterianus  (Tapparone-Canefri  1875).] 


83 


8 


Figures  5  and  6.     Detail  drawing  of  the  protoconch  of  Hexaplex  nlcyi  from  SDNHM  81617b. 

FicH!RES  7  AND  8.     Detail  drawing  of  the  protoconch  of  Hexaplex  fulvcscens  (Sowerby  1834).  SDNHM 

81620. 

Fica  RE  9.     Detail  drawing  of  the  protoconch  of  Hcxaplex  kusterianus  (Tapparone-Canefri  1875).  SDNHM 

78106. 


Description.— '^\\e\\  broadly  biconic;  grayish  white  with  three  pale  brown  bands 
distributed  evenly  over  body  whorl,  bands  mostly  distinguishable  within  outer  aperture. 
Protoconch  of  holotype  eroded.  Six  weakly  sloping,  tabulate  postnuclear  whorls;  suture 
impressed.  Aperture  ovate,  comparatively  large;  outer  lip  crenulate,  reflecting  external 
sculpture;  inner  lip  white,  mostly  appressed;  anal  sulcus  well-defined.  U-shaped.  Si- 
phonal  canal  open,  of  moderate  length,  bent  to  left  and  recurved,  with  five  previous 
canal  terminations;  umbilical  chink  present.  Axial  sculpture  of  nine  strong,  spinose 
varices  on  body  whorl  which  cross  shoulder  and  terminate  at  suture.  Five  lamellose 
major  cords  terminate  in  open  spines  on  each  varix;  between  major  cords  are  numerous 
minor  cords  of  varying  width.  Entire  shell  including  shoulder  finely  scabrous.  A  prom- 
inent single  row  of  open  spines  on  siphonal  canal  with  space  separating  this  row  of 
spines  from  those  on  body  whorl. 


84 


Co/or. —  Grayish  white  with  three  weak  brown  bands. 

Tvpe  material. -Ho\oXyx)Q:  SDNHM  81618.  Paratypes:  SDNHM  81617a  and 
81617b;  AMNH  213801;  USNM  819632. 

Other  material  examined.  — TweWe  specimens  from  the  type  locality,  Kenneth 
Riley  collection. 

Type  locality.  — 64  km  offshore.  United  Arab  Emirates  in  The  Gulf  (Persian);  depth 
15  m;  clinging  to  petroleum  field  rigs. 

Dimensions  (in  mm).— 


Length 


Width 


51.0 

36.8 

47.0 

36.5 

21.0 

15.8 

39.0 

29.5 

33.0 

25.0 

m 

26.5 

19.5 

to 

43.9 

31.4 

SDNHM  81618  (holotype) 
SDNHM  81617a  (paratype) 
SDNHM  81617b  (paratype) 
USNM  819632  (paratype) 
AMNH  213801  (paratype) 
12  uncatalogued  specimens 
in  the  K.  Riley  coll. 

Etymology —^SiTned  for  Mr.  Kenneth  Riley,  a  petroleum  engineer  who  collected 
the  specimens. 

Discussion.  — The  color  of  this  species  ranges  from  pure  white  to  pale  brownish 
white  with  indistinct  brown  bands.  The  varices  number  from  six  to  nine  and  the 
postnuclear  whorls  from  five  to  six.  Protoconch  of  paratype  SDNHM  81617b  has  two 
and  one-third  smooth  whorls,  somewhat  tabulate  and  of  nearly  equal  diameter. 

This  new  species  is  similar  to  Hexaplex  trunculus  (Linne  1758)  from  the  Medi- 
terranean Sea.  Hexaplex  trunculus,  however,  has  a  higher  spire,  is  less  tabulate,  the 
body  whorl  is  more  convex  and  bulges  at  the  shoulder,  and  the  aperture  is  larger. 

Further  comparison  can  be  made  with  Hexaplex  fulvescens  (Sowerby  1 834),  which 
occurs  in  the  western  Atlantic  and  the  Gulf  of  Mexico,  and  H.  kiisterianus  (Tapparone- 
Canefri  1875)  from  the  Indian  Ocean.  Hexaplex  fulvescens  has  a  much  larger  shell  (to 
200  mm),  the  shoulder  is  non-tabulate,  suture  weakly  impressed,  aperture  denticulate, 
and  canal  much  longer.  In  addition,  the  varices  are  concave  on  their  receding  side,  and 
the  shell  is  white  with  reddish  brown  lines  over  the  spiral  threads  of  the  entire  shell. 
Hexaplex  fulvescens  has  a  protoconch  of  three  convex  whorls  which  show  very  weak 
axial  grooves  (Figs.  7,  8).  Hexaplex  kiisterianus  has  a  broad  shell  with  a  low  spire  and 
indistinct  suture.  The  body  whorl  is  very  broad  and  the  aperture  large.  The  protoconch 
of  H.  kiisterianus  has  an  indistinct  number  of  convex  whorls  (Fig.  9). 


Family  Coralliophilidae  Chenu,  1859 
Genus  Babelomurex  Coen,  1922 

Type  species.  — Fusus  babelis  Requien,  1848  by  original  designation. 


Babelomurex  deroyorum  new  species 
Figures  10-14 

Description.  S\\e\\  broadly  fusiform;  spire  moderately  high;  suture  weakly  im- 
pressed. Protoconch  of  holotype  eroded;  teleoconch  of  six  whorls;  aperture  large,  ovate; 
outer  lip  sharp,  with  spiral  grooves  resulting  from  open  spines  at  margin.  Anal  sulcus 
broad,  shallow;  inner  lip  demarked  by  elongate  node;  canal  short,  open,  recurved; 
siphonal  fasciole  with  chink  and  five  short  canal  terminations.  Axial  sculpture  often 
varices  with  weakly  defined  margins  on  body  whorl;  penultimate  body  whorl  with  nine 
varices;  strong  shoulder  keel  separated  by  gap  from  two  close-set  keels  below.  Narrow, 
flatly  triangular,  relatively  long  spines  arise  at  varical  margins;  spines  on  keel  directly 
below  shorter,  and  those  on  most  anterior  keel  progressively  shorter;  presence  of  ter- 
minal portions  of  secondary  keel  obscuring  suture;  fine  squamous  threads  covering 
entire  surface,  including  upper  and  lower  surface  of  spines;  spiral  threads  coarser  on 
canal. 


85 


Figures  10-13.     Babelomurex  deroyorum  D'Attilio  &  Myers.  Dorsal  (10)  and  apertural  (11)  views  of  ho- 
lotype.  SDNHM  81613.  Dorsal  (12)  and  apertural  (13)  views  of  paratype,  SDNHM  81616a. 


Co/or.  — White,  showing  some  sHght  attrition;  small  patches  are  covered  with  a  red 
hydrocoral  or  bryozoan;  paratypes  with  a  lustrous  white  aperture. 

Type  material.  - Holotype:  SDNHM  81613.  Paratypes:  SDNHM  8 1 6 1 6a.  8 1 6 1 6b, 
and  8 16 16c;  USNM  819633a  and  819633b;  AMNH  213802a  and  213802b. 

Type  locality.  — Y:>vQdgQd  from  75-100  m  off  Isla  Isabella  (Albemarle  Island).  Tagus 
Cove.  Galapagos  Islands;  January  1969. 

Dimensions  (in  mm).— 

Length  Width 

SDNHM  81613  (holotype)  28.8  29.3 

SDNHM  81616a  (paratype)  17.0  12.5 

SDNHM  81616b  (paratype)  12.2  8.9 

SDNHM  81616c  (paratype)  8.4  7.4 

AMNH  213802a  (paratype)  13.8  11.0 

AMNH  213802b  (paratype)  9.8  9.2 

USNM  819633a  (paratype)  12.7  9.0 

USNM  819633b  (paratype)  8.0  5.5 

Etymology.  — ^dixned  for  the  collectors,  Mr.  and  Mrs.  Andre  DeRoy.  who  through 
their  collecting  efforts,  have  contributed  to  our  understanding  of  the  endemic  molluscan 
fauna  of  the  Galapagos  Islands. 


86 


Figure  14.     Detail  drawing  of  the  protoconch  of  Bahelonturcx  dcroyorum  from  SDNHM  81616c. 


Discussion.  — In  addition  to  the  holotype,  which  is  a  dead  empty  shell  showing 
some  attrition,  seven  smaller  specimens  were  studied,  the  largest  of  which  possesses 
five  whorls.  The  keel,  showing  spines,  appears  clearly  on  the  third  postnuclear  whorl. 
Protoconch  of  SDNHM  81616c  has  three  and  one-half  whorls  with  a  spiral  cord  in  the 
center  and  a  second  spiral  cord  below;  axial  threads  are  diagonal  to  the  protoconch, 
knob-like  across  the  spiral  cords.  This  new  species  differs  from  all  other  eastern  Pacific 
and  western  Atlantic  species  in  the  narrow,  elongate  spines,  and  finer  squamous  sculp- 
ture. This  species  was  recovered  from  the  same  area  as  Babc/oniurc.x  santacruzensis 
(Emerson  and  D'Attilio  1970)  which  has  a  single  row  of  spines  on  the  shoulder  and  is 
more  closely  related  to  B.  dalli  (Emerson  and  D'Attilio  1 963)  from  the  western  Atlantic. 
Other  eastern  Pacific  species  are  B.  oldroydi  (Oldroyd  1929),  B.  costata  (Blainville 
1832),  and  B.  hindsi  (Carpenter  1857).  Babeloiuwex  oldroydi  is  found  off 
the  California  coast  and  has  a  larger,  heavier,  coarser  shell  than  B.  deroyoruni  and 
possesses  three  spinose  keels.  Babelonuire.x'  costata  is  similar  to  B.  oldroydi,  but  with 
less  developed  and  variably  keeled  spiny  cords;  B.  hindsi  is  smaller  than  any  of  these 
species. 


Babeloniurex  jeanneae  new  species 
Figures  1 5-20 

Description. —  S\\q\\  biconically  fusiform;  height  from  top  of  aperture  to  canal  ter- 
mination, 1 0  mm.  Protoconch  of  holotype  eroded;  six  postnuclear  angulate  whorls  with 
large  spinose  keel  at  shoulder  angle;  suture  obscured  by  anteriorly  directed,  scabrous- 
edged  spinose  keel  corresponding  to  secondary  keel  on  body  whorl;  aperture  relatively 
large,  ovate;  columella  pillar  straight  except  for  slight  concavity  midway;  inner  lip  edge 
weakly  erect  anteriorly,  outer  lip  (not  entirely  mature)  extending  into  the  open  spines; 
siphonal  canal  short,  broad,  open,  recurved;  fasciole  strongly  sculptured,  with  four 
older  canal  terminations;  umbilical  chink  narrow  but  deep;  seven  varices  on  body 
whorl,  eight  on  penultimate  whorl;  all  varices  terminating  in  lengthy  spines.  Spiral 
sculpture  of  a  primary  row  of  broad,  contiguous,  open  spines  forming  the  keel;  growth 
of  spines  episodic,  giving  them  a  scaly  or  foliated  appearance.  Outer  portion  of  primary 
spine  bent  in  direction  of  growth  and  entire  spiny  keel  recurved  posteriorly;  second 
and  third  row  of  spines  below;  bases  of  secondary  spines  contiguous  and  forming  keels. 
Five  progressively  smaller  scabrous  rows  of  spines  on  canal;  one  similar  scaly  row 
between  second  and  third  row  of  spines;  six  rows  of  weak  scabrous  spiral  cords  above 
shoulder  terminating  at  apertural  margin. 


87 


Figures  15  and  16.     Babclomurex  jeanneae  D'Attilio  &  Myers.  Dorsal  (15)  and  apertural  (16)  views  of 
hoiotype.  SDNHM  79499. 


CoA)/-.  — Creamy  white,  stained  with  pale  ochre  in  depressed  part  of  shoulder  and 
body  at  receding  side  of  margin. 

'Type  material. -Ho\o\\vq:  SDNHM  79499.  Paratypes:  SDNHM  79500  and  81402. 

Other  ntaterial  exatuined. —  OnQ  specimen  in  the  Rose  D'Attilio  collection,  one 
specimen  in  the  Barbara  W.  Myers  collection  and  one  specimen  in  the  Donald  Pisor 
collection. 

Type  loeality.  —  BohoX  Straits  between  the  Islands  of  Cebu  and  Bohol  in  the  Phil- 
ippine Islands. 

DiiJiefisions  (in  »ini).— 


.ength 

Width 

21.5 

25.5 

22.9 

25.9 

23.5 

22.0 

17.0 

29.0 

23.0 

25.0 

22.0 

24.5 

SDNHM  79499  (hoiotype) 
SDNHM  81402  (paratype) 
SDNHM  79500  (paratype) 
R.  D'Attilio  coll. 
B.  W.  Myers  coll. 
D.  Pisor  coll. 

Etyiuology.— Named  for  Jeanne  Pisor.  who  with  her  husband  Donald  Pisor.  have 
made  noteworthy  contributions  to  the  molluscan  collections  of  the  San  Diego  Natural 
History  Museum. 

Discussion.  — There  may  be  one  or  two  spiny  cords  below  the  shoulder  keel:  the 
number  of  cords  below  the  secondary  keel  varies  from  four  to  six:  growth  striae  may 
be  strong.  SDNHM  79500,  with  a  mature  outer  lip.  retains  a  while  intritacalx.  a  chalky 
white  surface  layer  in  some  mollusks  (D'Attilio  and  Radwin  1971).  with  no  color  other 
than  the  white  surface.  The  specimen  in  the  R.  D'Attilio  collection  has  unusually  broad, 
lengthy  spines.  The  protoconch  on  this  specimen  has  two  and  one-half  whorls:  the 
earliest  portion  is  smooth,  followed  by  a  sculptured  portion  with  fi\e  close-set  a.xial 
striae  and  two  spiral  cords,  beaded  where  crossed  by  striae.  The  specimen  in  the  Barbara 
W.  Myers  collection  is  white  flushed  with  a  pale  violet-pink,  and  the  spines  curve 
strongly  in  the  direction  of  growth. 

Babelomure.x  jeanneae  probably  belongs  to  a  complex  of  species,  the  best  known 
of  which  is  B.  pagodus  [of  authors,  not  B.  pagodus  (A.  Adams  1853)  (see  D'Attilio 
1983)].  Characters  similar  to  B.  pagodus  are  the  possession  of  a  spinose  keel  at  the 
shoulder  and  a  secondary  keel  midway  on  the  body  whorl  followed  anteriorly  by  a 


88 


17 


Figure  1 7.     Detail  drawing  of  the  protoconch  of  Babelomurexjeanneae  from  specimen  m  the  Rose  D'Attiho 

collection. 

Figures  18  and  19.     Babelomurex  jeanneae.  Specimen  from  the  Rose  D'Attilio  collection.  Detail  of  spine 

formation  showing  scales  or  foliations  (18).  Detail  of  broad  spines  as  viewed  from  above  (19). 

FiGLiRE  20.     Babelomurex  jeanneae.  Detail  showing  spines  curved  in  the  direction  of  growth  from  specimen 

in  the  B.  W.  Myers  collection. 


series  of  progressively  diminishing  spinose  cords.  Babelomurex  pagodus  also  has  a 
white  shell  shaded  with  ochre  in  the  concave  portions  of  the  intervarical  areas.  Varical 
spinose  projections  also  number  around  eight  and  are  sharply  triangulate.  Babelomurex 
pagodus  is  well  known  and  common  in  southeastern  Japan.  Based  on  literature  records 
its  geographic  range  is  extensive,  being  known  throughout  the  central  and  western 
Pacific.  As  far  as  is  known  from  the  extensive  collecting  done  in  Japan,  B.  jeanneae 
does  not  occur  there.  Cebu  specimens  of  B.  pagodus  resemble  in  shell  morphology 
specimens  from  southeastern  Japan. 

Babelomurex  fruiticosus  (Kosuge  1979),  described  from  specimens  obtained  in 
the  Straits  of  Bohoi  between  Cebu  and  Bohol  Islands,  has  some  similarity  to  B.  jeanneae 
in  its  possession  of  a  spinose  keel  at  the  shoulder  and  one  midway  on  the  body  whorl. 
However,  the  spire  is  comparatively  higher,  the  area  below  the  shoulder  keel  shorter, 
appearing  compressed,  and  there  are  only  two  spinose  cords.  The  spines  are  narrow, 
often  very  elongate,  bent  or  recurved,  and  their  receding  side  ornamented  with  sharply 
pointed  narrow  spinelets  imparting  to  the  spines  a  resemblance  to  deer  antlers.  In 
addition,  the  entire  shell  is  suffused  with  rich  pink  or  pink-violet,  or  at  times  creamy 
white  with  red-brown  at  the  keel  concentrated  on  the  receding  side  of  the  varix. 

Babelomurex  cristatis  (Kosuge  1979)  also  has  conchological  characters  similar  to 
B.  jeanneae.  However,  the  eight  shoulder  keel  spines  in  B.  cristatus  are  shorter,  strongly 
up-turned  and  recurved,  with  spinelets  on  their  receding  side;  the  secondary  keel  on 
the  mid-area  of  the  body  whorl  is  mostly  non-spinose  except  in  mature  specimens. 
Two  or  three  inconspicuous  cords  occur  below  the  mid-cord,  and  a  nearly  obsolete 
spiral  cord  may  be  present  on  the  canal;  the  intervarical  areas  are  relatively  broad  and 
the  shell  surface  is  finely  and  scabrously  striate;  the  color  is  light  to  dark  tan  (flesh), 
occasionally  being  a  deeper  brown  in  the  concave  areas  between  the  strongly  formed, 
rounded,  axial  costae. 


89 


Figures  21  and  22.     Coralliophila  armeniaca  D'Attilio  &  Myers.  Dorsal  (21)  and  apertural  (22)  views  of 
holotype.  SDNHM  79507. 

Genus  Coralliophila  H.  and  A.  Adams.  1853 

Type  species  by  subsequent  designation  Iredale  1912:  ""Murex  neritoideus  Chem. 
[nitz]"  =  Murex  neritoideus  Gmelin,  1791,  not  Linne,  1767  =  Fusus  neritoideus  La- 
marck, 1816  (syn.  Purpura  violacea  and  P.  diversiforniis  Kiener,  1836). 

Coralliophila  armeniaca  new  species 
Figures  21-24 

Description. —  S\\q\\  fusiform  above,  compressed  anteriorly.  Protoconch  of  holotype 
eroded;  six  postnuclear  convex  whorls;  suture  wavy,  distinct;  body  whorl  large,  sharply 


Figure  23.     Detail  drawing  of  the  protoconch  of  Coralliophila  armeniaca  from  SDNHM  795041. 
Figure  24(a)  AND(b).     Detail  drawing  of  the  operculum  o^  Coralliophila  armeniaca  from  holotype.  SDNHM 
79507.  Internal  (a)  and  external  (b)  views. 
Figure  25.     Detail  drawing  of  the  protoconch  of  Coralliophila  rosacea  (Smith  1903)  from  SDNHM  72131. 


90 


FiGL'RES  26  AND  27.     Coralliophila  cawleae  D'Attilio  &  Myers.  Dorsal  (26)  and  apertural  (27)  views  of 
holotype.  SDNHM  79505. 


incurved  below,  flaring  out  at  siphonal  fasciole.  Aperture  large  with  ten  lengthy  lirae 
within;  lirae  at  the  crenulate  outer  lip  more  numerous,  reflecting  external  spiral  sculp- 
ture; inner  lip  erect  on  lower  two-thirds,  adherent  above.  Anal  sulcus  very  weak  pos- 
teriorly at  juncture  of  outer  lip  and  columella;  siphonal  canal  open,  relatively  short  and 
broad,  recurved,  with  numerous  strong  canal  terminations  on  the  flaring  fasciole,  um- 
bilical chink  present.  Shell  with  axial  sculpture  of  six  swollen,  rounded  ribs  set  close 
together,  terminating  at  base  of  body  whorl  below  and  diminishing  in  strength  as  they 
abut  the  whorl  above.  Spiral  sculpture  of  primary  and  secondary  cords  form  entire 
surface  of  the  shell;  about  15  primary  cords  with  intercalary  secondary  cords  on  the 
body  whorl  below  the  shoulder;  similar  cords  covering  the  shoulder;  all  cords  weak  to 
strongly  scabrous.  Operculum  thin,  translucent,  amber-brown  with  concentric  ridges 
externally;  internally  with  two  small  cords;  horseshoe  shaped. 

Color.  —  Deep  apricot  with  slight  orange  cast;  paratypes  range  from  violet  to  apricot- 
pink.  Aperture  pale  orange  at  edge  of  outer  lip,  white  within  and  on  the  columella. 

Type  material. -\\o\oXypQ:  SDNHM  79507.  Paratypes:  SDNHM  79504a-j;  USNM 
819634a  and  819634b;  AMNH  213803a  and  213803b. 

Type  locality.  — W\  specimens  from  approximately  75  m  depth  off^Cebu  Island,  in 
the  Bohol  Straits,  Philippine  Islands;  obtained  with  ground  nets. 

Ditnensions  (in  mm).— 


SDNHM  79507  (holotype) 
SDNHM  79504a  (paratype) 
SDNHM  79504b  (paratype) 
SDNHM  79504c  (paratype) 
SDNHM  79504d  (paratype) 
SDNHM  79504c  (paratype) 
SDNHM  79504f  (paratype) 
SDNHM  79504g  (paratype) 
SDNHM  79504h  (paratype) 
SDNHM  79504i  (paratype) 
SDNHM  79504J  (paratype) 
AMNH  213803a  (paratype) 
AMNH  213803b  (paratype) 
USNM  819634a  (paratype) 
USNM  819634b  (paratype) 


Length 

Width 

20.5 

11.9 

20.0 

10.2 

18.5 

10.5 

17.2 

8.9 

12.5 

6.4 

13.9 

7.3 

12.8 

6.8 

12.1 

6.2 

10.0 

5.0 

9.0 

5.0 

9.8 

5.0 

13.5 

7.0 

8.0 

4.5 

15.0 

8.0 

7.0 

4.0 

91 


Figure  28.     Detail  drawing  of  the  protoconch  of  Coral/ioplula  caraleac  from  hoiotype.  SDNHM  79505. 
FiciRE  29.     Coral/iop/u/a  ncniouica  (Lamarck  1816).  Apertural  view  of  a  juvenile  SDNHM  66538. 
Pick  RF  30.     Detail  drawing  of  protoconch  of  C  neritoidea  from  SDNHM  66538. 

Fkh  RE  31  (a)  AN[5  (b).     Detail  drawing  of  operculum  of  C.  iwriloidca  from  SDNHM  66538.  Internal  (a) 
and  external  (b)  views. 


Etymology.  — ¥vom  arnieniacus,  referring  to  the  color  of  ripe  apricot  fruit. 

Discussion.— This  species  appears  closely  allied  to  Coralliophila  fritschi  (von  Mar- 
tens 1874)  and  C.  rosacea  (Smith  1903),  both  from  South  Africa.  Another  apparently 
closely  related  species  is  C.  arbutum  (Woolacott  1954)  {  =  Rhombothais  arhiiiiini  Woo- 
lacott  1954)  from  New  South  Wales,  Australia.  These  three  species  differ  from  C. 
anncniaca  by  their  possession  of  broadly  fusiform  shells  with  a  larger  body  whorl, 
widest  at  mid-height.  In  contrast,  C.  atincniaca  has  a  relatively  high  spire  and  the  shell 
is  broadest  well  below  mid-height.  In  addition,  the  angulate  shoulder  of  C.  arbimini 
gives  it  a  biconic  shape.  Protoconch  of  SDNHM  79504i  has  three  and  one-half  whorls: 
the  first  whorl  is  smooth  and  rounded  whereas  the  remaining  whorls  have  two  spiral 
cords  crossed  by  axial  ridges  and  the  nodes  are  poorly  developed  where  the  axial  and 
spiral  sculpture  cross.  The  protoconch  of  C.  rosacea  is  illustrated  for  comparison  (Fig. 
25). 

Coralliophila  caroleae  new  species 
Figures  26-28 

Description.  — 'shcW  small,  biconic;  spire  concave,  low  to  moderate  height:  body 
whorl  swollen  with  moderately  angled  shoulder  tapering  to  a  short,  open  canal.  Pro- 
toconch of  nearly  three  whorls,  ridged  and  beaded.  Six  postnuclear  whorls,  moderately 
angled:  suture  not  clear  as  each  succeeding  whorl  encroaches  and  somewhat  submerges 
previous  whorl  up  to  body  whorl  where  suture  is  clearly  defined.  Aperture  wide,  with 
sinuous  crenulate  outer  lip:  inner  lip  smooth,  adherent  posteriorly:  canal  short  and 
open.  Siphonal  fasciole  composed  of  a  cur\ing  scabrous  ridge:  umbilical  chink  mod- 
erately deep.  Axial  sculpture  beginning  on  first  postnuclear  whorl  with  nine  ribs,  in- 
creasing to  14  on  subsequent  whorls  and  fading  at  fourth  postnuclear  whorl,  becoming 


92 


FiGL'RE  32.     Coralliophila  erosa  (Roding  1798).  Apertural  view  of  a  juvenile  SDNHM  77174. 

Figure  33.     Detail  drawing  of  the  protoconch  of  Coralliophila  erosa  from  SDNHM  77174. 

Figure  34  (a)  and  (b).     Detail  drawing  of  the  operculum  of  C.  erosa  from  SDNHM  77 1 74.  Internal  (a)  and 

external  (b)  views. 


barely  visible  on  body  whorl.  Spiral  sculpture  of  scabrous  major  and  minor  cords  above 
and  below  periphery,  with  strong  major  cord  at  periphery  of  each  whorl.  Body  whorl 
with  1 1  cords  between  suture  and  periphery,  and  28  cords  from  periphery  to  canal; 
width  of  cords  variable;  interspaces  narrow. 

Color.  — Dull  orange;  aperture  orange;  inner  lip  pale  orange. 

Type  material. -Holoxype:  SDNHM  79505.  Paratypes:  SDNHM  79503  and  81614. 

Other  material  examined.  — One  specimen  in  the  Donald  Pisor  collection  and  one 
specimen  in  the  James  Springsteen  collection. 

Type  locality.  — Boho\  Straits  between  the  Islands  of  Cebu  and  Bohol  in  the  Phil- 
ippine Islands,  dredged  with  bottom  nets  at  75-100  m. 

Dimensions  (in  mm).- 


SDNHM  79505  (holotype) 
SDNHM  79503  (paratype) 
SDNHM  81614  (paratype) 
D.  Pisor  coll. 
J.  Springsteen  coll. 

£'/>'mo/o^.  — Named  for  friend  and  co-worker,  Carole  M.  Hertz. 

Discussion.— This  new  species  is  similar  to  two  other  deep  water  species  from 
Mactan  Island,  Cebu,  Philippine  Islands:  Coralliophila  elvirae  D'Attilio  and  Emerson, 
1980,  and  C.  solutistoma  Kuroda  and  Shikama,  1966.  Although  C  soliitistoma  was 
described  from  Japan,  it  has  been  recently  discovered  in  the  Philippine  Islands  (D'At- 
tilio  and  Emerson  1980). 

The  spire  of  C.  elvirae  is  higher  and  convex  rather  than  concave  as  in  C.  caroleae. 
The  aperture  of  C  elvirae  is  restricted  to  a  comparatively  longer,  narrow  opening,  with 
the  inner  and  outer  lip  parallel,  whereas  C.  caroleae  has  a  broad  aperture  and  swollen 
body  whorl.  The  protoconch  of  C.  elvirae  consists  of  two  and  one-fourth  whorls  with 
weakly  beaded  cords;  there  is  only  one  cord  on  the  final  whorl  (D'Attilio  and  Emerson 
1980).  The  protoconch  of  C  caroleae  has  two  and  one-half  to  three  whorls,  the  beading 
on  the  cords  is  much  stronger,  and  the  final  whorl  has  two  cords.  Coralliophila  solu- 
tistoDia  has  a  heavier,  larger  shell  with  a  higher  spire  and  less  inflated  body  whorl;  the 
axial  ribbing  is  much  stronger  and  the  ribbing  is  continuous  over  the  entire  shell.  The 
protoconch  is  more  coarsely  beaded  (D'Attilio  and  Emerson  1980). 


Length 

Width 

19.2 

13.6 

14.3 

11.0 

16.8 

9.2 

15.1 

11.8 

12.5 

7.9 

93 


Notes  on  Two  Additional  Species  of  Coralliophila 

Two  well-known  species  oi  Coralliophila,  widely  distributed  throughout  the  Indo- 
Pacific,  are  discussed  below.  The  two  species  are  usually  found  in  such  eroded  and 
encrusted  condition  that  morphological  characters  are  obscured.  Hence,  an  adequate 
description  of  the  protoconch  and  early  whorls  has  been  lacking  in  the  literature.  We 
are  fortunate  in  having  at  our  disposal  (SDNHM)  juvenile  specimens  of  each,  and  we 
have  thus  appended  information  regarding  their  shell  morphology,  protoconch  and 
operculum  to  add  to  a  general  understanding  of  the  genus. 

Coralliophila  erosa  Roding,  1  798 
Figures  32-34 

This  species  is  distributed  throughout  the  Indo-Pacific,  mostly  intertidal.  Mor- 
phologically it  exhibits  considerable  inter-  and  intrapopulation  variation.  The  shell  is 
often,  as  the  name  suggests,  encrusted  or  eroded,  obscuring  the  finely  scabrous  surface. 
It  has  a  characteristic  sinuous  outer  lip,  the  upper  portion  of  which  is  deeply  concave. 
The  operculum  is  thin  with  close  concentric  ridges  externally;  internally  there  are  six 
U-shaped  ridges.  The  protoconch  is  distinguished  by  having  one  and  three-fourths  low 
and  weakly  convex  whorls;  spire  is  depressed.  An  uneroded  juvenile  specimen  (SDNHM 
77174)  from  Hawaii  is  illustrated. 

Coralliophila  neritoidea  Lamarck,  1816 
Figures  29-31 

This  species  has  a  distribution  and  habitat  similar  to  Coralliophila  erosa.  The  white 
shell  is  most  often  encrusted  with  calcareous  organisms.  When  not  encrusted  or  eroded, 
the  surface  of  the  shell  is  spirally  scabrous.  After  the  first  two  to  three  postnuclear 
whorls  there  often  follows  a  rapid  expansion  of  the  body  whorl.  The  purple  aperture 
is  less  prominent  in  immature  specimens.  The  operculum  is  a  dark  chocolate  brown 
with  a  lateral  mid-central  nucleus.  Internally  it  has  a  thickened  ridge  and  four  ovate 
ridges  spaced  over  the  remaining  surface.  The  protoconch  has  two  to  two  and  one-half 
smooth,  acutely  conical  whorls,  followed  by  a  whorl  with  a  midway  rib.  A  juvenile 
specimen  (SDNHM  66538)  is  illustrated. 

Acknowledgments 

The  following  friends  and  collectors  have  contributed  towards  the  completion  of 
this  paper:  Mr.  Kenneth  Riley,  for  donating  the  specimens  used  in  the  description  of 
Hexaplexrileyi;  Mr.  and  Mrs.  Andre  DeRoy  from  Santa  Cruz  Island,  Galapagos  Islands, 
for  donating  specimens  of  Babelomurex  deroyorunr,  Mr.  Donald  Pisor  of  San  Diego, 
California,  and  Mr.  Loyal  J.  Bibbey  of  Imperial  Beach,  California,  for  donating  spec- 
imens of  Babelomurex  Jeanneae:  Mr.  Gene  Everson  of  Lauderhill.  Florida,  and  Mr. 
Donald  Pisor  for  contributing  specimens  of  Coralliophila  armeniaca:  Mr.  Donald  Pisor 
and  Mr.  Jim  Springsteen  of  Manila,  Philippine  Islands,  for  donating  specimens  of 
Coralliophila  caroleae. 

We  are  especially  grateful  to  Mr.  David  K.  MuUiner  of  San  Diego,  California,  for 
the  photographs  used  in  this  paper.  We  further  thank  Dr.  Richard  C.  Brusca,  Chairman, 
Department  of  Marine  Invertebrates,  San  Diego  Natural  History  Museum;  Dr.  William 
K.  Emerson,  American  Museum  of  Natural  History;  and  Dr.  Emily  H.  Yokes,  Tulane 
University,  for  editorial  review  of  the  manuscript.  Marjorie  Rea  kindly  typed  the 
manuscript. 

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lima  from  the  Cumingian  collection.  Proc.  Zool.  London  24(8):343-42 1 . 

Soc.  London  for  1851   19:267-272.  BlainviUe.  H.  M.    1832.    Disposition  methodique 

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mollusca.  Van  Voorst,  London,  vol.  1.  484  pp.  Pourpre.  Ricinule.  Licorne  et  Concholepus  de 


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TRANSACTIONS 

'^\,PRARY  OF  THE  SAN  DIEGO 

^  SOCIETY  OF 

|_ ^  ,5  'mA  NATURAL  HISTORY 

UNIVERSITY 


Volume  20  Number  6  pp.  95-98  18  January  1984 


The  Fossil  Leptostracan  Rhabdouraea  bentzi  (Malzahn,  1958) 

Frederick  R.  Schram 

Department  of  Geology;   Paleontology  Section.   San  Diego  Natural  History  Museum,   San  Diego.   CA 
92112  USA 

Eric  Malzahn 

Aufder  Heide  33.  D-3004  Isernhagen,  German  Federal  Republic 

Abstract.  The  type  material  of  the  only  known  fossil  leptostracan  phyllocarid,  Rhabdouraea  bentzi, 
is  reexamined.  This  species  was  originally  placed  in  the  genus  Nebalia.  The  distinctive  caudal  rami  and 
the  form  of  the  carapace,  however,  require  separate  generic  and  familial  status  for  this  Permian  material. 
A  redefinition  of  the  living  family  Nebaliidae  is  also  provided. 

Introduction 

The  fossil  record  of  the  Phyllocarida  has  always  presented  problems  for  phylo- 
genetic  analysis  because  of  the  incomplete  nature,  generally,  of  the  fossils  (Rolfe  1981). 
This  has  been  especially  vexing  for  the  assessment  of  the  living  order  Leptostraca  where, 
with  a  single  exception,  fossils  are  non-existent.  Malzahn  (1958)  described  Nebalia 
bentzi  on  the  basis  of  a  single  specimen  from  the  Upper  Permian  Zechstein  of  West 
Germany.  Further  discussion  of  this  species  was  provided  by  Glaessner  and  Malzahn 
(1962).  Initial  assignment  of  this  material  to  the  living  genus  Nebalia  reflected  a  con- 
servative approach  pending  the  discovery  and  study  of  more  and  better  material. 
Unfortunately,  despite  continuing  efforts  at  collection  and  study  of  this  fauna  by  the 
junior  author,  no  additional  specimens  have  turned  up.  However,  restudy  of  the  known 
material  has  enabled  us  to  clarify  the  relationship  of  this  Permian  species  to  the  living 
leptostracans. 

Systematic  Paleontology 

Class  Malacostraca  Latreille,  1806 

Subclass  Phyllocarida  Packard,  1879 

Order  Leptostraca  Claus,  1880 

Family  Rhabdouraeidae  n.  fam. 

Diagnosis. —  S>?LmQ  as  that  of  the  genus. 
Type  genus.— Rhabdouraea  n.  gen. 

Rhabdouraea  new  genus 

D/fl^/7m«.— Carapace  short,  not  covering  abdomen.  Caudal  rami  rod-like,  at  least 
as  long  as  abdomen. 

Etymology.  — Yrom.  the  Greek  rhabdos  (=rod)  and  uraea  (=tail),  gender  feminine. 
Type  species.— Nebalia  bentzi  Malzahn,  1958. 

Rhabdouraea  bentzi  (Malzahn.  1958) 

Diagnosis.— Since  only  one  species  is  known,  the  diagnosis  of  the  species  is  the 
same  as  that  of  the  genus. 


96 


<> 


Figure.  1 .     Partial  reconstruction  of  Rhahdouraea  bentzi  (Malzahn).  A.)  lateral  view  of  abdomen.  B.)  dorsal 
view  of  telson  and  caudal  rami.  Scale  2  mm. 


Holotype.—Z45,  in  the  collection  of  the  Niedersachsisches  Geologisches  Lande- 
samt,  Hannover.  From  a  depth  of  280-282  m.  Shaft  4,  of  the  Friedrich-Heinrich  Mine, 
near  Ort  Hoerstgen,  Niederrhein,  German  Federal  Republic.  Zechstein  1,  Upper  Perm- 
ian. 

Description.— The  posterior  margin  of  the  carapace  is  visible  on  the  type,  and 
covers  the  posterior  aspect  of  the  thorax  while  leaving  the  anterior  pleomeres  exposed 
(Figs.  1  and  2).  The  pleomeres  are  subequal,  each  about  0.7  mm  in  length.  The  posterior 
margin  of  the  abdominal  tergites  are  raised  as  a  slight  ridge.  The  abdominal  pleura  are 
rounded  anteriorly  and  somewhat  acuminate  posteriorly.  At  least  the  four  anterior 
pleopod  protopods  are  long  and  robust;  the  fifth  and  sixth  are  present,  appear  to  be 
somewhat  smaller  than  those  of  the  anterior  limbs,  but  cannot  be  clearly  discerned 
because  of  the  preservation  of  the  specimen.  The  seventh  pleomere  appears  to  lack 
appendages.  The  distal  branches  of  the  anterior  pleopods  are  robust,  and  marked  by  a 
line  of  lateral  pits  (which  may  have  been  sockets  of  marginal  setae).  The  telson  is  only 
slightly  shorter  than  the  pleomeres,  and  with  a  slight  fossa  or  depression  on  the  dorsal 
surface  between  the  bases  of  the  caudal  rami.  The  telson  bears  terminally  a  set  of  large, 
rod-like,  papillose  caudal  rami.  The  length  of  these  rami  cannot  be  determined  exactly 
since  their  distal  ends  are  broken  off.  However,  the  longer  ramus  (now  broken  in  two 
pieces)  is  at  least  3.5  mm  and  indicates  the  rami  were  probably  at  least  as  long  as  the 
abdomen. 

Family  Nebaliidae  Baird,  1850 

Diagnosis.— Carapace  generally  large,  covering  anterior  pleomeres  as  well  as  the 
thorax.  Caudal  rami  relatively  short,  as  flaps  with  marginal  setae. 
Type  genus.— Neba/ia  Leach,  1814. 


Discussion 

The  two  distinct  features  of  Rhahdouraea  bentzi  are  the  short  carapace  and  the 
long,  rod-like  caudal  rami,  so  diagnostic  in  fact  as  to  warrant  separate  generic  and 
familial  status  for  this  species. 


97 


FiGiRF  2.  Rhabdouraea  bentzi  (Malzahn).  holotype.  Z45.  A.)  Right  side  [right  caudal  ramus  visible  on 
original  illustrations  since  broken  off.  but  still  retained  with  specimen],  lOx.  B.)  Left  side,  closeup  of  anterior 
pleopods  and  caudal  ramus,  16x;  c— carapace,  cr— caudal  ramus,  pi  — pleuron  of  second  pleomere,  pr— 
pleopodal  protopods.  r— distal  rami  of  pleopods.  t  — telson. 


The  living  leptostracans  typically  have  a  large  carapace  that  completely  encloses 
the  body,  except  for  the  posteriormost  portions  of  the  abdomen,  and  lobate.  setose 
caudal  rami.  The  only  exception  to  these  features  occurs  in  the  genus  Nehaliopsis.  This 
pelagic  form  has  the  posterolateral  aspect  of  the  carapace  truncate,  exposing  the  pleo- 
meres  and  ventral  portions  of  the  posterior  thoracomeres.  In  addition,  the  caudal  rami 
in  Nebaliopsis  are  thin  and  leaf-like.  These  features  are  prompting  Hessler  (in  prep.) 


98 


to  place  Nehaliopsis  in  its  own  family,  separate  from  the  benthic  nebaliid  genera.  The 
differences  noted  on  Rhahdouraea,  incomplete  though  the  fossil  is,  are  of  a  magnitude 
at  least  as  great  as  that  which  separates  Nehaliopsis  from  Nehalia.  Paranebalia,  and 
Neba/ic/la,  and  thus  justify  separating  this  Permian  taxon  by  itself 

All  the  living  nebaliids,  benthic  or  pelagic,  have  relatively  short  or  modest  sized 
caudal  rami  with  setose  margins.  These  rami  function  in  a  manner  analogous  to  uropods, 
and  assist  the  animal  in  swimming.  The  very  long,  papillose,  rod-like  rami  in  Rhah- 
douraea are  unlike  any  of  those  seen  among  leptostracans,  and  are  more  akin  to  those 
seen  in  notostracan  branchiopods. 

Several  features  of  Rhahdouraea  hentzi  ally  this  species  to  the  leptostracan  phyl- 
locarids,  viz.,  the  four  well-developed  anterior  pleopods,  the  two  apparently  small 
posterior  pleopods,  the  lack  of  the  seventh  pleopods,  the  robust  pleopod  protopods, 
the  lateral  row  of  pits  on  the  pleopod  distal  branches,  and  the  telson  not  developed 
dorsally  between  the  caudal  rami.  The  recognition  of  the  separate  familial  status  of 
Rhahdouraea  hentzi  seconds  the  suspicions  of  Rolfe  (1969)  that  this  species  was  not 
referable  to  any  of  the  Recent  genera.  This  requires  an  emendation  of  the  diagnosis  of 
the  living  family  Nebaliidae  noted  above. 

Acknowledgments 

Drs.  Robert  Hessler  and  Ian  Rolfe  examined  the  material  and  offered  valuable 
counsel  on  the  taxonomic  issues.  Work  was  supported  in  part  by  NSF  grant  DEB  79- 
03602  (FRS). 

Literature  Cited 

Malzahn,  E.    1958.    Eine  neuer  jungpalaozoischer  Rolfe,  W.  D.  I.     1969.    Phyllocarida.  Pp.  R296- 

Krebs  aus  dem  niederrheinischen  Zechstein.  R331  in  R.  C.  Moore  (ed.).  Treatise  on  Inver- 

Zeit.  deutsch.  geol.  Ges.  1 10:352-359.  tebrate  Paleontology,  Part  R,  Arthropoda  4(1), 

Glaessner,  M.  F.,  and  E.  Malzahn.     1962.    Neue  Geol.  Soc.  Am.  and  Univ.  Kansas  Press,  Law- 

Crustaceen  aus  dem  niederrheinischen  Zech-  rence. 

stein.  Fortschr.  Geol.  Rheinld.  u.  Westf.  6:245-  .    1981.    Phyllocarida  and  the  origin  of  the 

264.  Malacostraca.  Geobios  14:17-27. 


TRANSACTIONS 
OF  THE  SAN  DIEGO 
SOCIETY  OF 

NATURAL  HISTORY 

LIBRARY 

UNlVERBiTY   Volume  20  Number  7  pp.  99-134       18  January  1984 

Phylogeny,  evolution  and  biogeography  of  the  marine  isopod 
Subfamily  Idoteinae  (Crustacea:  Isopoda:  Idoteidae) 

Richard  C.  Brusca 

Department  of  Marine  Invertebrates,  San  Diego  Natural  History  Museum.  San  Diego,  California  92112  USA 

Abstract.  The  patterns  of  spatial  distribution  attained  by  the  genera  of  Idoteinae  are  discussed  in 
hght  of  a  cladistic  analysis  of  the  suborder  Valvifera  and  the  subfamily  Idoteinae.  A  schematic  pattern 
analysis  technique  is  demonstrated  and  reveals  the  probability  of  multiple  origins  of  similar  pleonal 
morphologies  among  various  genera  of  Idoteinae.  Reduction  in  the  maxillipedal  palp  has  occurred 
numerous  times  within  the  Idoteinae.  while  loss  of  the  biramous  uropodal  condition  has  probably 
occurred  twice.  A  geographic  cladogram  of  temperate  Gondwanan  shores  is  proposed.  An  evolutionary- 
biogeographic  narrative  is  presented,  in  which  a  set  of  hypotheses  is  developed  to  describe  the  history 
of  the  Idoteinae  in  time  and  space.  The  subfamily  Idoteinae  appears  to  form  two  principal  lines  of 
descent,  both  arising  in  the  Triassic  or  Jurassic.  One  of  these  lines  remained  closely  tied  to  the  Southern 
Hemisphere  (primarily  Old  World)  temperate  marine  shores  from  which  the  Idoteinae  is  derived.  The 
other  line  invaded  the  Northern  Hemisphere  and  various  New  World  environments,  and  more  recently 
(Cenozoic)  underwent  a  radiation  in  the  American  tropics.  The  success  of  this  latter  lineage  (e.g., 
Erichsonella.  Eusymmerus,  Parasymmerus.  Cleantioides)  may  be  due  to  certain  morphological  and  life 
history  adaptations  not  found  in  New  World  species  of  the  former  line  (e.g.,  Idotea.  Synidotea).  The 
Valvifera  probably  originated  in  the  temperate  Southern  Hemisphere,  at  least  by  Permean/Triassic 
times.  Global  distribution  patterns  of  some  genera  can  be  ascribed  most  parsimoniously  to  vicariance 
processes,  and  in  others  to  dispersal,  ecological  phenomena,  or  a  combination  of  processes.  Other  factors 
have  apparently  also  affected  modern  distributional  patterns  of  idoteine  genera,  for  example,  extinctions. 
Biogeographical  data  can  be  used  to  elucidate  viable  alternative  cladistic  hypotheses  to  those  generated 
solely  on  parsimonious  patterns  of  synapomorphy.  Biogeographic  data  can  also  be  used,  in  conjunction 
with  the  cladogram,  to  identify  probable  ancestral  taxa. 

Introduction 

Few  groups  of  marine  invertebrates  have  enjoyed  analysis  by  cladistic  techniques, 
despite  the  current  popularity  of  the  method.  The  only  previous  attempt  to  examine 
the  phylogenetic  relationships  of  an  isopod  taxon  by  strict  cladistic  (Hennigian)  meth- 
odology was  that  of  Williams  ( 1 970),  who  analyzed  the  relationships  of  North  American 
epigean  species  oi Asellus  (Asellota):  but,  as  was  common  with  early  attempts  at  Hen- 
nigian analyses,  he  used  a  weakly  defined  method  of  character  state  polarity  assessment. 
Despite  this,  he  was  still  able  to  construct  a  very  plausible  ph\  logenetic  hypothesis,  or 
cladogram.  for  the  14  species  he  treated. 

One  reason  for  the  paucity  of  cladistic  studies  on  marine  invertebrates  is  the 
necessity  to  work  with  a  group  whose  taxonomic  relationships  arc  reasonably  well 
known  within  the  context  of  the  larger  hierarchical  taxon  to  which  the  study  group 
belongs.  Without  this  knowledge,  an  assessment  of  character  polarity  is  difficult  to 
obtain,  and  without  polarities,  construction  of  both  cladograms  and  phylogenetic  (evo- 
lutionary) trees  must-  be  based  on  speculatively  generated  hypothetical  ancestors  (ex- 
pressed or  implied).  Although  several  marine  invertebrate  taxa  are  well  understood 
systematically  (e.g..  certain  families  of  molluscs,  crabs,  barnacles),  the  great  majority 
are  not.  Several  crustacean  isopod  groups  are  also  well  understood  in  this  regard.  The 
subfamily  Idoteinae  (suborder  Valvifera;  family  Idoteidae)  is  one  such  group. 

The  present  study  attempts  to  answer  questions  about  the  evolutionary  history  of 
the  Idoteinae  using  cladistic  techniques.  Specific  questions  addressed  arc:  What  are  the 


100 


phylogenctic  patterns  and  relationships  of  the  genera  of  Idoteinae?  What  are  the  spatial 
patterns  and  relationships  of  the  genera  of  Idoteinae?  What  evolutionary  history  is 
suggested  when  these  patterns  are  compared  to  one  another  and  to  the  earth's  geological 
and  ecological  history?  Previous  studies  on  these  topics  include  comments  on  the 
phylogeny  and  biogeography  of  the  genus  Idotea  by  Menzies  (1950c/):  on  the  genus 
Synidotea  by  Menzies  and  Miller  (1972);  and  on  the  comparative  morphology  of  the 
valviferan  higher  taxa  in  general  by  Sheppard  (1957).  A  review  of  the  distribution  of 
shallow-water  idoteine  species  in  the  northeastern  Pacific  was  given  by  Brusca  and 
Wallerstein  (1979/)).  and  a  discussion  of  the  possible  ecological  and  historical  mech- 
anisms regulating  distribution  and  latitudinal  trends  in  morphology  and  behavior  in 
that  taxon  is  provided  by  Wallerstein  and  Brusca  (1982). 

Methods 

The  methods  used  in  this  study  are  largely  summarized  in  3  recently  published 
books  (Eldredge  and  Cracraft  1980,  Nelson  and  Platnick  1981,  Wiley  1981).  However, 
even  the  principal  spokesmen  of  current  cladistic  theory  are  not  without  disagreement 
on  both  details  of  procedure  and  certain  underlying  philosophical  issues.  As  Eldredge 
and  Cracraft  (1979)  point  out,  "No  two  cladists  agree  with  each  other  (or,  for  that 
matter,  with  Willi  Hennig)  on  every  point,  and  this  'school'  of  systematics  is  no  more 
a  monolith  than  that  of  the  more  traditional  'evolutionary  taxonomy'."'  The  overall 
concept  of  cladistic  or  "phylogenctic"  analysis  has  evolved  considerably  since  Hennig 
(1966).  and  indications  are  that  it  will  continue  to  change  for  some  time  to  come.  For 
these  reasons,  and  others.  I  offer  the  following  position  statements. 

While  the  present  study  is  cladistic  in  nature,  it  is  my  opinion  that  such  analyses 
are  most  useful  as  investigatory  techniques  and  do  not  represent  the  final  word  on 
phylogeny.  The  most  powerful  (and  important)  aspect  of  cladistic  methodology  is  its 
ability  to  posit  and  define  monophyletic  groups  in  an  unambiguous  and  testable  manner. 
Synapomorphy  patterns,  however,  do  not  constitute  the  sole  source  of  phyletic  infor- 
mation on  a  taxon.  but  rather  must  be  compared  to  other  kinds  of  data  and  analyses 
when  constructing  phylogenctic  trees,  evolutionary  scenarios,  AND  classifications.  A 
cladogram  depicts  only  a  sequence  of  character  appearances,  which  may  or  may  not 
correspond  to  speciation  events  (Hull  1979).  There  appear  to  be  4  principal  products 
that  can  result  from  phylogenctic  analyses:  cladograms,  phylograms  (evolutionary  trees), 
evolutionary  scenarios,  and  classifications.  The  cladogram  should  be  viewed  as  a  "best 
guess"  in  the  face  of  uncertainty  (Felsenstein  1973,  Harper  1979,  Hull  1979,  Simberloff 
et  al.  1981,  Endler  1982),  and  information  contained  in  any  of  these  other  products  of 
phylogenctic  analyses  can  legitimately  be  used  to  improve  any  other,  including  the 
cladogram  itself  (Hull  1979).  In  the  present  study  I  construct  a  cladogram,  a  phylogram, 
and  an  evolutionary  scenario  for  the  subfamily  Idoteinae.  and  use  the  latter  two  products 
to  shed  new  light  on,  and  make  ammendments  to,  the  cladogram. 

Nomenclature  and  general  terminology  are  taken  from  current  literature  on  val- 
viferan isopods  (see  above  references).  Morphological  structures  discussed  in  this  paper 
are  illustrated  in  Fig.  1.  Analysis  of  character  polarity  is  based  on  out-group  analysis 
(see  Eldredge  and  Cracraft  1980.  de  Jong  1980,  and  Watrous  and  Wheeler  1981).  I 
believe  that  out-group  comparisons  need  not  be  rigidly  constrained  by  nomenclatural 
rank  or  Linnean  hierarchical  structure,  but  are  applicable  at  all  levels  of  a  cladogram. 
Though  parsimony  is  a  potent  methodological  tool,  it  is  primarily  a  method  of  logical 
analysis,  not  a  biological  law  or  principle.  Application  of  parsimony  should  be  an  initial 
technique,  or  one  to  be  used  in  the  absence  of  other  data.  To  continue  to  hold  to  the 


FiciURF  1.  Aspects  of  the  morphology  of  idoteid  isopods  discussed  in  the  text,  a,  Synidotea  harfordi;  note 
multiarticulate  fiagclla  on  antennae  2  and  0+1  pleonal  morphology,  b,  Cleantioides  occidentalism  note  un- 
iarticuiate  (clavate)  flagcllae  on  antennae  2  and  3+1  pleonal  morphology,  c,  Colidotca  findleyi:  note  mul- 
tiarticulate fiagcllae  on  antennae  2  and  0+1  pleonal  morphology,  d.  Mandible  of  Kusyninwrus  antennatus; 


101 


note  large  4-toothed  incisor,  smaller  lacinia  mobilis.  and  stout  molar  process,  e.  Mandible  of  Colidotea 
findleyi;  note  5-toothed  incisor,  large  lacmia  mobilis.  and  stout  molar  process,  f,  Maxilliped  of  Colidotea 
findleyi\  note  4-articulate  palp,  g,  Uniramous  uropod  of  Colidotea  findleyi.  h.  Antenna  2  of  Colidotea  findleyi; 
note  multiarticulate  flagellum.  i.  Antenna  1  of  Colidotea  findleyi;  note  uniarticulate  flagellum.  j.  Antenna  2 
of  Erichsonella  cortezi:  note  uniarticulate  (clavate)  flagellum.  k,  Plcopod  2  of  Enchsonella  cortezi  (male); 
note  appendix  masculinum. 


102 


"simplest"  explanation  (i.e..  the  shortest  cladogram)  in  the  face  of  biological  evidence 
indicating  a  less  parsimonious  but  more  biologically  reasonable  explanation  is  both 
nonscientific  and  an  abuse  of  the  tool.  There  is  little  point  in  creating  a  falsifiable 
hypothesis  if  one  docs  not  accept  all  forms  of  data  that  can  falsify  it.  In  this  regard  I 
agree  with  Kitts  (1981)  that  phylogenetic  patterns  (and  hence  analyses)  ARE*historical 
in  nature  and  this  involves  describing  the  real  world;  one  could  not  be  writing  history 
if  one  supposed  every  relationship  between  events  to  have  transpired  in  the  "most 
direct"  manner.  Whether  or  not  any  particular  phylogeny  is  parsimonious  is  something 
to  be  found  out  in  the  course  of  a  historical  investigation,  it  is  not  something  to  be 
assumed. 

Finally,  my  technique  for  the  historical  biogeographic  analysis  follows  the  hypo- 
thetico-deductive  method,  primarily  as  described  by  Morse  and  White  ( 1979).  I  assume 
no  particular  paradigm  to  be  of  overriding  importance,  but  rather  attempt  to  interpret 
the  patterns  of  characters  and  distributions  in  the  most  parsimonious  (biological  par- 
simony) fashion  possible.  McDowall  (1978)  was.  of  course,  correct  in  noting  that  one 
can  never  know  with  certainty  whether  any  given  individual  component  track  is  the 
product  of  vicariance  or  dispersal.  However,  that  both  phenomena  exist  in  nature  can 
hardly  be  denied,  and  it  has  been  my  task  in  the  present  study  to  decide,  where  possible, 
which  of  these  two  phenomena  (or  others)  produced  the  biogeographic  patterns  seen 
today  in  the  genera  of  Idoteinae.  Needless  to  say.  I  have  been  severely  hampered  in 
this  regard  by  the  absence  of  cladograms  for  other  coastal  marine  taxa. 

Higher  Classification  of  the  Order 

ISOPODA  AND  the  SUBORDER  VALVIFERA 

Out-group  analysis  requires  acceptance  of  some  higher  level  classificatory  structure 
before  statements  regarding  the  relationships  of  lower,  inclusive  taxa  can  be  made.  For 
the  purposes  of  this  study,  I  accept  the  monophyletic  nature  {scnsii  Hennig,  1966)  of 
three  taxa:  the  order  Tanaidacea,  the  order  Isopoda,  and  the  suborder  Valvifera.  The 
monophyletic  nature  of  these  distinct  taxa  are,  to  my  knowledge,  unquestioned. 

While  the  nature  of  the  primitive  isopod  body  plan  (presented  below)  is  generally 
agreed  upon,  the  relationships  of  the  9  isopod  suborders  are  unknown  and  fraught  with 
speculation  (see  Schultz  1979  for  recent  summary  comments).  Various  authors  have 
described  the  nature  of  the  primitive  or  ancestral  isopod  morphotype,  which  is  char- 
acterized by  the  following  combination  of  characters:  carapace  wanting;  pereopods 
uniramous;  respiratory  structures  (heart  and  branchial  surfaces)  primarily  abdominal; 
pereopodal  coxae  forming  marginal  plates  on  pereonites;  first  and  second  antennae 
with  multiarticulate  flagella;  mandible  with  a  multiarticulate  palp;  appendix  masculina 
present  only  on  second  pair  of  pleopods;  uropods  biramous  (probably  attached  ter- 
minally or  subterminally  to  telson  or  pleotelson,  although  some  authors  suggest  a  lateral 
attachment  vis-a-vis  the  cirolanoids);  eyes  entirely  sessile;  all  pereopods  more  or  less 
similar;  pereon  of  7  free  somites  (thoracomeres  2-8);  pleon  of  6  free  somites  and  a 
telson  (or  possibly  5  free  somites  and  a  pleotelson);  maxilliped  with  a  large  basal  endite 
and  reduced  endopodal  articles  (the  latter  forming  the  5-articulate  palp);  maxilliped 
with  a  small,  ovate,  nonbranchial  epipodite  (the  "endognath");  penes  and  opening  of 
vas  deferens  on  thoracomere  8;  simple  foregut;  and  maxillary  glands  present  in  adults. 

This  generalized  ancestral  isopod  plan  was  first  developed  in  the  early  studies  of 
Bate  and  Westwood  (1861-1868),  Stebbing  (1893),  and  Caiman  (1909),  and  more 
recently  by  Schram  (1974)  and  Hessler  et  al.  (1979).  The  concept  of  this  morphotype 
is  supported  by  fossil  data  as  well  as  by  comparison  with  other  peracarid  and  mala- 
costracan  taxa.  It  is  also  compatible  with  all  three  "competing"  hypotheses  of  extant 
primitive  isopod  morphotypes  (i.e.,  cirolanoid,  phreatoicid,  asellote). 

The  Valvifera  stand  apart  as  perhaps  the  most  distinct  of  the  isopod  suborders  in 
several  regards.  Important  features  distinguishing  the  valviferan  body  plan  are  (see  Fig. 
1):  (a)  coxae  of  thoracic  legs  (pereopods)  with  both  dorsal  and  ventral  plates,  the  latter 
extending  over  the  sterna;  (b)  uropods  attached  laterally  on  pleotelson.  but  modified 


103 


to  form  ventral  opercular  plates  covering  the  pleopods;  (c)  vas  deferens  (and  penes) 
opening  on  abdomen  of  male,  rather  than  on  thorax  as  in  all  other  isopods,  excepting 
the  Oniscoidea  (i.e.,  on  pleonite  1  or  on  the  articulation  of  pleonite  1  and  thoracomere 
8);  (d)  flagella  of  first  antennae  reduced  to  one  or  a  few  vestigial  articles;  (e)  pleon  of  4 
or  fewer  free  somites  (plus  the  pleotelson);  (0  uropods  biramous  or  uniramous;  (g) 
maxillipedal  palp  of  3-5  articles:  (h)  second  antennae  uniramous.  fiagellum  multiar- 
ticulate  or  uniarticulate;  (i)  mandible  with  or  without  a  3-jointed  palp.  Attributes  (a) 
and  (b)  are  unique  synapomorphies  that  distinguish  the  Valvifera  from  all  other  isopod 
taxa;  attributes  (c)  through  (0  are  valviferan  synapomorphies  that  also  appear  in  one 
or  more  other  isopod  suborders  (apparent  convergences). 

The  current  classification  of  the  isopod  suborder  Valvifera  is  as  follows: 

Order  Isopoda  Latreille.  1817 
Suborder  Valvifera  Sars,  1882 

Family  Holognathidae  Thomson,  1904 
Family  Idoteidae  Fabricius,  1798 

Subfamily  Idoteinae  Dana,  1852 

Subfamily  Parachiridoteinae  Elkaim  and  Daguerre  de  Hureaux,  1976 

Subfamily  Glyptonotinae  Miers,  1881 

Subfamily  Chaetilinae  Dana,  1852  (=Macrochiridoteinae  Nordenstam.  1933) 

Subfamily  Mesidoteinae  Racovitza  and  Sevastos,  1910 
Family  Xenarcturidae  Sheppard,  1957 
Family  Arcturidae  G.  W.  Sars,  1897 
Family  Amesopodidae  Stebbing,  1905 
Family  Pseudidotheidae  Ohlin.  1901 

The  relationships  of  the  6  valviferan  families  have  long  been  unclear.  The  only 
cogent  discussion  of  the  topic  was  that  of  Sheppard  (1957).  The  cladogram  in  Fig.  2 
depicts  the  best  arrangement  that  I  have  been  able  to  devise  for  these  families,  being 
the  most  parsimonious,  and  admitting  no  convergences,  parallelisms  or  reversals.  A 
convincing  higher  level  classificatory  scheme  of  the  9  isopod  suborders  does  not  pres- 
ently exist,  and  carcinologists  disagree  over  the  relationships  among  these  taxa.  For 
this  reason,  the  Tanaidacea  was  used  as  an  out-group  to  construct  the  cladogram  of 
valviferan  families  (Fig.  2).  Tanaidacea  is  the  peracarid  order  "traditionally"  (Schram 
1981)  taken  to  be  the  most  probable  sister-group  to  the  Isopoda  (also  see  Slewing  1963 
and  Fryer  1964).  Whether  or  not  it  is  the  actual  sister-group  of  the  isopods  is  unim- 
portant for  its  use  in  out-group  comparison,  however,  as  it  is  clearly  a  closely  related 
taxon  within  the  unified  peracarid  line.  Character  polarity  assessments  based  on  tanaids 
were  compared  to  those  obtainable  by  using  the  Amphipoda.  Cumacea  and  hypothetical 
ancestral  isopod  as  out-groups  and  no  changes  in  polarity  were  required  when  these 
other  groups  were  used  in  place  of  tanaids.'  A  step-by-step  discussion  of  the  cladogram 
of  valviferan  families  follows,  the  numbering  in  the  text  following  that  of  Fig.  2. 

Tanaids  are  united  to  the  isopods  only  by  possession  of  their  peracarid  attributes, 
the  most  obvious  of  these  being:  (a)  pereopodal  coxae  with  thin  ventral  plates  (oos- 
tegites)  that  form  a  female  brood  pouch  for  the  developing  young:  (b)  mandibles  with 
lacinia  mobili  in  adult  stages  of  life  cycle,  and  (c)  young  released  from  the  marsupium 
in  subadult  "mancoid"  stage.  At  this  level  of  analysis  these  attributes  are  symplesio- 
morphies:  I  know  of  no  synapomorphies  unique  to  the  tanaids  and  isopods. 

The  Isopoda  are  united  by  the  features  listed  above  for  the  primitive  isopod 
morphotype.  The  first  4  of  these  are  synapomorphies,  as  follow:  (1)  carapace  wanting 
(vs.  present  in  tanaids):  (2)  pereopods  uniramous  (vs.  retaining  vestiges  of  exopods); 
(3)  respiratory  structures  (branchial  pleopods  and  heart)  primarily  abdominal  (vs.  tho- 
racic): (4)  pereopodal  coxae  forming  marginal  plates  on  pereonites  (vs.  not  forming 
plates). - 

The  Holognathidae  shares  in  common  with  its  sister-group  (the  remaining  5  val- 
viferan families)  the  4  valviferan  synapomorphies  listed  earlier:  (5)  pereopodal  coxae 
form  ventral  (sternal)  plates;  (6)  uropods  modified  into  opercular  plates  covering  pleo- 


104 


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II 


Figure  2.     Cladogram  of  families  of  Valvifera.  Closed  boxes  indicate  apomorphies;  open  boxes  plesio- 
morphies.  Numbering  of  characters  corresponds  to  text  discussion. 


pods;  (7)  vas  deferens  and  penes  opening  on  abdomen  of  male,  rather  than  on  thorax; 
(8)  flagella  of  first  antennae  reduced  to  one  or  a  few  vestigial  articles.  The  Holognathidae 
is  a  monotypic  family  containing  but  one  species,  H.  stewarti  (Filhol).  It  cannot  be 
distinguished  by  an  autapomorphy  of  its  own,  although  it  stands  apart  from  all  other 
species  in  the  suborder  Valvifera  by  its  retention  of  a  palp  on  the  mandible,  as  well  as 
numerous  other  primitive  attributes  (e.g.,  biramous  uropods,  5-articulate  maxillipedal 
palp,  pleon  of  4  free  somites  plus  the  pleotelson).  Holognathus  may  be  considered  a 
relict  taxon  within  the  Valvifera.  Nordenstam  (1933)  long  ago  recognized  the  lack  of 
apomorphies  in  Holognathus,  suggesting  that  it  might  best  be  incorporated  into  the 
Idoteidae.  Were  this  done,  however,  Holognathus  would  probably  have  to  be  ranked 
as  a  sixth  subfamily,  rather  than  included  in  the  Idoteinae  as  Nordenstam  suggested. 
The  presence  of  4  free  pleomeres  places  this  genus  at  a  position  ancestral  to  both  the 
Idoteinae  and  the  Glyptonotine-group  discussed  below  (see  Fig.  3).  The  remaining 
valviferan  families  are  thus  united  by  the  loss  of  the  mandibular  palp  (9).  Clearly,  were 
one  to  place  Holognathus  in  the  Idoteidae,  characters  5-8  would  become  synapomor- 
phies  uniting  Idoteidae  to  all  other  valviferan  families. 

The  Idoteidae  stand  apart  as  the  only  family  in  which  the  uropods  may  be  reduced 
from  the  primitive  biramous  state  to  a  uniramous  condition  (10),  and  in  which  a 
reduction  of  the  maxillipedal  palp  takes  place  (11).  The  Idoteidae  have  been  charac- 
terized by  two  other  "synapomorphic  trends":  trends  towards  fusion  of  the  pleonites 
and  towards  fusion  (or  loss)  of  the  flagellar  articles  on  the  second  antennae.  These 
reductions,  however,  are  convergent  to  patterns  that  also  occur  in  other  valviferan 
families  and  thus  have  not  been  used  to  construct  the  cladogram  (Fig.  2).  The  Idoteidae 


105 


4+0 


GIvP'onoline  group 


3+1 


to  Idoleinae 


3+1 


lenobmna 


3+1 


2+2 


1+3 


0*4 


0+3 


0+2 


0+1 


0+0 


i 

3+0 


LINEAGE  A  teduclion  of 
amennal  flagella 


KJ 


Cleanlioides 


I 

LINEAGE  B    no  reduction  of 
antennal  flagella 

i 

2+1  1+2 


1+2 


Lyidolea 


Cdolea  Enchsonella 

Parasymmerus 
Eusymmerus 


Cleantiella 


0+3 


0+2 


0+1 


0+0 


Idolea  Engidolea         Glyptidolea 

1  Paridotea  Synischia 

2+0  Pentias 

Crabvzos 
LUidotea 


Colidolea  Synisoma 


1+1 


0+2 


0+1 


1+0 


Synidolea 
Mop/isa 


Barnardidolea 


FiGL'RE  3.  Schematic  representation  of  possible  phylogenetic  pathways  for  pleonal  fusion  in  the  Idoteinae. 
Pleonal  formulas  are  written  above  pleonal  diagrams  and  indicate  number  of  free  and  number  of  fused 
pleomeres.  Genera  assigned  to  given  pleonal  morphology  are  indicated  below  diagrams.  See  text  for  discussion. 


Stands  out  most  strikingly  from  the  4  families  that  comprise  its  sister-group  in  the 
retention  of  numerous  plesiomorphic  attributes  (e.g.,  free  penes  retained  in  4  of  the  5 
subfamilies;  free  cephalon;  one  pair  of  appendix  masculina.  on  the  second  pleopods). 
Racovitza  and  Sevastos  (1910)  long  ago  recognized  the  primitive  nature  of  the  Idoteidae. 
regarding  it  as  an  '"ancient"  family.  The  Idoteidae  was  the  first  valviferan  family  to  be 
described,  subsequent  families  being  distinguished  from  it  by  elucidation  of  new  char- 
acters acquired  outside  the  Idoteidae.  Thus,  historically  a  diagnosis  of  the  Idoteidae 
has  been  developed  largely  upon  absence  of  characters  (a  phenomenon  common  among 
older  taxa).  The  discovery  of  new  distinguishing  attributes  (apomorphies)  for  the  Hol- 
ognathidae  and  Idoteidae  is  clearly  needed  and  will  provide  important  data  for  testing 
the  hypotheses  contained  in  the  cladogram. 

The  Xenarcturidae  and  its  sister-group  (Arcturidae-Amesopodidac-Pseudido- 
theidae)are  distinguished  by  the  following  synapomorphies:  ( 12)  first  pleopods  of  males 
bearing  "accessory  appendix  masculina"  (in  addition  to  the  true  appendix  masculina 
of  the  second  pleopods);  (13)  cephalon  fused  medially  to  pereonite  I  (second  thorac- 
omere);  (14)  peduncle  of  first  pleopods  greatly  elongated.  The  Xenarcturidae  is  a  mono- 
typic  taxon  distinguished  by  the  following  autapomorphies:  (15)  pereonites  I-IV  with 
lateral  margins  expanded  into  large  plates  covering  bases  of  pcreopods;  (16)  second 
antennae  with  flagella  reduced  to  single  articles;  and  (17)  flagella  of  second  antennae 
directed  towards  mouth,  rather  than  away  from  buccal  field. 

The  Arcturidae  and  its  sister-group  (Amesopodidae-Pseudidotheidae)  are  distin- 
guished by  a  synapomorphy  in  the  functional  grouping  of  the  pcreopods  (18).  Only  in 
these  taxa  are  pcreopods  I-IV  similar  and  directed  forward  to  form  a  functional  group 


106 


distinct  from  pereopods  V-VII.  In  all  other  valviferan  taxa  the  percopodal  functional 
grouping  is  I-III  and  I  V-VII.  Arcturidae  is  distinguished  by  two  synapomorphies:  (19) 
the  unique  body  shape  (cylindrical  or  tubular,  often  geniculate),  and  (20)  having  per- 
eonite  IV  generally  manifestly  enlarged  or  elongated. 

The  Amesopodidae  and  Pseudidotheidae  are  distinguished  by  the  synapomorphic 
condition  of  having  pereonites  II-IV  grossly  enlarged  (21).  Amesopodidae  is  a  mono- 
typic  family  {A.  richardsonae  Stebbing,  1905)  distinguished  by  the  autapomorphies  of 
highly  reduced  second  pereopods  (22),  and  the  complete  loss  of  pereopods  III  and  IV 
(23).  Pseudidotheidae  contains  two  genera  distinguished  by  the  synapomorphy  of  fusion 
of  the  first  two  articles  of  the  peduncle  of  the  second  antennae  (24). 

The  Family  Idoteidae  and  the  Subfamily  Idoteinae 

The  systematic  history  of  the  family  Idoteidae  can  be  traced  through  the  following 
works:  H.  Milne  Edwards  (1840),  Dana  (1853),  Bate  and  Westwood  (1868),  Harger 
(1880),  Miers  (1881).  Chilton  (1890),  Ohlin  (1901),  Richardson  (1905a),  Stebbing 
(1905),  Collinge  (1917),  Barnard  (1920),  Nordenstam  (1933),  Menzies  (1950fl),  Shep- 
pard  (1957),  and  Menzies  and  Miller  ( 1972).  The  American  idoteid  fauna  is  well  known, 
largely  due  to  the  work  of  Dana  (1853),  Harger  (1880),  Benedict  (1897),  Richardson 
(1899a.  /),  1900,  1901,  1904,  1905a.  b,  1909),  Hatch  (1947),  Menzies  and  Waidzunas 
(1948),  Menzies  (1950a.  b),  Menzies  and  Bowman  (1956),  Menzies  and  Frankenberg 
(1966),  Menzies  and  Miller  (1972),  and  Brusca  and  Wallerstein  (1977,  1979a./)).  All 
species  of  Idoteidae  are  marine,  although  two  species  of  the  subfamily  Mesidoteinae 
also  extend  their  distributions  into  fresh  water.  Saduna  {=Mesidotea)  entoniofi  has 
been  found  in  several  deep  Scandinavian  lakes,  and  Austridotea  lacustris^  occurs  from 
the  littoral  zone  to  fresh  water  rivers  and  lagoons  in  New  Zealand. 

The  subfamily  Idoteinae  contains  22  valid  genera  (Table  1).  The  great  majority 
are  shallow-water  and,  for  the  most  part,  intertidal  species.  Few  species  are  restricted 
to  depths  greater  than  30  m.  For  the  past  150  years  (since  the  work  of  Brandt  1833 
and  H.  Milne  Edwards  1840)  studies  on  this  group  have  consistently  found  that  the 
use  of  a  few  clearly  defined  characters  provided  a  basis  for  a  classification  that  has  been 
both  stable  and  reliable.  Thus  most  idoteine  genera  are  clearly  defined,  unambiguous, 
and  easily  distinguished  from  one  another.  As  will  soon  be  seen,  however,  not  all 
idoteine  genera  can  be  defined  by  unique  apomorphies.  The  few  genera  that  are  not 
clearly  differentiated  from  one  another  comprise  2  small  groups  of  largely  monotypic. 
Southern  Hemisphere  genera  that  are  in  need  of  reexamination.  Principal  characters 
used  to  distinguish  the  idoteine  genera  are  external  and  easily  recognized,  as  follows. 

Uropods.  —  Tht  uropods  of  Idoteinae  are  either  biramous  or  uniramous.  The  prim- 
itive biramous  condition,  while  being  clearly  distinct  from  the  uniramous  condition 
and  hence  useful  in  pattern  analysis,  is  not  understood  ontogenetically  (see  Caiman 
1909,  Racovitza  and  Sevastos  1910,  Tait  1917,  Nordenstam  1933,  and  Menzies  and 
Miller  1972).  Loss  of  one  uropodal  ramus  has  occurred  at  least  twice  among  the  val- 
viferan families,  in  the  Idoteidae  (subfamily  Idoteinae)  and  again  in  a  single  species  of 
Arcturidae  {Microarcturus  digitalis  Nordenstam  1933).  Whether  or  not  these  separate 
losses  were  by  the  same  "mechanism"  is  not  known. 

Pleon.  —  Isopod  taxa  are  characterized  by  varying  degrees  of  fusion  of  the  pleomeres 
and  telson.  Although  trends  towards  fusion  of  pleomeres  are  evident  throughout  the 
Isopoda  and  occur  in  every  suborder,  no  one  has  yet  attempted  to  analyze  these  mor- 
phoclines  in  a  systematic  fashion  (Fig.  3).  In  the  Valvifera,  there  are  always  four  or 
fewer  free  pleomeres,  plus  the  pleotelson.  The  term  "pleotelson"  refers  to  that  region 
of  the  pleon  consisting  of  the  telson  and  its  fused  pleomeres.  For  many  years,  the  pattern 
of  discrete  character  states  manifested  by  fusion  of  pleomeres  in  the  idoteine  genera 
has  been  taken  to  represent  a  morphocline  that  is  a  sequence  of  phenotypes  presumed 
to  reflect  the  probable  evolutionary  pathway  of  descent.  The  polarity  or  direction  of 
this  morphocline  is  clearly  shown  by  out-group  analysis  to  be  towards  levels  of  in- 
creasing pleomere  fusion.  Fusion  of  the  pleomeres  is  often  (but  by  no  means  always) 
indicated  by  the  presence  of  partly  free  lateral  margins,  distinguishable  by  the  presence 


107 


Table  1.  Summar\-  of  "traditional  characters"  used  to  difFcrentiate  the  genera  of  Idoteinae  (from  Menzies 
and  Miller  1972;  with  corrections).  See  text  for  additional  characters.  Edotca  includes  the  synonym  Ilpclys. 
and  Zcnobiana  includes  the  synonym  Cleantis.  Erichsonella  includes  the  synonyms  Erichsonia  Dana  and 
Ronalca  Men/ies  and  Bowman.  Pircs  (pers.  comm.)  has  a  manuscript  in  preparation  in  which  she  intends 
to  remove  the  monot>  pic  genus  Ronalca  from  synonymy  with  Erichsonella.  based  on  the  alleged  presence 
of  a  single  pair  of  lateral  incisions  in  R.  pseudoculata  (Boone).  I  have  not  examined  R.  pseiidoculata  myself 
Such  a  change  would  require  a  minor  revision  in  the  cladogram  (Fig.  17b).  by  adding  Ronalca  as  a  fork 
at  the  lip  of  the  line  leading  to  Eusynmicrus.  making  these  two  genera  sister-taxa. 


No. 

Flagelh 
Multi- 

am  of  Antenna  2 

Pleonal 

arti- 
cles 
in 

Uropods 

articu- 

Clav- 

Vesti- 

form- 

Mxp 

Unira-       Bira- 

Genus 

late 

ate 

gial 

ula 

palp 

mous       mous 

Barnaididotca  Menzies  &  Miller 

X 

1  +0 

3 

X 

Crabyzos  Bate 

X 

0  +  3 

5 

X 

Engidolea  Barnard 

X 

1  +  2 

5 

X 

Glyptidotca  Stebbing 

X 

0  +  3 

5 

X 

Moplisa  Morcira 

X 

0+  1 

3 

X 

Pentias  Richardson 

X 

0  +  3 

5 

X 

Pandotca  Stebbing 

X 

1  +  2 

5 

X 

Synidotea  Harger 

X 

0+  1 

3 

X 

Synischia  Hale 

X 

0  +  3 

5 

X 

Idolea  Fabricius 

X 

2+  1 

4-5 

X 

Colidotea  Richardson 

X 

0+  1 

4 

X 

Synisoma  Collinge 

X 

0  +  0 

2  +  "" 

4 

X 

Zenobianopsis  Hale 

X 

4+  1    • 

5 

X 

Euidotea  Collinge 

X 

0  +  3 

4 

X 

Cleantiella  Richardson 

X 

1  +2 

5 

X 

Erichsonella  Richardson 

X 

0  +  0 

4 

X 

Eusynmicrus  Richardson 

X 

0+  1 

4 

X 

Parasymmerus  Brusca  &  Wallenstein 

X 

0+  1 

3 

X 

Lyidolca  Hale 

X 

0  +  3 

3 

X 

Zcnobiana  Risso  (as  Zcnobia) 

X 

X 

3+  1 

5 

X 

Cleantioides  Kensley  &  Kaufman 

X 

3+  1 

4-5 

X 

Edotca  Guerin-Meneville 

X 

0+  1 

3 

X 

of  lateral  incisions  (generally  referred  to  as  "suture  lines'").  Presence  of  free  lateral 
margins  on  the  fused  pleomeres  is  taken  to  represent  incomplete  fusion  (i.e.,  medial 
fusion  only)  of  these  somites,  and  hence  a  less  derived  state  than  absence  of  the  free 
lateral  margins  (i.e.,  complete  fusion). 

Maxi//ipech.  —  The  palp  of  the  idoteid  maxilliped  is  composed  of  5  or  fewer  articles. 
Out-group  analysis  indicates  the  plesiomorphic  state  (occurring  in  all  families  except 
Idoteidae)  is  5  free  articles;  reduction  in  the  number  of  articles  thus  represents  a  derived 
condition.  It  is  not  known  with  certainty  whether  reduction  in  the  number  of  articles 
is  due  to  fusion  or  to  actual  loss,  although  Brusca  and  Wallerstein  ( 1 979c/)  have  suggested 
that  both  processes  may  exist  among  various  genera  (e.g.,  loss  in  Idotca,  fusion  in 
Zenobiaua). 

Anwnnac  — In  valviferans,  the  flagellum  of  the  second  antenna  is  either  (1)  nuii- 
tiarticulate  (the  primitive  condition);  (2)  reduced  to  a  single  clavate  article  with  1  to  4 
minute  "vestigial"  apical  articles;  (3)  reduced  to  a  single  clavate  article  only;  or  (4) 
reduced  to  just  the  minute  "vestigial"  articles.  Brusca  and  Wallerstein  i\979a)  point 
out  that  these  two  kinds  of  reduction  (clavate  vs.  vestigial  articles)  arc  probably  the 
result  of  two  different  processes,  the  former  being  a  case  of  fusion  of  the  tlagellar  articles, 
the  latter  being  an  actual  loss  or  reduction  in  the  number  of  articles,  creating  a  "vestigial" 
flagellum.  In  one  genus  (Zenobiaua)  both  the  clavate  condition  and  or  the  vestigal 
condition  may  both  occur,  suggesting  that  the  two  morphologies  are  somehow  linked, 
perhaps  both  dcvelopmentally  and  phylogenetically.  The  early  reduction  was  apparently 
a  situation  in  which  partial  fusion  of  the  flagellar  articles  produced  the  clavate  mor- 


108 


phology  while  retaining  a  few  vestigial,  unfused,  apical  articles  (as  seen  in  some  7.en- 
ohiana  species).  This  condition  could  have  progressed  in  either  of  two  directions  — loss 
of  the  vestigial  articles  to  leave  just  the  remaining  clavate  process  (as  in  Cleantiella, 
Erichsonella,  Euysninierus,  Parasyinnieriis,  Cleantioides,  and  Lyidotea),  or  loss  of  the 
clavate  process  to  leave  just  the  remaining  vestigial  articles  {Edotea  and  some  species 
of  Zenobiana).  The  relationships  between  the  clavate  and  vestigial  conditions  may  not 
be  resolvable  in  a  phylogenetic  sense,  and  may  represent  differing  avenues  of  a  flexible 
developmental  program.  However,  unlike  the  pleonal  and  maxillipedal  palp  charac- 
teristics (above)  which  show  varying  degrees  of  reduction,  the  antennal  flagella  are 
either  reduced  (states  2-4  above)  or  not  reduced. 

Co.xal  plates.  Although  the  coxal  plates  have  been  used  extensively  in  valviferan 
taxonomy,  use  of  these  structures  has  not  been  consistent.  Previous  workers  have  treated 
these  structures  in  a  variety  of  ways  and  one  worker's  description  is  not  always  com- 
parable to  another's.  These  problems  have  been  discussed  at  length  by  Nordenstam 
(1933),  Sheppard  (1957),  and  Brusca  and  Wallerstein  (1979a).  For  these  reasons,  the 
coxal  plates  are  not  considered  in  the  following  analysis. 

While  the  genera  of  the  Idoteidae  appear  to  be  reasonably  well-defined,  the  5 
nominate  subfamilies  are  not.  The  subfamily  Idoteinae  stands  apart  from  the  other  4 
in  numerous  features,  and  appears  to  represent  a  monophyletic  group.  The  other 
subfamilies  (Glyptonotinae,  Chaetilinae,  Parachiridoteinae,  and  Mesidoteinae)  cannot 
be  easily  separated  from  one  another,  nor  be  distinguished  unambiguously  in  a  clado- 
gram.  For  this  reason,  these  4  subfamilies  collectively  are  herein  considered  an  out- 
group  of  the  Idoteinae.  They  may  be  thought  of  as  representing  an  unresolved  poly- 
chotomy  on  the  cladogram  in  Fig.  4.  In  the  following  discussion  these  4  subfamilies 
are  treated  as  one  and  referred  to  as  the  "glyptonotine-group.""*  A  second  out-group 
used  to  construct  a  cladogram  of  Idoteinae  genera  is  the  Holognathidae  (see  Fig.  2). 
Numbers  in  the  following  discussion  correspond  to  that  on  the  cladogram  in  Fig.  4. 

The  glyptonotine-group  is  distinguished  by  the  following  synapomorphies:  (1)  ce- 
phalon  strongly  produced  laterally,  moving  eyes  to  dorsal  position;  (2)  body  broadened 
and  dorsoventrally  depressed;  (3)  pereopods  I-III  subchelate  or  prehensile.  It  retains 
the  symplesiomorphy  of  separate  penes.  The  Idoteinae  is  distinguished  by  the  following 
synapomorphies:  (4)  reduction  of  the  pleon  to  the  3+1  condition;  (5)  fusion  of  the 
penes  into  a  single  structure  (Fig.  4). 

Paleontological  data,  limited  as  they  are,  corroborate  the  out-group  comparison 
for  the  Idoteinae.  The  oldest  known  idoteid  fossils  are  referred  to  the  monotypic  genus 
Proidotea  {P.  haugi  Racovitza  and  Sevastos,  1910),  from  mid-  to  late  Oligocene  deposits 
of  eastern  Europe.  This  genus  clearly  falls  within  the  subfamily  Mesidoteinae  (the 
glyptonotine-group).  The  only  other  fossil  data  for  the  Valvifera  are  Pleistocene  to 
Recent  specimens  of  Saduria  {=AIesidotea)— probably  the  holarctic  5".  sabini  (Kroyer). 
In  both  of  these  genera,  the  pleon  is  composed  of  4  somites,  plus  the  pleotelson.  The 
maxillipedal  palp  of  Saduria  is  5-articulate.  The  uropods  of  Mesidoteinae  are  biramous, 
as  in  the  subfamilies  Glyptonotinae,  Chaetilinae,  and  Parachiridoteinae. 

The  genus  Zenobianopsis  Hale,  1946  is  not  indicated  on  the  cladogram  (Fig.  4). 
The  status  of  this  deep  water  taxon  is  uncertain.  Only  two  species  have  been  reported 
and  they  differ  markedly  in  morphology.  Species  of  Zenobianopsis  have  a  pleon  of  4 
free  somites,  plus  indications  of  a  fifth  (although  in  Z.  caeca  Hale,  1946,  these  somites 
appear  somewhat  reduced).  Other  attributes  indicate  that  Zenobianopsis  is  a  very 
primitive  member  of  Idoteidae  (Table  1),  presumably  with  its  origin  at  or  about  the 
time  of  separation  of  the  Idoteinae  from  the  glyptonotine-group.  The  early  isolation  of 
these  species  is  further  suggested  by  the  fact  that  both  are  known  only  from  deep 
subantarctic  waters  of  the  Indian  Ocean.  Zenobianopsis  is  indicated  by  a  dashed  line 
in  the  evolutionary  tree  in  Fig.  1 6;  further  research  may  place  the  two  species  in  separate 
genera. 

Within  each  of  the  two  principal  idoteine  lineages  depicted  in  Fig.  4,  the  trend 
towards  fusion  of  pleomeres  is  expressed  in  a  "directed''  fashion.  That  is,  the  greater 
the  degree  of  pleomere  fusion  in  a  taxon,  the  farther  up  the  cladogram  it  appears.  Fig. 


109 


CO 

c 
ex 

1 

o 

s 

:t3 
1 

CO 

o 

3: 

CD 

i 

Co 


N 


FiGi'RE  4.     Cladogram  of  Idoteinae  genera.  Only  apomorphies  are  indicated.  Numbering  of  characters  cor- 
responds to  discussion  in  text.  For  characters  uniting  Holognathidae  to  Idoteidae  see  Fig.  2  and  text. 


3  provides  a  schematic  representation  of  known  pleonal  morphologies  beginning  with 
the  4-segmented  pleon  found  in  the  glyptonoline-group.  The  pleonal  formulas  are 
written  in  two  digits,  separated  by  a  plus  sign.  The  first  digit  is  the  number  of  complete 
pleomeres  present  in  the  pleon  (not  counting  the  pleotelson);  the  second  digit  is  the 
number  of  lateral  incisions  present,  representing  remnants  of  incompletely  fused  pleo- 
meres. 

There  are  14  possible  combinations  or  pleonal  formulas  that  species  of  Idoteinae 
might  possess:  3+1.  3+0,  2+2,  2+1.  2+0,  1+3,  1+2,  1  +  1,  1+0.  0+4.  0+3.  0+2.  0+1. 
0+0.  However,  there  are  over  100  possible  different  steps  in  which  fusion  may  progress 
to  give  rise  to  these  14  combinations.  Each  of  these  100+  pleonal  morphologies  is 
derived  in  a  unique  manner  and  hence  each  constitutes  an  '"attribute"  (scnsii  Platnick. 
1979:542).  Not  every  pathway  is  represented  in  Fig.  3:  only  enough  steps  are  shown 
in  order  to  reach  the  existing  morphologies  of  the  known  idoteine  genera  in  a  parsi- 
monious fashion.  This  large  ''uncertainty"  problem,  as  well  as  the  seeming  parallelism 
of  pleonal  fusion,  can  be  resolved  by  careful  examination  of  the  schematic  pattern 
analysis  in  Fig.  3.  All  known  genera  of  Idoteinae  have  pleonal  formulas  that  must  have 
arisen  from  one  of  two  main  lines:  one  line  beginning  with  a  3+  1  configuration  (indicated 
on  Figs.  3  and  4  as  "lineage  A"),  and  the  other  beginning  with  a  2+1  configuration 
(indicated  on  Figs.  3  and  4  as  "lineage  B").  Assignment  of  genera  to  one  or  the  other 
of  these  lineages  can  be  based  on  the  antennal  features  described  above.  That  is.  the 
13  genera  with  multiarticulate  second  antennal  flagella  arc  hypothesized  to  represent 
a  lineage  or  series  distinct  from  the  8  genera  that  have  lost  the  multiarticulate  conditions. 
Any  other  assignment  of  these  genera  requires  accepting  convergent  evolution  of  these 
antennal  morphologies;  parsimony  is  maintained  by  presuming  these  antennal  mor- 
phologies to  have  arisen  only  once.  This  split  clearly  places  all  genera  on  one  or  the 
other  of  these  two  main  lines  of  descent  involving  pleomcrc  fusion,  and  further  suggests 
that  evolution  of  identical  pleonal  formulas  in  these  two  lines  of  descent  was  through 
different  steps,  and  hence  not  true  convergence  at  all. 

Not  all  14  possible  patterns  are  represented  by  extant  species.  In  fact,  only  9  are 
known  at  present  (4+1.3+1.2+1.  2+2.  1  +2.  1  +0.  0+3.  0+ 1 .  0+0).  Other  morphologies 


presumably  existed  in  the  past  (in  extinct  intermediate  taxa)  or  exist  at  present  but 
await  discovery.  Fig.  3  predicts  the  nature  (overall  morphology)  of  these  yet  to  be 
discovered  genera,  and  hence  sets  the  stage  for  testing  the  hypotheses  contained  therein. 

Fig.  3  is  not  a  cladogram  OR  an  evolutionary  tree.  It  is  merely  a  graphic  arrange- 
ment of  pleonal  morphologies  in  a  sequence  of  most  primitive  at  the  top,  witfi  increas- 
ingly derived  morphologies  (increased  fusion)  expressed  following  the  arrows  down 
and  across.  As  can  be  seen  in  Figs.  3  and  4,  reduction  in  pleomere  number  corroborates 
the  reduction  in  the  antennal  flagellum  (i.e.,  there  are  no  conflicts). 

Lineage  B  (Fig.  4)  is  distinguished  by  two  synapomorphies:  (6)  pleonal  fusion  has 
advanced  to  the  2+ 1  morphology,  and  (7)  the  uropods  have  lost  the  primitive  biramous 
morphology,  deriving  a  uniramous  condition.  Lineage  A  is  defined  by  the  synapo- 
morphy  of  (8)  antennal  flagellum  reduction  to  condition  2  described  above  (i.e..  fusion 
of  most  flagellar  articles  into  a  clavate  process  bearing  on  its  apex  a  few  remaining 
"vestigaf"  articles). 

Within  lineage  B,  the  genus  Idotea  is  distinguished  by  the  apomorphy  of  (9)  re- 
duction in  maxillipedal  palp  article  number;  the  sister-group  to  Idotea  has  reduction 
of  pleonal  morphology  beyond  the  2+1/2+0  condition  (10).  Baruardidotea,  Moplisa, 
and  Synidotea  have  maxillipedal  palp  reduction  to  3  articles  (1 1),  as  well  as  continued 
fusion  of  pleomeres  to  produce  a  1+0  pleonal  morphology  (12).  Baruardidotea  is 
distinguished  from  Moplisa  and  Synidotea  by  retention  of  the  1+0  pleonal  formula, 
while  the  latter  two  genera  have  achieved  the  0+ 1  condition  (13).  Moplisa  and  Synidotea 
can  be  distinguished  from  one  another  by  the  loss  of  the  molar  process  in  the  former 
(14). 

The  Paridotea-Engidotea  line  is  distinguished  from  its  sister-group  by  retention 
of  the  symplesiomorphic  pleonal  condition,  1+2  (vs.  the  synapomorphic  condition, 
0+3)  (15).  These  two  genera  cannot  be  clearly  distinguished  from  one  another  on 
morphological  criteria  as  they  are  currently  understood.  The  Colidotea-Synisoma  group 
is  distinguished  from  its  sister-group  by  two  synapomorphies:  (16)  lacinia  mobilis  of 
mandible  greatly  enlarged  (as  large  or  larger  than  incisor  process),  and  (17)  reduction 
to  a  4-jointed  maxillipedal  palp.  Colidotea  is  distinguished  by  (18)  a  0+1  pleonal 
morphology.  Synisonia  is  distinguished  by  two  synapomorphies:  (19)  pleonal  mor- 
phology 0+0,  and  (20)  an  elongate  pleon  ('/3  or  more  total  body  length).  Euidotea  is 
distinguished  from  its  sister-group  by  (21)  possession  of  a  4-jointed  maxillipedal  palp 
(vs.  5-jointed  in  the  Glyptidotea-Pentias-Crabyzos-Synischia  line).  The  latter  4  genera 
cannot  be  distinguished  by  morphological  criteria  as  they  are  currently  understood  and 
are  hereafter  referred  to  as  the  Glyptidotea-group. 

Within  lineage  A,  Zenohiana  can  be  distinguished  only  by  the  symplesiomorphous 
retention  of  biramous  uropods:  whereas  its  sister-group  has  achieved  the  synapomor- 
phic condition  of  uniramous  uropods  (22).  Some  species  of  Zenohiana  have  lost  the 
minute  apical  articles  on  the  tip  of  the  second  antennae,  while  others  retain  these 
terminal  articles.  I  am  in  agreement  with  previous  authors  that  these  differences  do 
not  warrant  the  splitting  of  Zenohiana  into  several  genera  (e.g..  Bate  and  Westwood 
1861-1868,  Issel  1913,  Collinge  1917,  Barnard  1925).  I  have  not  taken  the  presence 
or  absence  of  the  "vestigial"  articles  into  consideration  in  construction  of  the  cladogram. 

Cleantioides,  as  defined  here,  contains  only  two  species,  C.  occidentalis  (Richard- 
son) and  C.  planicaitda  (Benedict).  This  genus  is  distinguished  by  2  synapomorphies: 
(23)  reduction  of  the  fourth  pair  of  pereopods  to  nonambulatory  appendages,  and  (24) 
reduction  of  the  maxillipedal  palp  to  4  articles  in  one  species  (C  occidentalis).  It  retains 
the  symplesiomorphic  3+ 1  pleonal  morphology,  while  its  sister-group  has  achieved  the 
2+2  or  less  stage  (25). 

Cleantiella  is  distinguished  by  the  apomorphy  of  pleonal  reduction  to  the  1+2 
condition  (26).  Lyidotea  and  its  sister-group  bear  2  synapomorphies:  (27)  the  pleon 
has  achieved  the  0+3  morphology,  and  (28)  the  maxillipedal  palp  has  been  reduced  to 
the  3-  or  4-articulate  condition.  Lyidotea  bears  an  apomorphic  condition  in  which  the 
last  perconal  somite  has  become  fused  with  pleomere  1  (29);  its  sister-group  is  distin- 
guished by  the  0+2  or  less  pleonal  morphology  (30). 


Ill 


Erichsonella  is  distinguished  by  the  apomorphy  of  (31)  complete  pleonal  fusion, 
resulting  in  the  0+0  condition,  although  some  species  retain  a  faint  trans\erse  furrow 
presumably  representing  the  fused  juncture  of  pleomere  1  to  the  remainder  of  the  pleon. 
Erichsonelld's  sister-group  {Eiisymmerus-Parasytnmerus-Edotea)  has  achieved  the  0+  1 
morphology  (32).  Edotea-Parasyninierus  are  defined  by  the  synapomorphy  of  maxil- 
lipedal  palp  reduction,  to  the  3-articulate  condition  (33).  Edotca  is  distinguished  by  its 
acute  subtriangular  pleon  (34).  Eusyninicriis  is  distinguished  by  the  unique  3-pointed 
spine  that  arises  between  the  lacinia  and  incisor  process  on  the  left  mandible  (35). 

This  cladogram  (Fig.  4)  reduces  the  number  of  character  convergences  to  two. 
Reduction  in  the  number  of  articles  on  the  maxillipedal  palp  appears  to  take  place  time 
and  again,  throughout  the  various  idoteine  lines,  and  no  sensible  cladogram  design  can 
eliminate  it.  In  Fig.  4  it  occurs  four  times  in  lineage  A  and  three  times  in  lineage  B. 
For  this  reason,  it  was  given  lower  priority  than  all  other  characters  used  in  the  analysis. 
Maxillipedal  palp  reduction  is  common  throughout  the  order  Isopoda  and  represents 
a  kind  of  convergence  or  parallelism  known  as  canalized  evolutionary  potential.  This 
character  also  expresses  both  "inside"  and  "outside"  parallelism  (sensu  Brundin  1976. 
1981).  The  second  convergence  (or  parallelism)  is  loss  of  the  ancestral  biramous  uropod 
condition,  which  takes  place  in  (and  characterizes)  lineage  B,  and  then  again  in  the 
Cleantioides  through  Eusymmerus  line  of  lineage  A  (synapomorphy  22). 

While  the  cladogram  in  Fig.  4  is  highly  parsimonious,  what  is  clearly  needed  is 
the  elucidation  of  additional  synapomorphies  to  further  test  the  contained  hypotheses. 
Six  terminal  taxa  (or  groups)  cannot  be  distinguished  by  unique  apomorphies  at  this 
time.  As  Sheppard  (1957)  and  Brusca  and  Wallerstein  (1979a)  pointed  out,  a  critical 
examination  of  the  nature  of  the  pereopodal  coxae  in  the  various  idoteid  genera  will 
surely  prove  enlightening  and  undoubtedly  provide  us  with  a  new  suite  of  character 
states  with  which  to  test  evolutionary  relationships  among  the  genera.  The  exact  nature 
of  the  reduction  in  flagellar  articles  on  the  second  antennae  needs  to  be  investigated 
(particularly  regarding  Zenobiana,  Cleantioides,  and  Edotea),  and  this  too  might  shed 
new  light  on  the  cladogram.  This  reduction,  as  well  as  reduction  in  maxillipedal  palp 
articles  and  free  pleomeres,  tends  to  follow  Brown's  (1965)  "Rule  of  Evolutionary 
Reduction."  Finally,  two  unresolved  polychotomies  exist  within  the  Idoteinae  that  can 
be  resolved  only  by  synonymizing  genera  or  by  the  elucidation  of  new  characters  to 
differentiate  these  taxa.  It  is  my  belief  that  a  careful  study  of  the  coxal  plates,  lacinia 
mobili.  second  antennae,  and  pleopods  3-5  could  resolve  all  of  these  problematic  areas 
among  the  idoteine  genera. 

BlOGEOGRAPHY  AND  EVOLUTION  OF  THE  IdOTEINAE  GeNERA 

Idoteids,  like  other  isopods,  brood  their  developing  young  in  a  marsupium.  from 
which  they  are  hatched  as  "mancas,"  which  are  subjuveniles  that  resemble  miniature 
adults  except  for  lacking  the  seventh  pair  of  pereopods.  There  is  no  evidence,  ecological 
or  morphological,  that  the  manca  stage  is  planktonic.  and  in  those  species  that  have 
been  reared  in  the  laboratory,  hatchlings  are  always  benthic  crawlers  like  their  parents. 
The  only  published  records  of  idoteids  in  plankton  are  for  the  2  widespread  species, 
Cleantioides  planicaiida  and  Idotea  metallica,  and  the  2  species  belonging  to  the  ques- 
tionable genus  Zenohianopsis.  Both  mancas  and  adults  of  most  species,  however,  are 
capable  of  swimming  in  short  bursts  over  small  distances  (e.g.,  between  algal  fronds). 
When  idoteids  are  dislodged  from  the  substratum  by  waves  or  surge  they  sink  or  swim 
quickly  to  the  bottom  (Jansson  and  Matlhiesen  1971,  Salemaa  1979,  Sywula  1964.  Lee 
and  Gilchrist  1972.  Wallerstein  and  Brusca  1982).  Idoteids  are  clearly  a  component  of 
the  benthic  community  in  marine  shallow-water  habitats,  where  their  niche  is  that  of 
a  cryptic  herbivore  and  occasional  scavenger.  This  suggests  that  idoteids,  particularly 
intertidal  species,  are  probably  not  good  dispersers. 

Because  the  following  discussion  deals  with  the  Idoteinae  at  the  generic  level,  it 
must  remain  fairly  general.  Even  at  this  level,  however,  these  generalizations  describe 
patterns  of  geographic  distribution  that  can  be  correlated  with  the  cladogram  (Fig.  4), 


1 1: 


Figure  5.     Distribution  of  valviferan  families  (excluding  Idoteidae):  Amesopodidae  [O],  Arcturidae  [•], 
Holognathidae  [D],  Pseudidotheidae  [O],  Xenarcturidae  [A]. 


and  which  can  most  parsimoniously  be  ascribed  either  to  vicariance  or  non-vicariance 
events.  Geological  dates  of  events  described  in  this  section  have  been  used  to  transform 
the  cladogram  into  the  evolutionary  tree  depicted  in  Fig.  16.  It  is  not  my  purpose  to 
present  a  detailed  analysis  of  each  genus  here;  that  must  await  a  species-by-species 
study  of  each  genus  (e.g.,  see  Brusca  1983  for  the  genus  Colidotea).  The  latter  will 
clearly  be  an  enormous  undertaking,  but  one  for  which  the  following  analysis  could 
provide  a  starting  point. 

The  Valvifera  as  a  whole  show  strong  correlation  to  southern  temperate  latitudes, 
3  of  the  6  families  being  restricted  to  that  geographic  region,  1  known  only  from  the 
shores  of  India,  and  the  other  2  being  widespread  (Fig.  5).  The  idoteine  genera  show 
a  similar  geographic  trend  (Figs.  7-14).  Numerous  studies  have  recently  appeared  that 
examine  the  relationships  of  distributional  patterns  of  terrestrial  flora  and  fauna  to  the 
geological  history  of  the  Southern  Hemisphere  (e.g.,  Keast  1973,  Raven  and  Axelrod 
1972,  Cracraft  1974,  1980).  However,  similar  studies  on  marine  groups  have  been  few. 
For  these  reasons,  Gondwanan  shores  are  discussed  in  some  detail.  A  brief  review  of 
the  breakup  of  Gondwana  follows,  based  on  Kennett  (1977),  Smith  and  Briden  (1977), 
Norton  and  Sclater  (1979),  Knox  (1979),  Durham  (1979),  Grant-Mackie  (1979)  and 
Hallam  (1981). 

Throughout  most  of  the  Permian,  the  Triassic,  and  most  of  the  Jurassic  periods, 
Gondwana  was  unified  as  a  single  land  mass.  The  proto-southern  continents  were  all 
situated  at  higher  latitudes  than  they  are  today.  Although  Permian  glaciation  probably 
existed  in  the  highest  southern  latitudes,  that  cold  era  came  to  a  fairly  abrupt  halt  by 
the  Jurassic.  By  mid-Mesozoic  times  climates  in  the  Southern  Hemisphere  had  changed 
markedly,  as  a  long-lasting  global  warming  trend  became  established.  Paleontological 
evidence  suggests  that  by  the  middle  or  late  Jurassic  the  northern  shores  of  Gondwana 
had  already  begun  acquiring  a  warm-water  Tethyian  biota,  thus  restricting  temperate 
coastal  biota  to  the  southernmost  latitudes.  A  common  temperate  marine  fauna  prob- 
ably inhabited  the  contiguous  coastline  of  Gondwana  south  of  55°-65°S  latitude.  This 
shallow  coastal  sea  washed  the  shores  of  southernmost  South  America,  southeast  Africa, 
eastern  Antarctica  and  eastern  Australia.  The  continuous  coastal  topography,  coupled 
with  non-glacial  and  generally  equable  climates  of  the  Jurassic,  facilitated  *'warm-water 
cosmopolitanism"  along  the  shores  of  northern  Gondwana.  and  "cold-water  cosmo- 
politanism" along  the  shores  of  southern  Gondwana.  These  two  regions  were  probably 
physically  isolated  from  one  another  until  at  least  the  early  Tertiary,  when  separation 
of  Australia  from  Antarctica  instituted  a  direct  high  latitude  southern  connection  be- 
tween the  Indian  and  Pacific  oceans.  The  distribution  of  modern  coastal  temperate 


113 


faunas  in  the  Southern  Hemisphere  can  thus  be  hypothesized  to  be.  at  least  in  part, 
the  product  of  the  fragmentation  of  an  early  to  mid-Mesozoic  temperate  Gondwanan 
track. 

That  part  of  Gondwana  composed  of  Antarctica,  Australia,  New  Zealand  and  South 
America  rotated  southward  during  the  Mesozoic,  although  as  early  as  the  Late  Jurassic 
these  land  masses  were  beginning  to  separate  as  Antarctica-Australia-India  began  un- 
coupling from  Africa  and  South  America.  By  the  Early  Cretaceous  (about  120  MY  A) 
sea  floor  spreading  had  started  to  form  the  south  Atlantic  Ocean.  At  about  the  same 
time,  India  began  to  decouple  from  Antarctica-Australia-Africa.  Marine  conditions 
developed  between  India  and  Antarctica- Australia  by  105  MY  A.  Africa  was  clearly 
separated  from  Gondwana  about  90  MYA.  New  Zealand  split  from  Antarctica-Aus- 
tralia 70-80  MYA.  By  the  mid-Cretaceous  the  south  Atlantic  Ocean  was  open  along 
its  entire  length  as  Africa  and  South  America  pulled  away  from  one  another.  The  free 
ocean  connection  (the  "Vema  Gap")  between  the  north  and  south  Atlantic  was  estab- 
lished by  the  Late  Cretaceous  as  the  transverse  Rio  Grande-Walvis  Ridge  sank  below 
the  1  km  depth  (approximately  78-80  MYA).  Australia  was  last  to  decouple  from 
Antarctica,  about  50-60  MYA.  During  the  Paleocene  these  land  masses  were  probably 
separated  by  a  shallow  narrow  seaway;  however,  the  South  Tasman  Rise  acted  as  an 
effective  barrier  to  the  development  of  a  circum-Antarctic  current  until  about  30-50 
MYA,  subsequent  to  which  the  southern  circum-polar  current  began  to  develop.  The 
modern  deep-flowing  Antarctic  Circumpolar  Current  could  not  have  been  established 
until  the  opening  of  the  Drake  Passage,  22-28  MYA,  when  deep-sea  conditions  de- 
veloped between  southern  South  America  and  Antarctica  (plate  boundaries  and  timing 
of  geotectonic  events  in  this  region  are  still  very  controversial). 

During  most  of  the  course  of  events  described  above,  Antarctica  continued  a  slow 
drift  southwards,  reaching  a  position  close  to  its  present  location  by  Late  Cretaceous: 
it  has  remained  nearly  stationary  throughout  the  Cenozoic.  Thus,  deep  marine  con- 
ditions began  to  develop  in  the  Southern  Ocean  by  the  Late  Paleocene,  although  a 
corridor  of  scattered  highlands  and  shallow  seas  persisted  between  Australia  and  Ant- 
arctica until  about  the  Late  Eocene  or  Early  Oligocene. 

South  America  has  moved  progressively  westward  since  the  Late  Cretaceous.  India 
collided  with  Asia  about  this  same  time.  Coincidental  with  this  southern  fragmentation, 
the  highest  post-Carboniferous  sea  levels  ever  to  occur  (Campanion  Era,  about  75 
MYA)  submerged  roughly  'A  of  the  present  continental  area  below  shallow  epiconti- 
nental seas.  Deep  water  flows  from  the  Pacific  into  the  Atlantic  across  Central  America 
commenced  about  55  MYA,  only  to  be  halted  in  the  Pliocene  when  the  Panama  Isthmus 
emerged. 

The  shallow-water  marine  fauna  would  not  have  responded  to  the  breakup  of 
Gondwana  in  the  same  manner  as  the  terrestrial  fauna,  because  the  initial  separation 
of  the  land  masses  created  intervening  shallow  seaways  that  fostered  the  spread  of 
marine  biota  before  separation  of  the  biotas  occurred  (for  a  comparison  with  the 
fragmentation  of  the  terrestrial  fauna  of  Gondwana  see  Cracraft  1 974  and  Rosen  1 978). 
As  South  America  and  Africa  moved  northward,  and  Antarctica  moved  southward, 
relative  to  one  another,  the  effect  on  temperate  coastal  marine  life  was  most  likely  to 
have  created  two  disjunct  faunas.  One  of  these  faunas,  the  western  coastal  fauna,  would 
have  inhabited  the  shores  of  southern  South  America,  southern  Africa,  and  north- 
western Antarctica.  The  eastern  fauna  would  have  been  restricted  to  Australia,  eastern 
Antarctica  and  southernmost  India.  New  Zealand,  at  this  time,  was  situated  in  warm- 
water  latitudes.  By  the  mid-Cretaceous,  the  Southern  Ocean  had  become  extensive 
enough  to  break  the  western  fauna  into  two  separate  temperate  regions:  southern  South 
America  (and  probably  the  region  of  the  Antarctic  Peninsula)  and  southern  Africa.  The 
eastern  Gondwana.  mid-Cretaceous,  temperate  coastal  region  remained  unchanged 
even  though  New  Zealand  had  split  from  the  warm-water  shores  of  Australia.  By  the 
mid-  to  Late  Cretaceous  the  shallow  sea  between  India  and  Antarctica  had  deepened 
to  isolate  these  regions  entirely  from  one  another.  India  having  drifted  into  the  tropical 
waters  of  the  Indian  Ocean.  The  coastal  environment  of  southern  Australia  remained 


114 


S' 

"&         ;f 

*     A® 

/ 

/ 

OLIGOCENE 


EOCENE 


PALEOCENE 


LATE  CRETACEOUS 


So.  America 


s.  Australia, 


e.   Antarctica 
l^    India 


MID  CRETACEOUS 


7 


EARLY  CRETACEOUS 


So.   America,  Africa, 
nw.  Antarctica 


s.   Australia,  New  Zealand, 
e.   Antarctica,   India 


JURASSIC 


Figure  6.  Proposed  geographic  cladogram  of  temperate,  shallow-water,  Gondwanan  coastal  areas.  Note 
the  significant  differences  between  this  cladogram  of  shallow  continental  shores  (marine  geographic  areas) 
and  Rosen's  (1978)  cladogram  of  Gondwanan  continents  (terrestrial  geographic  areas).  Seelexl  for  discussion. 


tied  to  northern  Antarctica  until  the  Cretaceous-Tertiary  boundary,  or  perhaps  even 
to  the  Early  Oligocene.  These  relationships  are  pictured  in  Fig.  6. 

Because  paleoclimates  and  oceanographic  conditions  were  not  the  same  during  the 
Mesozoic  and  Cenozoic  as  they  are  today,  latitudinal  distributions  of  shallow-water 
marine  life  would  have  been  under  quite  different  thermal  regimes.  This  ecological 
component  must  be  taken  into  account  when  considering  paleobiogeographical  recon- 
structions. A  brief  review  of  Mesozoic-Cenozoic  climates  follows;  for  more  detailed 
information  consult  Fleming  (1975,  1979),  Margolis  et  al.  (1978),  Grant-Mackie  ( 1 979). 
Durham  (1979),  Frakes  (1979),  Zinsmeister  (1982),  and  references  therein. 

Overall,  middle  and  late  Mesozoic  climates  were  extremely  equable,  with  ocean 
temperatures  exceeding  those  of  the  present  day  by  7°-15°C.  Polar  ice  caps  did  not 


115 


Figure  7.  Composite  distribution  map  of  all  idoteine  genera,  except  the  3  cosmopolitan  taxa  (Idoiea. 
Synidotea,  Zenobiana).  Dashed  lines  indicate  limits  of  warm-water  (tropical/subtropical)  regions;  open  cir- 
cles =  genera  of  lineage  A;  closed  boxes  =  genera  of  lineage  B. 


exist  and  both  terrestrial  and  marine  biotas  exhibited  little  evidence  of  latitudinal 
zonation.  Cosmopolitanism  was  common.  Cool  sea  water  existed  in  the  southern  oceans 
only  south  of  about  50°S  latitude.  Early  Cretaceous  sea  bottom  temperatures  were 
approximately  10°-15°C  warmer  than  today;  Late  Cretaceous  sea  bottom  temperatures 
were  approximately  7°C  warmer  than  today.  The  Eocene  marked  the  beginning  of  a 
global  cooling  trend  that  ultimately  led  to  the  Cenozoic  glaciations  and  cooling  of  deep 
ocean  bottom  waters  to  their  present  thermal  regimes  (Shackleton  and  Kennett  1975). 
As  late  as  the  Eocene  (38-55  MYA),  all  but  the  southernmost  Antarctic  seas  were 
warm-temperate  in  nature.  The  steep  thermal  drop  over  the  Eocene/Oligocene  boundary 
was  probably  related  to  the  establishment  of  the  Antarctic  Circumpolar  Current  and 
subsequent  South  Polar  glaciation.  Although  land  masses  occupied  both  poles  by  the 
Early  Paleocene,  large-scale  Antarctic  glaciations  probably  did  not  begin  until  the  Mio- 
cene. The  Antarctic  Convergence  began  moving  northward  in  the  Oligocene.  Cooling 
trends  continued  throughout  the  Cenozoic,  apparently  marked  by  several  periods  of 
severe  chilling. 

Fossil  reefs,  requiring  relatively  warm  surface  waters,  grew  to  paleolatitudes  of 
about  40°  throughout  the  Paleogene,  as  in  the  Cretaceous.  In  the  northeast  Pacific, 
tropical  environments  (i.e.,  surface  temperatures  >20°C)  extended  to  approximately 
45°N  in  the  Eocene,  and  have  contracted  equatorward  steadily  since  that  time.  Abun- 
dant paleoceanographic  and  stratigraphic  data  exist  to  indicate  that  New  Zealand  shores 
were  bathed  primarily  by  warm  subtropical  waters  throughout  the  Cretaceous  and 
Paleocene  (Fleming  1962,  1975,  Durham  1979,  Knox  1979),  although  a  distinct  cooling 
trend  began  in  the  middle  to  Late  Miocene,  eventually  resulting  in  present-day  tem- 
perate coastal  conditions.  At  best,  only  southernmost  New  Zealand  might  have  expe- 
rienced temperate  waters  from  the  Eocene  to  the  Miocene.  By  early  Pleistocene  sub- 
antarctic  waters  had  reached  about  40°S  latitude  on  New  Zealand  shores.  The  middle 
Oligocene  marked  the  end  of  "'high  latitude  subtropical  communities,"  and  by  the  Late 
Miocene  modern  water  bodies  and  their  associated  biological  provinces  had  begun  to 
be  established.  The  Australian  Subantarctic  water  mass  had  formed  by  about  1  5  MYA. 
For  the  past  10-15  million  years  the  Southern  Ocean  temperature,  circulation  and 
zonational  water  masses  have  remained  essentially  the  same. 

If  the  distributions  for  all  species  of  Idoteinae  are  plotted  on  the  globe,  nearly  every 
sea  and  shore  is  seen  to  be  inhabited  by  one  or  more  genera.  Fig.  7  is  a  composite 
distribution  map  plotting  the  ranges  of  all  taxa  of  Idoteinae  EXCEPT  the  3  cosmopolitan 
gQnQV2i  —  Idotea,  Synidotea.  and  Zenobiana.  These  3  taxa  are  omitted  from  the  com- 


116 


Figure  8.     Distributions  of  Cleantiella  [•],  Lyidotea  [•],  and  Cleantioides  [O]. 


posite  distribution  map  because  they  provide  no  particular  insights  into  a  global  pattern 
analysis  at  this  level.  Distribution  maps  of  the  individual  genera  are  given  in  Figs.  8- 
14.  and  are  discussed  below.  It  will  be  seen  from  these  distribution  maps  and  the 
following  discussion  that  most  idoteine  genera  are  endemic  to  only  one  or  a  few  coastal 
regions.  The  dashed  lines  on  Fig.  7  indicate  the  limits  of  the  tropical/subtropical  regions, 
based  on  Ekman  (1953),  Briggs  (1974),  and  Brusca  and  Wallerstein  (1979/)).  While 
some  disagreement  exists  regarding  the  exact  limits  of  the  tropical  zones,  the  latitudes 
that  I  have  chosen  clearly  separate  the  "warm-water"  faunal  regions  of  the  world's 
shores  from  the  "cold-water"  regions.  The  only  serious  arguments  with  these  delimi- 
tations might  be  in  whether  one  regards  the  Mediterranean  Sea  as  warm-temperate  or 
subtropical.  The  best  comprehensive  discussion  of  Mediterranean  zoogeography  is 
probably  that  of  Ekman  (1953),  who  in  describing  the  complex  mixtures  of  northern 
and  southern  elements,  couldn't  comfortably  label  this  sea  either  warm-temperate  or 
subtropical.  The  nature  of  south  African  faunal  designations  has  also  been  a  matter  of 
some  controversy,  and  was  recently  reviewed  by  Brown  and  Jarman  (1978). 

Several  general  patterns  are  revealed  when  Fig.  7  is  examined.  In  the  New  World, 
endemic  idoteine  genera  are  distributed  fairly  equally  in  both  warm  and  cold  waters. 
In  the  Old  World,  however,  there  is  a  marked  absence  of  records  from  the  tropics. 
Only  3  genera  on  this  map  represent  Old  World  tropical  taxa:  Cleantiella  and  Clean- 
tioides both  occur  on  the  east  Asian  coast,  while  Lyidotea  is  known  from  a  single  species 
in  northeastern  Australia  (see  Fig.  8).  A  second  pattern  seen  is  that  Old  World  endemic 
genera  tend  to  form  3  distinct  clusters,  as  follows:  (1)  a  Southern  Hemisphere  cold- 
water  group,  (2)  a  European  cool-  to  cold-water  group,  and  (3)  a  northeast  Asian  cool- 
to  warm-water  group.  Only  2  of  the  genera  depicted  in  Fig.  7  occur  in  both  the  Old 
and  New  Worlds:  the  southern  temperate  Paridotea  and  the  northern  tropical  Clean- 
tioides. Because  most  species  and  genera  of  Idoteidae  are  restricted  to  temperate  or 
polar  seas,  the  family  has  long  been  considered  a  "cold-water  centered  taxon"  (see 
Brusca  and  Wallerstein,  1919b  and  references  therein).  It  is  of  particular  interest  that 
the  Old  World  and  New  World  tropical  regions  are  inhabited  by  entirely  different 
genera.  The  New  World  tropical  genera  are  Cleantioides.  Erichsonella,  Parasyninierus, 
and  Eusymmerus.  The  Old  World  tropical  genera  are  Cleantiella,  Lyidotea,  Idotea, 
Synidotea,  and  Zenohiana.  As  the  latter  3  are  largely  cosmopolitan  taxa,  the  obvious 
question  is,  "Why  have  species  in  these  genera  been  unable  to  invade  the  tropics  in 
the  New  World?"  Let  us  now  examine  the  distributions  of  the  individual  idoteine 
genera. 

Idotea  is  nearly  a  cosmopolitan  genus  (Fig.  9).  It  occurs  in  most  waters  of  the  Old 
World,  including  the  tropics,  but  in  the  New  World  it  is  notably  absent  from  the  warm 
seas  between  the  Tropics  of  Cancer  and  Capricorn  in  both  the  Pacific  and  Atlantic. 


17 


Figure  9.     Distribution  of  Idotea. 

Brusca  and  Wallerstein  ( 1 919b)  reviewed  literature  documenting  the  eurythermal  nature 
o{  Idotea  species,  and  suggested  that  biological  factors  (competition  or  predation),  rather 
than  temperature,  could  be  excluding  the  2  New  World  temperate  isopod  genera,  Idotea 
and  Synidotea,  from  the  Western  Hemisphere  tropics.  Wallerstein  and  Brusca  (1982) 
subsequently  provided  experimental  and  comparative  morphological  evidence  that 
predation  by  crustacivorous  fishes  is  possibly  restricting  species  o^  Idotea  from  the  New 
World  tropics.  In  the  latter  study  they  proposed  a  model  that  envisioned  "faunal  waves" 
of  tropical  predator  species  (primarily  fishes)  moving  northward  from  the  equatorial 
region  during  warm  Pleistocene  interglacials,  pushing  the  southern  latitudinal  range 
end  points  of  non-adaptable  temperate  species  northward  ahead  of  them,  and  thus 
excluding  Idotea  and  Synidotea  from  the  New  World  equatorial  region  (see  Addicott 
1 970  and  Zinsmeister  1 974  for  a  discussion  of  similar  north-south  faunal  displacements 
in  Mollusca).  The  nearly  global  distribution  of  Idotea,  as  well  as  its  position  on  the 
cladogram  (Fig.  4)  relative  to  other  genera  in  lineage  B  (see  discussion  below)  suggests 
that  Idotea  is  an  old  genus  (pre-Cretaceous).  It  should  be  emphasized  that,  while  the 
genus  Idotea  is  cosmopolitan,  the  species  in  this  taxon  are  themselves  largely  endemic 
to  restricted  coastlines.  Only  one  species  of  Idotea  is  cosmopolitan,  /.  nieta/llca.  Its 
cosmopolitanism  has  been  explained  by  drift  dispersal  of  the  algae  on  which  it  lives 
(Naylor  1972,  Poore  1981). 

Nine  of  the  12  remaining  genera  in  lineage  B  are  endemic  to  the  Southern  Hemi- 
sphere (Table  2)  and  are  probably  descendants  of  a  pan-austral,  cold-water,  Gondwanan. 
pre-Cretaceous  fauna.  The  concept  of  a  pan-austral  terrestrial  biota  was  apparently  first 
recognized  by  Hooker  (1853,  1860)  and  Huxley  (1868).  Though  largely  suppressed  by 
100  years  of  "Wallaceian  dispersalism,"  the  concept  has  finally  experienced  a  rebirth 
owing  largely  to  the  work  of  Pantin  et  al.  (1960)  and  Brundin  (1966.  1970.  1972<:/.  /). 
1976).  For  a  recent  summary  of  many  subsequent  papers  see  N.Z.  DSIR  (1979).  Only 
recently,  however,  has  serious  documentation  and  discussion  of  generalized  temperate 
pan-austral  marine  tracks  begun  (e.g.,  Zinsmeister  1976,  1982,  Fleming  1975.  Knox 
1975).  The  mid-Cretaceous/early  Tertiary  fauna  that  inhabited  this  temperate,  shallow- 


Table  2.     Distribution  of  the  8  Idoteinae  Genera  of  Lineage  B  Endemic  to  the  Southern  Hemisphere. 


Geographic 

Pah- 

Eiii- 

Craliv- 

Svn- 

Glypti- 

Barnardi- 

Engi- 

occurrence 

Moplisa 

doiea 

dotea 

zos 

ischia 

dotea 

dotea 

doica 

Southern  South  America 

+ 

+ 

South  Africa 

+ 

+ 

+ 

+ 

+ 

Southern  AustraHa 

+ 

+ 

+ 

+ 

New  Zealand 

— 

■~ 

+ 

118 


Figure  10.     Distributions  of  Glyptidotea  and  Barnardidotea  [•],  Engidotea  [•].  Pentias  [A],  Zenobianopsis 
[D],  and  Paridolea  [O]. 


water,  southern  Gondwanan  region  has  been  referred  to  as  the  WeddelHan  Province 
by  Zinsmeister  (1976). 

One  of  the  eadiest  appearing  genera  in  this  posX-Idotea  radiation  was  Synidotea, 
which  like  Idotea  is  nearly  cosmopolitan  but  notably  absent  from  the  New  World 
tropics  (Fig.  1 1 ).  The  same  comments  that  apply  to  Idotea  above  are  probably  applicable 
to  Synidotea.  The  origins  of  Synidotea  appear  old  enough  that  the  Early  Cretaceous 
circum-Arctic  seaway  probably  served  as  one  of  several  principal  dispersal  routes,  this 
being  reflected  in  the  modern  distribution  of  the  genus,  which  has  by  far  the  majority 
of  its  species  concentrated  in  the  Northern  Hemisphere.  This  hypothesized  age  of  origin, 
plus  the  restriction  of  both  Barnardidotea  and  Mop/isa  (Synidotea^s  sister-group)  to 
the  Southern  Hemisphere,  argues  strongly  for  the  origin  of  Synidotea  in  the  southern 
seas.  This  contrasts  with  the  opinion  of  Gurjanova  (1935)  and  Menzies  and  Miller 
(1972),  who  suggested  that  the  place  of  origin  of  Synidotea  was  the  north  Pacific. 
Menzies  and  Miller's  opinion  appears  to  have  been  based  solely  on  the  fact  that  most 
species  of  Synidotea  presently  occur  in  this  area,  which  they  considered  to  be  its  "center 
of  origin."  The  present  analysis,  however,  does  corroborate  Menzies  and  Miller's  (op. 
cit.)  dating  of  the  origin  of  this  genus.  Synidotea's  sister-group,  Moplisa,  consists  of  a 


FiGURt  11.     Distribution  of  .S'lYj/^o/f'a. 


119 


Figure  12.     Distribution  of  Crabyzos  [•],  Synischia  [•],  Ewdotea  [D],  Synisoma  [O],  Colidotea  [A],  and 
Moplisa  [O]. 


single  species.  M.  sphaewmiformis  (Mane-Garzon.  1946).  so  far  known  only  from  a 
short  stretch  of  coastline  in  southern  Brazil  (Fig.  1 2).  The  cosmopolitanism  ofSynidoiea 
(vs.  the  restricted  distribution  of  the  monotypic  Moplisa).  and  the  absence  of  an  apo- 
morphy  to  distinguish  Synidotea  from  Moplisa  strongly  suggests  that  Moplisa  evolved 
either  in  sympatry  or  as  a  peripheral  isolate  from  a  continuing  stock  of  Synidotea. 
These  relationships  are  pictured  in  the  phylogram  (Fig.  16)  and  are  consistent  with  the 
cladogram. 

Although  Glyptidotea,  Crabyzos,  Pentias,  and  Synischia  cannot  be  separated  by 
cladistic  analysis  (Fig.  3),  the  biogeographic  data  provide  evidence  regarding  their 
origins.  Glyptidotea  is  endemic  to  temperate  South  Africa  (Fig.  10);  Crabyzos  and 
Synischia  are  endemic  to  temperate  Australasia  (Fig.  12).  These  patterns  suggest  that 
these  3  genera  arose  subsequent  to  the  initial  east-west  split  of  Gondwana  (i.e..  post- 
Jurassic).  The  absence  o{  Glyptidotea  from  the  South  American  component  of  the  west 
Gondwanan  track  suggests  that  this  genus  arose  subsequent  to  the  separation  of  these 
two  land  masses  (i.e..  mid-Cretaceous  or  later).  The  closely  related  genus  Pentias  is 
today  restricted  to  temperate  shores  of  northern  Japan  (Fig.  10).  The  simplest  expla- 
nation for  its  occurrence  would  seem  to  be  fortuitous  jump  dispersal  across  the  warm 


Figure  13.     Distribution  of  Ze«oWa«a. 


120 


Figure  14.     Distribution  of  Eusymmerus  and  Parasymmerus  [•],  Edotea  [O],  and  Erichsonella  [•]. 


waters  of  the  equatorial  region,  perhaps  during  a  period  of  global  cooling  and  tropical 
compression  such  as  the  late  Tertiary  (although  the  western  Pacific  was  probably  far 
less  affected  by  marine  cooling  than  other  regions  on  the  globe).  A  second,  remote 
possibility  is  that  it  (or  its  ancestors)  reached  Japan  via  China,  which  may  have  originally 
been  part  of  the  Gondwanan  continent  (see  Crawford  1974). 

None  of  the  above  southern  taxa  occur  along  the  warmer  shores  of  the  Indian 
Ocean.  Their  distributions  suggest  that  these  genera  are  Cretaceous  to  post-Cretaceous 
descendants  of  the  widespread  Jurassic-Early  Cretaceous  temperate  Gondwanan  track 
discussed  earlier  in  this  section.  The  ancestor(s)  inhabiting  this  track  is  indicated  in  the 
evolutionary  tree  by  "GonAnc"  (Fig.  16). 

Because  several  genera  cannot  be  clearly  distinguished  from  one  another  by  the 
cladogram,  it  is  difficult  to  seek  congruence  between  lineage  B  and  the  geographic 
cladogram  of  Fig.  6.  However,  a  partial  correlation  (38%)  is  nonetheless  evident  (Fig. 
\5a,  b).  Table  2  provides  a  distributional  summary  of  the  8  Idoteinae  genera  of  lineage 
B  that  are  Southern  Hemisphere  endemics.  Fig.  15  gives  reduced  taxon-area  and  geo- 
graphic cladograms  for  the  groups  in  question.  The  cladogram  congruence  with  the 
Synidotea-Baniardidotea-Moplisa  line  corroborates  the  hypothesis  that  the  latter  two 
taxa  arose  subsequent  to  the  splitting  of  South  America  from  Africa,  while  Synidotea 
continued  to  persist  unchanged.  Correlation  also  exists  for  Synischia.  If  one  accepts 
the  probable  dispersal  of  Crahyzos  from  Australia  to  New  Zealand,  the  correlation 
between  these  cladograms  is  raised  to  50%.  Paridotea  has  retained  its  circumpolar 
distribution,  corroborating  its  early  (pre-Cretaceous)  origin. 

The-occurrence  of  Euidotea  on  African,  Australian,  and  New  Zealand  shores  (Fig. 
12),  and  its  absence  from  South  American  coasts  today,  argues  for  either  (1)  its  origin 
prior  to  the  break-off  of  Africa  and  its  subsequent  extinction  in  South  America,  or  (2) 
its  origin  on  African  shores  after  the  break-off  of  that  continent  and  its  subsequent 
spread  to  Australasia  via  West  Wind  Drift.  Recall  that  New  Zealand  probably  did  not 
achieve  its  present  temperate  coastal  thermal  regime  until  well  after  the  deep  water 
barriers  were  formed  that  isolated  it  from  Australia,  probably  not  until  the  Eocene  at 
the  earliest.  Thus,  introduction  of  the  cold-water  genus  Crahyzos  could  have  been  via 
dispersal  from  Australia  in  fairly  recent  times  (mid-  to  late  Tertiary).  This  same  rea- 
soning must  apply  to  the  other  temperate  New  Zealand  genera,  Paridotea  and  Euidotea. 
The  fact  that  none  of  these  three  taxa  are  endemic  to  New  Zealand  (all  3  also  occur  in 
southern  Australia)  lends  support  to  this  probability. 

The  sister-groups  Colidotea  and  Synisoma  are  restricted  to  warm-temperate  and 
subtropical  waters  of  the  New  and  Old  World  respectively  (Fig.  12).  This  restriction, 
plus  the  widespread  occurrence  of  Synisoma  (Europe,  Far  East),  suggests  that  these 


,21 


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Figure  1 5a.  Reduced  geographic  cladogram  of  4  extant  temperate  shores  derived  from  Gondwana  {see  Fig. 
6).  15b.  Reduced  taxon-area  cladogram  of  Southern  Hemisphere  endemic  idoteine  genera  (see  Fig.  3  and 
Table  2). 


taxa  are  vicariant  descendants  of  a  widespread,  circumglobal.  northern  Tethyian  track. 
The  absence  of  either  of  these  genera  on  eastern  American  shores  is  most  easily  ex- 
plained by  local  extinction  on  that  coast.  Closure  of  the  Panama  seaway  in  the  Pliocene, 
or  simple  isolation  from  the  parental  stock  as  the  North  Atlantic  opened  up.  could 
have  provided  the  setting  for  the  origin  of  these  two  taxa  as  we  know  them  today. 
Judging  by  its  extreme  disjunct  distribution.  Synisoma  has  remained  unchanged  since 
at  least  the  Paleocene,  when  the  Gibralter  gap  closed.  Thus,  it  appears  that  the  ancestors 
of  these  two  taxa  were  members  of  the  Tethyian  fauna,  at  least  marginally,  while  the 
extant  descendants  have  been  pushed  northward  to  the  subtropical  and  warm -temperate 
portions  of  their  former  range. 

In  summary,  the  genera  comprising  lineage  B  appear  to  have  remained  closely  tied 
to  their  ancestral  (Jurassic)  temperate  Gondwanan  distribution.  Historical  hypotheses 
that  are  consistent  with  the  cladogram  suggest  that  vicariancc  alone  can  account  for 
the  modern  distributional  patterns  of  only  a  few  taxa.  most  haxing  modern  distributions 
clearly  tied  to  both  former  vicariant  and  dispersal/extinction  events.  These  probable 
extinctions,  dispersal  events,  and  biological  interactions  affecting  ranges  add  to  the 
uncertainty  in  ascribing  geographic  origins  to  the  relatively  old  taxa  comprising  lin- 
eage B. 

Turning  to  lineage  A.  we  see  one  broadly  distributed  genus,  the  primitive  Zcno- 
biana.  which  occurs  in  both  temperate  and  tropical  waters  of  the  Old  World,  but  is 
restricted  to  temperate  South  America  in  the  New  World  (Fig.  13).  Zenobiana  is  the 
only  genus  of  Idoteinae  to  retain  the  primitive  biarticulatc  uropods.  .As  was  the  case 
with  Idotea,  Zenobiana's  cladistic  relationship  and  widespread  distribution  initially 


122 


"9        5; 


6       5; 


i      a      5      i 


Figure  16. 
discussion. 


Proposed  phylogeny  of  the  genera  of  Idoteinae,  based  on  Fig.  1 7  and  other  data.  See  text  for 


suggests  that  Zenobiana  was  pre-  to  Early  Cretaceous  in  origin  (i.e..  pre-Gondwanan 
fragmentation).  Zenobiana  cannot  be  much  older  than  this  as  it  clearly  shows  no  alliance 
to  the  Permo-Triassic  Tethyian  Sea  radiation,  owing  to  its  near  absence  from  the  tropical 
Indo-West  Pacific  region;  only  one  species  occurs  in  Indo-Pacific  waters.  Z.  nata/ensis 
(Barnard  1925).  Thus.  Zenobiana's  limited  invasion  of  tropical  environments  appar- 
ently took  place  after  the  final  closure  of  the  Tethyian  Sea.  in  the  early  to  mid-Tertiary. 
Its  alliance  to  and  probable  origin  on  temperate  shores  is  further  indicated  by  its  present 
distributional  pattern.  All  genera  that  appeared  subsequent  to  the  establishment  of  the 
globally  distributed  Zenobiana  (the  7  remaining  genera  in  lineage  A)  are  warm-water 
taxa,  or  at  least  probably  initially  evolved  as  warm-water  taxa  (i.e..  Edotea).  The  absence 
of  identifiable  apomorphies  to  distinguish  Zenobiana  indicates  that  this  genus  was  both 
the  ancestor  to,  and  persisted  after  the  origin  of  its  sister-group. 

Cleantioides  consists  of  only  2  species:  C.  occidentalis  is  endemic  to  the  tropical 
eastern  Pacific;  C  planicauda  is  a  widespread  tropical  species  known  from  both  coasts 
of  the  Americas  and  from  eastern  Asia.  Cleantiella  and  Lyidotea  are  western  Pacific 
descendants  of  a  Cleantioides-Mke  ancestor.  Cleantiella  is  restricted  to  the  warm  shores 
of  USSR  and  China,  and  most  likely  arose  as  a  post-Cretaceous  northwest  Pacific 
endemic  (Fig.  8).  Lyidotea  is  restricted  to  the  warm  waters  of  northeastern  Australia. 
The  absence  of  Cleantiella  and  Lyidotea,  both  warm-water  taxa,  from  the  equatorial 
region  of  the  west  Pacific  is  enigmatic  and  suggests  3  possibilities:  (1)  dispersal  of  the 
ancestor  of  Lyidotea  from  Asian  shores  to  Australian  shores;  (2)  the  former  existence 
of  a  N-S  tropical  Cleantiella-like  track  joining  these  two  regions,  with  subsequent 
fragmentation  into  northern  and  southern  tropical  isolates  with  extirpation  in  the  equa- 
torial region;  or  (3)  an  artifact  of  collection  records.  The  paucity  of  records  for  species 
in  any  other  genus  of  Idoteidae  from  this  region,  and  the  unlikelihood  of  alternatives 
1  and  2  above,  suggest  the  third  possibility  may  be  the  correct  interpretation.  According 
to  Poore  (in  litt.)  virtually  no  collecting  has  ensued  in  this  region  since  the  Siboga 
Expedition  (ca.  1900). 


123 


All  remaining  genera  in  lineage  A  are  New  World  endemics  and  represent  descen- 
dants of  the  post-Cretaceous  ancestors  of  this  line.  Edotca  has  an  anti-tropical  distri- 
bution, occurring  on  both  North  and  South  American  shores  (Fig.  14).  Brusca  and 
Wallerstein  (1979a)  have  discussed  this  unusual  distribution,  concluding  that  Edotea, 
although  evolving  in  the  New  World  tropics  as  part  of  the  early  transisthmian  track 
biota  (the  "Tertiary  Caribbean  Province"  of  Woodring  1954,  1966;  the  "Panamanian 
Track"  of  Croi/.at  et  al.  1974),  was  promptly  excluded  from  this  warm-water  region 
by  competition  with  the  better  adapted  tropical  genera  like  Clcantioides,  Eusymmcrus, 
Parasymmenis,  and  Erichsonella.  Modern  species  of  Edotea  now  occur  only  in  tem- 
perate latitudes  to  the  north  and  south  of  the  New  World  tropics.  "Better  adapted" 
includes  the  possession  of  such  predator  avoidance  adaptations  as  small  body  size, 
early  reproduction,  and  cryptic  morphologies  (see  Wallerstein  and  Brusca  1982,  for 
details  of  these  adaptations). 

Edotea.  Erichsonella  (Fig.  14),  and  Cleantioides  are  all  components  of  the  Tertiary 
Caribbean  Province  (Woodring  1966)  that  have  retained  their  amphi-American  dis- 
tributions. Numerous  eastern  Pacific-western  Atlantic  analog  or  geminate  species  can 
be  identified  within  these  genera.  Eusymmenis  and  Parasymnwrus  {Fig.  14)  are  tropical 
eastern  Pacific  endemics.  Parasyninierus  probably  evolved  subsequent  to  the  Pliocene 
closure  of  the  Panama  seaway.  However,  if  the  relationships  depicted  on  the  cladogram 
are  correct,  Eusymmerus  probably  evolved  prior  to  the  closure  of  the  seaway  (in  order 
to  be  the  sister-group  or  ancestor  of  Edotea-Parasymmerus).  Its  present  restriction  to 
the  eastern  Pacific  implies  that  either  (1)  it  evolved  there  and  never  became  part  of  the 
transisthmian  fauna,  or  (2)  it  was  part  of  the  transisthmian  biota  but  has  subsequently 
become  extinct  in  the  western  Atlantic.  The  latter  seems  far  the  more  probable  sequence 
of  events.  Woodring  (1966)  has  pointed  out  that  many  molluscan  genera  presently 
restricted  to  the  tropical  eastern  Pacific  occurred  throughout  the  eastern  Pacific-western 
Atlantic  Tertiary  Caribbean  Province  during  Miocene  times.  These  geographic  data,  as 
well  as  the  cladistic  relationships,  imply  that  Edotea  was  both  the  ancestor  to  Para- 
symmerus  and  remained  essentially  unchanged  subsequent  to  the  origin  of  Parasym- 
merus  (Fig.  16).  It  is  unfortunate  that  fossil  material  for  isopods  is  rare,  for  such  data 
could  provide  excellent  corroboration  or  refutation  of  the  above  hypothesis.-^ 

The  data  summarized  in  the  cladogram  and  biogeographic  review  clearly  imply 
that  there  were  both  Old  World  and  New  World  warm-water  genera  derived  from  an 
ancestral  global  ^^Zenobiana-Cleantioides  line."  The  extant  Old  World  fragments  of 
this  line  are  Cleantiella  and  Lyidotea\  the  extant  New  World  fragments  are  Erichsonella, 
Edotea,  Parasymmerus,  and  Eusymmerus. 

The  relationships  in  the  cladogram  suggest  that  these  latter  4  New  World  endemic 
taxa  were  derived  as  the  sister-group  of  Lyidotea  (an  Old  World  endemic).  This  ne- 
cessitates trans-Pacific  (west  to  east)  dispersal  of  the  ancestor  of  these  4  New  World 
genera.  An  alternate  scenario  would  have  the  New  World  taxa  of  lineage  A  descended 
not  from  the  geographically  restricted  Lyidotea  line  or  its  Old  World  ancestor,  but  from 
the  widespread  ancestral  Zenobiana-Cleantioides  line.  This  could  be  accomplished 
simply  by  reconstructing  a  portion  of  lineage  A,  as  shown  in  Fig.  1  7.  This  new  cladogram 
of  lineage  A  is  longer  than  the  original  (Fig.4)  in  requiring  9  pleonal  transformations, 
versus  8  in  the  original.  The  new  cladogram  requires  four  transformations  (but  two 
convergences)  in  maxillipedal  palp  reduction,  as  in  the  original.  Thus,  for  lineage  A  we 
are  faced  with  a  cladogram  slightly  more  parsimonious  (one  less  step)  but  requiring  a 
major  successful  trans-Pacific  dispersal  event  to  establish  the  entire  New  World  tropical 
lineage  of  idoteine  genera,  versus  a  cladogram  slightly  less  parsimonious  but  requiring 
only  an  easily  explained  vicariant  origin  of  the  New  World  line  from  a  pre-drift  cos- 
mopolitan track.  This  is  perhaps  a  case  of  strict  methodological  parsimony  versus 
"biological  parsimony."  Given  the  multitude  of  avenues  for  pleonal  reduction.  I  see 
no  compelling  reason  to  accept  one  cladogram  over  the  other.  Such  problematic  re- 
lationships can  probably  be  resolved  only  by  the  elucidation  of  new  generic-level  at- 
tributes, and  by  careful  examination  of  the  morphological  and  biogeographic  relation- 
ships of  the  individual  species  in  these  genera. 


124 


Discussion 

The  above  phylogenetic  and  biogeographic  analysis  sheds  light  on  several  problems 
in  understanding  the  evolution  of  the  Idoteinae.  First  is  the  problem  of  discordant 
character  trends.  Although  trends  toward  fusion  of  body  somites  and  appendage  articles 
(pleon,  antennae,  maxillipedal  palp)  have  been  noted  for  many  decades,  it  was  not  until 
Menzies  and  Miller  (1972)  summarized  and  tabulated  these  data  that  the  discordant 
trends  in  these  different  morphological  features  were  recognized,  suggesting  probable 
high  levels  of  homoplasy  in  the  Idoteinae  (see  Table  1).  Generating  schematic  pattern 
analyses  (Figs.  4  and  17a)  reveals  that  identical  pleonal  morphologies  can  be  derived 
in  a  great  many  ways.  For  example,  previous  workers  have  assumed  the  0+1  pleonal 
morphology  of  Colidotea  to  have  been  derived  in  the  same  manner  as  the  0+ 1  pleonal 
morphology  of  Eusynimeriis,  Parasymnierus,  and  Edotea  (i.e.,  to  be  homologous), 
despite  the  fact  that  the  former  has  a  multiarticulate  second  antennal  flagellum  whereas 
the  latter  3  genera  have  uniarticulate  (clavate)  second  antennal  flagella,  the  latter  pre- 
sumably being  a  fundamentally  significant  difference.  In  fact,  the  0+1  pleonal  mor- 
phology can  be  derived  in  any  of  about  a  dozen  different  ways,  and  in  the  above  case 
it  is  clear  that  the  derivation  of  Colidotea  was  probably  quite  different  from  that  of  the 
other  3  genera  (see  Figs.  3,  4,  16  and  17).  Use  of  this  schematic  approach  to  pattern 
analysis  in  the  Idoteinae  also  facilitates  placement  of  the  primitive  genus  Zenohiana 
in  a  single  lineage  apart  from  Cleantioides,  even  though  both  taxa  have  identical  pleonal 
morphologies  and  have  been  confused  with  one  another  in  the  past.  Further,  the  sche- 
matic analysis  provides  clear  visual  descriptions  of  all  potential  intermediate  stage 
morphologies  for  the  Idoteinae  genera,  lending  high  predictability  (hence  testability) 
to  hypotheses  contained  in  the  above  analysis. 

That  isopods,  and  peracarids  in  general,  were  already  diverse  and  important  taxa 
by  the  late  Paleozoic  has  been  fairly  well  documented  by  Schram  (1970,  1974).  By  as 
early  as  the  Pennsylvanian,  the  Peracarida  had  radiated  into  most  if  not  all  its  currently 
recognized  orders  (except  perhaps  the  Amphipoda).  As  Schram  (1974)  stated,  ".  .  . 
peracarid  radiation  was  probably  initiated  in  Devonian  time,  when  it  is  generally 
thought  caridoid  eumalacostracans  were  taking  origin,"  ".  .  .  the  Peracarida  are  now 
known  to  be  among  the  most  ancient  of  the  eumalacostracans,"  and  ".  .  .  the  superorder 
Peracarida  was  a  major  element  in  the  late  Paleozoic  radiation  of  eumalacostracans 
and  contributed  the  principal  caridoid  types  of  that  time."  The  earliest  isopods  known 
from  the  fossil  record  are  phreatoicids  from  the  middle  Pennsylvanian;  spelaeogripha- 
cians,  tanaids,  and  mysidaceans  first  appear  in  the  lower  Carboniferous. 

The  origins  of  the  subfamilies  of  Idoteidae  hypothesized  in  this  study  are  consistent 
with  Schram's  data,  and  also  with  dates  offered  by  Schultz  (1979),  who  stated  that  most 
isopod  families  were  well-defined  before  the  Triassic. 

The  present  analysis  reveals  the  existence  of  two  principal  lines  of  descent  within 
the  Idoteinae  (Fig.  16),  lines  that  separated  from  one  another  early  on  (Jurassic  or 
perhaps  even  Triassic).  One  of  these  lines  remained  closely  tied  to  its  origins  in  the 
Old  World  Southern  Hemisphere  temperate  biota,  while  the  other  invaded  New  World 
environments  and  undertook  a  radiation  in  the  New  World  tropics  (producing  genera 
such  as  Cleantioides,  Erichsonel/a,  Eusymmerus,  and  Parasyiuinerus).  Brusca  and  Wal- 
lerstein  (1979/?)  hypothesized  that  idoteids  might  only  recently  have  invaded  the  New 
World  tropics,  perhaps  in  close  association  with  a  similar  invasion  of  these  regions  by 
the  brown  algae  (Phaeophyta),  which  probably  serve  as  the  principal  habitat  and  food 
resource  for  most  idoteine  species.  That  the  late  Triassic-Early  Jurassic  periods  might 
have  had  a  widespread  southern  temperate  Idoteinae  distribution  of  little  or  no  ende- 
mism  is  hardly  surprising.  As  Hallam  (1981)  points  out,  Jurassic  marine  faunas  as  a 
whole  were  decidedly  more  cosmopolitan  than  those  of  the  present  day.  These  early 
origins  in  temperate  waters  explain  the  long-held  belief  that  the  Idoteidae  and  Idoteinae 
are  "cold-water  centered  taxa."  Plate  movements  became  increasingly  influential  in 
promoting  endemism  within  the  Idoteinae  from  Late  Jurassic  through  Late  Cretaceous, 
as  Pangaea  fragmented.  The  high  degree  of  endemism  among  Idoteinae  genera  contrasts 


125 

sharply  with  claims  by  Taylor  and  Forester  (1979)  that  cold-water  psychrospheric 
isopod  biofacies  (generic  assemblages)  tend  to  be  similar  regardless  of  latitude  or  depth. 

This  study  has  shed  some  light  on  the  place  of  origin  of  both  the  Idoteinae  and 
the  Idoteidae,  as  well  as  the  Valvifera  itself.  Distributions  of  the  Idoteidae  subfamilies 
(excluding  the  Idoteinae)  are  given  in  Fig.  18;  distributions  of  the  five  other  valviferan 
families  are  given  in  Fig.  5.  As  can  be  seen,  two  of  the  five  Idoteidae  subfamilies  are 
distinctly  Gondwanan  in  distribution  (Glyptonotinae  and  Chaetilinae).  one  is  predom- 
intly  Laurasian  (Mesidoteinae),  one  is  ubiquitous  (Idoteinae).  and  one  is  endemic  to 
the  shores  of  Morocco  (Parachiridoteinae).  Of  the  six  valviferan  families,  two  are 
cosmopolitan  (Idoteidae  and  Arcturidae),  and  the  remaining  four  are  all  Gondwanan, 
including  the  most  primitive  family,  the  Holognathidae.  No  valviferan  family  is  dis- 
tinctly Laurasian  in  distribution.  These  data  taken  together  strongly  suggest  that  the 
Valvifera  originated  in  the  temperate  Southern  Hemisphere  at  least  by  early  or  mid- 
Pangaean  times  (Permian/Triassic),  if  not  earlier. 

Hurley  and  Jansen  (1977)  reviewed  the  zoogeography  of  the  isopod  family  Sphae- 
romatidae  on  Southern  Hemisphere  coastlines.  Their  data  on  generic  and  species  dis- 
tributions reflect  patterns  similar  to  those  seen  here  in  the  Idoteinae.  Hurley  and  Jansen 
found  generic  affinity  between  all  southern  continents,  as  well  as  generic  endemism  on 
all  shores  (endemism  levels  reported  are  12  genera  [48%]  in  Australia;  7  genera  [41%] 
in  South  Africa;  3  genera  [20%]  in  New  Zealand;  and  1  genus  [1  1%]  in  Chile).  Hurley 
and  Jansen  invoked  strict  dispersalist  mechanisms  to  account  for  these  distributions, 
however,  and  stated,  "The  possibility  of  invoking  continental  drift  as  an  agency  is 
hardly  necessary  in  view  of  the  fact  that  littoral  species  are  involved.'"  What  this 
statement  is  meant  to  imply  is  not  clear.  Presumably  the  authors  are  suggesting  that 
littoral  organisms  are  expected  to  disperse  across  ocean  barriers  with  great  regularity, 
thus  negating  the  roles  of  continental  drift  and  vicariance  in  establishing  endemism. 
This  view  seems  inconsistent  with  their  data,  however,  which  record  high  levels  of 
endemism  at  both  generic  and  species  levels  on  southern  continental  shores  (species 
endemism  on  Australian  shores  was  reported  as  91%;  South  African  shores,  80%;  New 
Zealand,  89%;  Chile,  62%).  In  any  event.  Hurley  and  Jansen  (1977)  did  not  present 
any  phylogenetic  analyses  of  the  taxa  in  question,  making  it  impossible  to  evaluate 
alternative  biogeographic  scenarios  for  the  Southern  Hemisphere  Sphaeromatidae. 

The  present  study  reveals  several  situations  wherein  ancestral  taxa  apparently 
persisted  while  new  (sister)  taxa  evolved  as  peripheral  isolates  (or  perhaps  in  sympatry) 
in  restricted  geographic  regions.  For  example,  Synidotea  almost  certainly  persisted 
unchanged  during  the  events  that  produced  Moplisa  and  Barnardidotea.  Similar  situ- 
ations exist  for:  Paridotca  and  Engidotea:  Zenobiana  and  its  sister-group;  and  Edotea 
and  Parasynifnerus  (see  Fig.  16). 

This  study  suggests  that  both  dispersal  and  vicariant  forces  probably  played  im- 
portant roles  in  creating  modern-day  distributional  patterns  of  idoteine  genera.  Dis- 
persal via  the  Antarctic  Circumpolar  Current  appears  to  have  played  only  a  minor  role, 
however,  as  endemism  on  southern  shores  is  high.  The  unique  (highly  endemic)  nature 
of  these  southern  continental  shores  was  established  long  ago.  Even  New  Zealand, 
which  spans  13  degrees  of  latitude  and  sits  just  1760  km  off  Australia,  is  noted  for  its 
high  endemicity  of  coastal  species.  Examples  include  24%  endemism  for  polychaetes 
(Augener  1924),  50%  for  crabs  (Chilton  and  Bennet  1929),  64%  for  echinoderms  (Mor- 
tensen  1925),  and  89%  for  sphaeromatid  isopods  (Hurley  and  Jansen  1977).  Because 
the  present  analysis  was  at  the  generic  level  rather  than  the  species  level,  and  because 
several  Southern  Hemisphere  genera  cannot  be  clearh  distinguished  in  a  cladogram. 
resolution  of  all  geographic  patterns  has  not  been  achieved.  What  is  clearly  needed  are 
similar  analyses  for  each  of  the  idoteine  genera  (e.g.,  see  Brusca  1 983).  It  is  of  particular 
interest  to  note  that  the  major  lines  of  descent  within  the  Idoteinae  appeared  prior  to 
the  mid-  to  Late  Cretaceous  global  warming  trend,  no  doubt  further  facilitating  the 
temperate-based  distribution  of  this  taxon.  The  present  study  adds  further  evidence  to 
a  large  body  of  data  demonstrating  the  concept  of  the  Crustacea  being  a  taxon  fraught 
with  convergences.  As  Schram  (1978)  deftly  pointed  out,  "The  central  dominating 


26 


1+2 


0+3 


Cleantiella  Lyidotea 

3+1  2+2  1+3  0+4  0+3 


loss  of 
biramous  uropods 


Cleantioides 


I 

0+2 


retention  of 
biramous  uropods 


3+1 


I 

0+1 


Zenobiana 


Edotea 
Parasymmerus 
Eusymmerus 

0+0 


Erichsonella 


N 


ES 

to 


tb 
I 

I 


to 

I 


I 
I 

.5 


I 


-5 

CO 


I 
^ 


b. 


12,13,14 


FkiIkk  17a.  Alternative  schematic  representation  for  lineage  A,  based  on  biogeographic  analysis.  17b. 
Alternative  cladogram  for  lineage  A,  based  on  17a.  Synapomorphies  are  as  follow:  (1)  uropods  uniramous, 
(2)  pereopods  IV  reduced,  (3)  maxillipedal  palp  reduced  in  some  species  to  4  articles,  (4)  pleon  2  +  2.  (5) 
maxillipcdal  palp  reduced  to  4  articles.  (6)  pleon  0+1.  (7)  pleon  1+2.  (8)  pleon  0+0.  (9)  maxillipedal  palp 
reduced  to  3  articles.  (10)  pleon  with  unique,  acute,  subtriangular  shape,  (11)  left  mandible  with  3-pointed 
spine  arising  between  lacinia  and  incisor,  ( 1 2)  pereonite  VII  fused  to  pleonite  1,(13)  maxillipedal  palp  reduced 
to  3-4  articles,  (14)  pleon  0+3.  See  text  for  discussion. 


127 


Ficii'RE  18.     Distribution  of  the  subfamilies  of  Idoteidae  (except  Idoteinae):  Mesidoteinae  [•].  Chaetilinae 
[a],  Glyptonotinae  [O],  Parachiridoteinae  [O]. 


theme  of  arthropod  evolution  is  the  muhiphcity  of  convergent  development.  No  phy- 
letic  scheme,  monophyletic  or  polyphyletic,  can  escape  this." 

Wiley  (1981)  divided  biogeography  into  three  subdisciplines:  descriptive,  histor- 
ical, and  ecological  biogeography.  He  felt  that  the  goals  and  interests  of  the  ecological 
biogeographer  lie  more  with  ecology  than  with  systematics,  whereas  the  opposite  is 
true  of  the  historical  biogeographer.  Finally,  he  suggested  that  systematics  has  little  to 
contribute  directly  to  the  field  of  ecological  biogeography.  I  disagree  with  Wiley,  and 
feel  I  have  shown  here  and  through  a  series  of  studies  that  there  is  a  logical  sequence 
of  basic  taxonomy/descriptive  biogeography/ecological-historical  biogeography.  all  these 
studies  existing  within  the  realm  of  systematics  (i.e.,  Brusca  and  Wallerstein  1977. 
1979fl,  b,  Wallerstein  and  Brusca  1982,  Brusca  1983).  Hessler  and  Wilson  (in  press) 
provide  further  evidence  of  these  relationships  by  their  implication  of  both  ecological 
and  historical  factors  in  reviewing  the  probable  causes  for  differences  in  distribution 
of  major  crustacean  taxa.  Pregill  and  Olson  (1981)  came  to  similar  conclusions  with 
regards  to  the  Caribbean  land  vertebrates,  as  did  Stock  (1981)  regarding  the  Caribbean 
crustacean  stygobionts.  While  the  methods  of  ecological  and  historical  biogeography 
may  differ,  the  two  endeavors  are  clearly  complementary;  to  attempt  one  at  the  expense 
of  the  other  is  to  invite  error. 

Only  a  few  genera  of  Idoteinae  can  be  confidently  tied  to  vicariance  events  asso- 
ciated with  the  breakup  of  Pangaea.  Only  one  New  World  genus  {Parasymmenis)  can 
be  clearly  tied  to  such  major  geological  events.  Vicariant  relationships  seem  to  be  more 
easily  distinguished  at  the  species  level  {see  Brusca  1983).  It  would  seem  that,  at  least 
in  idoteid  isopods,  too  much  time  has  passed  and  too  many  unknowable  events  tran- 
spired (new  taxa  have  evolved  and  gone  extinct,  extant  taxa  have  undergone  local 
extinctions  in  selected  portions  of  their  range,  dispersal  events,  etc.)  to  confidently 
extract  clear  vicariant  patterns  at  the  level  of  supraspecific  taxa.  Furthermore,  those 
idoteinc  taxa  that  do  appear  to  be  products  of  vicariance  phenomena  can  only  be  tested 
by  comparison  to  cladograms  and  distributional  data  for  other  intertidal  groups.  Cra- 
craft  (1982)  has  recently  detailed  such  a  procedure.  Briefly,  allopatric  vicariant  specia- 
tion  (type  la  of  Bush  1975)  implies  the  appearance  of  a  barrier.  In  the  case  of  littoral 
isopods,  these  barriers  would  be  new  stretches  of  ocean  or  new  land  barriers  across 
shallow  seaways.  Such  a  barrier  would  be  expected  to  influence  the  vicariance  patterns 
of  numerous  intertidal  taxa.  and  one  would  thus  predict  that  concordant  pairs  of  sister- 
taxa  would  exist  on  either  side  of  the  barrier.  In  contrast,  speciation  resulting  from  a 
dispersal  event  to  a  new  area  (a  founder  individual  or  population;  type  lb  speciation 
of  Bush  1975)  is  generally  taken  to  be  a  random  event.  Hence,  one  would  predict  not 


128 

to  find  concordant  vicariance  patterns  from  one  clade  to  another.  Absence  of  such 
concordance  with  other  intertidal  taxa  thus  would  suggest  that  speciation  was  initiated 
not  by  a  vicariance  event,  but  by  a  dispersal  event.  Thus,  the  need  for  cladograms  of 
other  marine  invertebrate  groups  becomes  evident,  and  without  such  studies  one  cannot 
critically  assess  the  hypothesized  processes  responsible  for  the  patterns  present  today 
in  the  idoteine  taxa.  Finally,  the  low  levels  of  congruence  between  the  cladogram  of 
taxa-area  and  geographic  cladogram  can  also  be  explained  by  the  simple  hypothesis 
that  Gondwanan  distributions  of  ancestral  idoteid  taxa  were  NOT  widespread,  but 
fragmented  and  local.  This  possibility,  of  course,  denies  a  popular  premise  of  generalized 
tract  theory,  that  ancestral  ranges  can  be  assumed  to  be  the  sum  of  the  ranges  of  the 
descendants. 

Footnotes 

'  There  have  been  several  recent  attempts  to  cast  doubt  on  the  monophyletic  nature  of  the  Peracarida 
(Watling  1981,  Dahl  and  Hessler  1982,  Hessler  in  press).  This  is  not  the  place  to  present  detailed  analyses 
and  criticisms  of  each  of  these  studies.  However,  it  should  be  pointed  out  that  in  Watling's  "cladistic"  analysis 
only  certain  selected  characters  were  used,  numerous  character  transformations  were  illogical  and  unexplained, 
character  states  attributed  to  various  taxa  were  incorrect,  and  a  clear  method  of  character  polarity  assessment 
was  not  provided.  Although  the  other  recent  studies  (op.  cit.)  employed  a  variety  of  approaches,  often  in  the 
guise  of  cladograms,  none  attempted  to  analyze  character  state  polarities  in  a  strict  logical  order  or  with  any 
clearly  expressed  methodology.  Schram  (1981)  was  concerned  with  the  recognition  of  basic  structural  plans 
within  the  Eumalacostraca,  and  his  classification  is  based  on  a  strictly  random  array  of  character  associations. 
The  phenogram  he  chose  for  conversion  into  a  classification  is  the  one  he  felt  "most  comfortable  with." 
Hessler's  (in  press)  "cladogram"  of  the  Peracarida  (his  fig.  5)  is  entirely  unjustified  by  the  data  he  presents. 
Among  other  problems,  no  attempt  was  made  to  achieve  parsimony  and  the  "cladogram"  actually  requires 
more  convergences  (at  least  24)  than  its  contained  character  transformations  (23).  In  the  case  of  Watling 
(1981)  and  Hessler  (in  press)  it  appears  as  though  evolutionary  scenarios  were  conceived  first,  these  then 
being  transformed  into  dendrograms  (incorrectly  called  cladograms)  upon  which  the  appropriate  "apomor- 
phies"  were  overlain.  Hessler's  dendrogram  of  the  Peracarida  is  essentially  Siewing's  (1963  and  earlier 
publications)  concept  of  peracarid  relationships.  This  procedure,  of  course,  ignores  parsimony  considerations 
and  is  the  exact  reverse  of  what  a  phylogenetic  (cladistic)  analysis  is  meant  to  accomplish. 

The  fact  is.  there  exist  many  unique  synapomorphies  that  unite  the  peracarid  orders:  (1)  maxilliped  with 
basis  produced  into  an  anteriorly  directed,  bladelike  endite;  (2)  lacinia  mobilis  present  in  adults;  (3)  oostegites 
formed  on  inner  pereopodal  coxal  margin;  (4)  young  brooded  in  a  brood  chamber  or  "marsupium"  (the 
location  of  the  brood  chamber  varies  from  a  simple  oostegial  pouch  to  invaginations  of  the  sterna,  modified 
oviducts,  or  even  to  the  inner  carapace  region  in  the  thermosbaenaceans);  (5)  direct  development,  with  no 
true  postnaupliar  larval  stages;  (6)  release  of  the  young  as  "manca";  (7)  whiplike  immobile  sperm,  devoid 
of  fibrils  and  mitochondria  (this  character  needs  further  documentation);  (8)  a  large  suite  of  embryological 
attributes  {see  below);  and  (9)  a  large  suite  of  attributes  associated  with  the  functional  morphology  of  the 
pereopods.  These  functional  and  morphological  skeletomuscular  adaptations  are  associated  with  a  system 
unique  to  the  Peracarida,  in  which  the  body-coxa  articulation  has  lost  the  caridoid  "gimbal"  joint  and  become 
either  immobilized  or  capable  of  only  limited  abduction/adduction,  and  the  coxa-basis  articulation  become 
monocondylic  but  capable  of  performing  a  complete  suite  of  motions.  This  peracaridan  system  for  ambulation 
IS  present  in  the  incipient  condition  in  mysidaceans.  Hessler  (1982)  views  these  particular  peracaridan  leg 
synapomorphies  as  adaptations  necessitated  by  the  development  of  the  marsupium  and  its  attendant  oos- 
tegites. Although  thermosbaenaceans  have  lost  the  oostegites,  they  still  retain  the  infolded  monocondylic 
articulation  of  the  pereopodal  coxa  and  basis  {see  Hessler  1982.  for  details). 

The  loss  of  the  oostegites  in  the  thermosbaenaceans  is  probably  a  response  necessitated  by  unique 
morphological  (fusion  of  pereopodal  articles)  and  functional  (locomotory)  adaptations  of  the  pereopodal 
endites  in  this  group.  Loss  of  oostegites  and  concomitant  relocation  of  the  brood  chamber  is  not  unique  to 
the  thermosbaenaceans  among  the  Peracarida  (it  occurs  in  several  hyperiid  amphipods  and  isopod  higher 
taxa).  This  matter  has  been  competently  dealt  with  by  Fryer  (1964)  and  need  not  be  repeated  here  (also  see 
Hessler  1982  and  Slewing  1958). 

The  presence  of  lacinia-like  movable  spines  in  the  larvae  of  a  few  species  of  euphausids  and  shrimps 
suggests  two  possibilities:  (1)  the  lacinia  of  adult  peracarids  is  a  paedomorphic  attribute  {sensu  Gould  1977) 
retained  from  a  nonperacaridcan  ancestor  with  lacinia  in  larval  stages  only,  or  (2)  these  are  superficially 
similar  convergent  features.  The  absence  of  postnaupliar  larval  stages  in  the  Peracarida,  and  their  direct 
development  to  a  juvenile  hatching  stage,  argue  for  their  origin  not  from  a  eucarid  ancestor  but  from  a  line 
separate  from  the  eucarids.  Hence,  the  second  hypothesis  is  the  more  parsimonious.  The  structural  simplicity 
of  the  "lacinia"  of  eucarid  larvae,  versus  the  complexity  of  the  peracarid  lacinia  supports  this  contention. 
Should  one  choose  to  consider  the  movable  mandibular  spine  of  certain  adult  bathynellaceans  to  be  true 
lacinia  (see  Siewing  1963,  Schminke  1972,  and  Dahl  and  Hessler  1982),  one  is  confronted  again  with  two 
possibilities:  (1)  the  adult  lacinia  is  a  feature  representing  a  potential  synapomorphy  for  a  bathynellacean- 
peracarid  line,  or  (2)  this  character  is  convergent  in  these  two  groups.  I  would  accept  either  interpretation 
as  a  working  hypothesis.  The  latter  seems  the  more  likely  considering  the  presence  of  movable  spines  in 
certain  eucarid  larvae  and  the  apparent  plasticity  of  the  spines  of  the  mandibular  row.  The  point  is,  however, 
that  acceptance  of  either  hypothesis  would  not  affect  the  monophyletic  status  of  the  Peracarida!  Frankly, 


129 

with  regards  to  the  monophyly  of  the  Peracarida.  all  the  fuss  over  the  lacinia  seems  "much  ado  about 
nothing." 

The  Peracarida  retain  an  "underlying  unity  in  development"  (Anderson  1973)  distinct  in  numerous  ways 
from  the  Eucarida  and  Syncarida.  This  unity  has  been  well  documented  and  succinctly  summarized  by 
Anderson  (1973).  Unique  attributes  of  pcracaridan  development  appear  in  virtually  all  stages  of  embryo- 
genesis,  and  include  among  other  things:  the  early  segregation  of  primordial  germ  cells  as  a  definite  pre- 
sumptive area;  the  teloblastic  development  of  the  postnaupliar  segments:  the  vitellophage  modifications  in 
the  early  development  of  the  midgut,  the  unique  embryogeny  of  the  digestive  glands:  the  distinct  cmbryogeny 
of  the  ectoderm;  and  the  formation  of  a  second  pair  of  "dorsal  organs"  (the  ectodermal  dorsolateral  organs). 

The  "mancoid"  stage  appears  to  be  little  more  than  the  product  of  alterations  in  embryogeny  and  timing 
in  the  release  of  the  young.  Its  absence  in  mysids  and  amphipods  may  be  tied  to  a  more  rapid  embryological 
development  (or  to  a  delayed  postembryonic  development)  in  these  two  taxa  {see  Steele  and  Steele  1975), 
which  may  also  be  linked  to  the  presence  of  ventrally  curved  embryos  and  completion  of  cleavage  in  the 
early  stages  in  these  groups  (i.e.,  rapid  early  holoblastic  cleavage).  Furthermore,  although  amphipods  leave 
the  marsupium  with  all  7  pairs  of  pereopods  "in  place."  there  is  a  great  deal  of  variation  in  the  structure 
and  development  of  this  appendage,  particularly  in  the  hyperiids.  Laval  (1980)  has  even  recognized  hyperiid 
"larvae,"  with  distinct  hatching  stage  morphologies.  Much  of  the  distinction  between  Laval's  "larvae"  and 
the  adults  involves  the  nature  of  the  pereopods,  and  some  hyperiids  certainly  appear  to  have  "virtual 
mancoids."  The  seventh  pereopods  seem  especially  plastic  in  hyperiids  and  amphipods  in  general.  The 
rudimentary  nature  of  the  seventh  pereopods  in  "juvenile"  amphipods  of  certain  species  has  been  known  at 
least  since  the  work  of  Bate  (1861)  on  Vihilia.  One  could  also  posit  the  origin  of  the  mancoid  stage  subsequent 
to  the  origin  of  the  mysids  and  amphipods  during  peracaridan  cladogenesis.  This  would  remove  this  attribute 
from  the  list  of  synapomorphies  defining  the  Peracarida,  but  it  would  certainly  not  destroy  the  monophyletic 
nature  of  that  ta.xon;  rather,  it  would  simply  make  the  "mancoid  stage"  a  synapomorphy  defining  a  subset 
within  the  Peracarida. 

An  embryological  attribute  that  might  suggest  alliance  among  the  amphipods,  mysids  and  eucarids  is 
the  retention  (from  the  larval  stages)  of  the  functional  antennal  glands  in  these  three  taxa.  Since  all  Crustacea 
have  antennal  glands  during  their  embryogeny,  the  retention  of  these  glands  into  adulthood  in  these  taxa 
hardly  seems  surprising  and  is  most  likely  either  a  convergence  or  simply  a  symplesiomorphy  retained  from 
a  common  ancestor  (i.e.,  a  plesimorphy  not  lost  until  the  appearance  of  the  mysids  and  amphipods  had  been 
achieved  during  peracaridan  cladogenesis). 

I  believe  that  a  carefully  (and  correctly)  accomplished  cladistic  analysis  of  the  Peracarida  will  reveal  the 
amphipods  to  be  the  nearest  relative  if  not  the  sister-group  of  the  isopods.  No  such  analysis  has  been  published 
to  date,  although  one  is  in  preparation  by  F.  Schram.  Watling  (1981),  Hessler  (in  press)  and  others  have 
chosen  to  ignore  or  deemphasize  the  fundamental  synapomorphies  unique  to  these  two  taxa  (e.g.,  sessile 
compound  eyes;  pereonites  with  coxal  plates:  pereopods  entirely  uniramous;  carapace  entirely  lost;  mandible 
of  the  transverse  biting  type),  and  rely  instead  on  differences  and  retained  plesiomorphies  in  their  analyses. 
As  indicated  in  Fig.  2,  I  do  not  believe  that  there  are  ANY  shared  derived  characters  unique  to  the  tanaids- 
isopods. 

-  Note  that  were  the  amphipods  taken  to  be  the  sister-group  to  the  isopods,  attributes  1,  2  and  4  would 
become  synapomorphies  uniting  these  two  taxa. 

'  The  genus  Aiistndotea  is  in  need  of  further  study;  it  may  have  to  be  removed  to  the  Idoteinae. 

■*  Poore  (in  litt.)  has  suggested  that  the  4  non-idoteine  subfamilies  are  synonymous  and  should  be 
combined.  At  the  time  of  this  writing  I  am  in  general  agreement  with  Poore,  but  do  not  address  the  matter 
here. 

'•  The  single  Old  World  species  of  Edotea  (E.  dilatata  Thomson,  1884)  has  been  shown  to  be  the  female 
of  Crabyzos  elongatus  (Miers  1876)  (see  Hurley  1961:292). 

Acknowledgments 

A  number  of  people  took  time  to  read  and  criticize  numerous  early  drafts  of  this 
paper.  The  additional  data  and  discussions  they  provided  me  were  invaluable  to  the 
successful  completion  of  this  study,  and  I  am  indebted  to  them  for  their  sincerity  and 
effort:  J.  L.  Barnard,  T.  Bowman,  N.  Bruce,  G.  Brusca,  P.  Delaney,  J.  Garth.  R.  Hessler, 
J.  Haig,  E.  Iverson,  B.  Kensley.  P.  McLaughlin.  M.  Miyamoto.  A.  M.  S.  Pires.  J.  Savage, 
and  especially  F.  Schram.  G,  Pregill.  and  G.  Poore.  The  philosophical  perspecti\es  of 
these  reviewers  more  often  than  not  differed  markedly  from  one  another,  and  I  hope 
that  I  have  succeeded  in  attending  to  each  of  their  criticisms  in  a  fair  and  appropriate 
manner.  However,  the  views  expressed  in  this  paper  are.  of  course,  my  own  respon- 
sibility. I  also  want  to  thank  the  former  students  of  my  graduate  seminars  in  systcmatics 
and  biogeography  for  their  often  highly  emotional  discussions  on  the  subjects  addressed 
in  this  paper,  particularly  E.  Iverson,  M.  Miyamoto,  B.  White,  and  B.  Wallerstein. 
Many  people  assisted  me  in  obtaining  specimens  on  loan:  I  especially  want  to  thank 
the  Invertebrate  Zoology  staff  of  the  British  Museum  (Natural  History).  G.  Wilson.  W. 
Cooke  (NOSC-Hawaii  Lab).  T.  Bowman,  and  G.  Schull/.  Marie  Hoff-Steinauer  exe- 
cuted the  art  work  (Exhibits  Department,  San  Diego  Natural  History  Museum).  This 
study  was  funded  by  a  grant  from  the  National  Science  Foundation  (DEB  80-17835). 


130 


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^1^      ^^^  TRANSACTIONS 

OF  THE  SAN  DIEGO 
SOCIETY  OF 
NATURAL  HISTORY 


Volume  20  Number  8  pp.  135-144     18  January  1984 


Rhamdia  reddelli,  new  species,  the  first  blind  pimelodid  catfish 
from  Middle  America,  with  a  key  to  the  Mexican  species 

Robert  Rush  Miller 

.v    /  ^n  /•«   /        /c  HARVARD 

Museum  oj  Zoology  ana  Division  of  Biological  Sciences, 

The  University  of  Michigan.  Ann  Arbor.  Michigan  48109  USA  i_NllV 

Abstract.  A  new  blind,  depigmented  catfish  is  described  and  illustrated  from  Cueva  del  Nacimiento 
del  Rio  San  Antonio,  Oaxaca,  Mexico,  and  compared  with  its  closest  relative.  Rhamdia  laticauda.  It 
resembles  that  species  in  having  strong,  retrorse  serrae  on  the  pectoral  spine,  and  a  shallowly  notched 
caudal  fin,  but  differs  in  the  longer  head,  longer  adipose  fin,  larger  cephalic  sensory  pores,  and  longer 
and  more  numerous  gill  rakers  (1  1-16  vs.  9-12).  The  karyotype  (2n  =  58)  of  the  new  species  and  a  key 
to  described  Mexican  species  are  given.  A  list  of  nominal  and  misidentified  Mexican  species  is  presented 
and  R.  laticauda.  R.  parryi.  and  R.  guatcmalensis  are  illustrated. 

Resumen.  Una  nueva  especie  de  bagre  anoftalmo  y  depigmentado  de  la  familia  Pimelodidae  se 
describe  de  la  Cueva  del  Nacimiento  del  Rio  San  Antonio,  Oaxaca,  Mexico.  Se  distingue  de  Rhamdia 
laticauda.  especie  estrechamente  relacionada  a  ella,  por  la  cabeza  mas  grande  y  larga,  la  aleta  adiposa 
mas  larga,  poros  cefalicas  mas  grandes,  y  por  la  longitud  y  el  numero  de  las  branquispinas  (11-16  vs. 
9-12).  El  numero  de  las  cromosomas(2n  =  58),  clavesparaladeterminacion  delosespeciesde  Rhamdia. 
y  illustraciones  de  R.  laticauda.  R.  parryi,  y  R.  guatcmalensis  se  presentan,  y  las  especies  describidas 
de  Mexico  se  listan. 

Heresay  reports  of  blind  catfish  from  Mexico  and  Central  America  have  persisted 
since  before  the  turn  of  the  century.  As  yet,  however,  the  only  described  eyeless  species 
from  this  region  has  been  Prietella  phreatophila  Carranza  (1954),  from  northeastern 
Mexico,  a  member  of  the  Nearctic  family  Ictaluridae  (6  genera,  nearly  40  species). 
Troglobitic  species  apparently  have  evolved  at  least  three  times  in  this  family  (Lundberg 
1982;  Reddell  1981:243-244,  gives  many  references  to  Prietella).  The  Neotropical 
catfish  family  Pimelodidae  is  much  larger  (about  56  genera  and  290  species  according 
to  Nelson  1984)  but.  thus  far,  only  three  blind  species  have  been  described  (see  Thines 
1955,  for  references).  Two  of  these  are  from  Sao  Paulo,  Brazil  {Pimelodella  kronei  and 
Caecorhamdella  brasiliensis),  the  third  from  Trinidad  {Rhamdia  urichi).  A  cave  pop- 
ulation with  the  eye  variably  reduced  was  described  recently  from  Belize  as  Rhamdia 
laticauda  lyphla  (Greenfield  et  al.  1983). 

For  many  years  it  was  common  practice  to  assign  cavernicolous  fishes  to  distinct 
genera  even  though  they  typically  differed  from  their  epigean  relatives  only  in  lacking 
eyes  and  being  depigmented.  That  viewpoint  has  changed  markedly  in  recent  years  (see 
discussions  by  Roberts  and  Stewart  1 976,  and  Banister  and  Bunni  1 980)  and  a  number 
of  blind  fishes  originally  placed  in  monotypic  genera  have  been  reassigned  to  their  more 
widespread  surface  relatives.  In  proposing  the  genus  Caecorhamdia  for  Rhai)idia  urichi, 
Norman  (1926)  wrote  that  his  genus  differed  from  Rhamdia  only  in  lacking  eyes  and 
that  C.  urichi  was  "almost  identical"  with  Rhamdia  quelen,  the  type  species  o{ Rhamdia. 
Mees  (1974:152,  160)  agreed,  placed  Caecorhamdia  in  synonymy  with  Rhamdia.  and 
designated  Norman's  species  as  RhaDidia  quelen  urichi.  Haseman  (1911:325)  stated 
that  Typhlobagrus  kronei  is  indistinguishable  from  Pimelodella  lateristrigata,  except 
for  the  loss  of  sight,  and  recommended  that  the  cave  form  be  relegated  to  subspecific 
status. 


136 


Figure  1.     Rhamdia  reddelli.  A,  Holotype  (5?),  UMMZ  21 1 164,  98.5  mm  SL. 


Catfishes  have  anatomical,  physiological,  and  behavioral  characteristics  that  prea- 
dapt  them  to  life  in  darkness  (e.g.,  well  developed  organs  of  taste  and  touch,  nocturnal 
activity,  crevice-seeking  habits).  It  is  not  surprising,  therefore,  that  about  40  percent 
(17  of  38  species)  of  the  blind  fishes  that  inhabit  fresh  water  are  siluroids.  Among  the 
five  genera  of  pimelodids  inhabiting  Middle  America,  only  Rhamdia  is  widely  distrib- 
uted and  evolutionarily  successful  (Bailey  and  Miller,  1979).  The  Mexican  and  Central 
American  representatives  of  this  genus  are  under  review  by  Reeve  M.  Bailey  and  myself; 
some  of  our  conclusions  receive  advance  notice  in  this  paper. 

The  eyes  of  Rhamdia  are  normally  small  and  of  secondary  importance  in  their 
life.  Field  observations  in  Honduras  by  Carr  and  Giovannoli  (1950)  oi  Rhamdia  bra- 
chycephala  indicate  that  this  species  (a  synonym  of  R.  cabrerai—see  below)  is  ''exclu- 
sively cavernicolous  and  thigmotactic"  in  its  swift-water  habitat. 

The  troglobitic  species  described  below  differs  from  its  epigean  relatives  in  many 
features  other  than  depigmentation  and  loss  of  eyes.  It  may  be  known  as: 

Rhamdia  reddelli  new  species 
Figures  1-3 

Synonymy.— Rhamdia  new  species— LeGrande,  1981:42  (chromosome  and  arm 
numbers,  based  on  UMMZ  199016).  Reddell,  1981:244-245  (mentioned;  type  locality 
mapped). 

Holotype.  — UMMZ  21 1 164,  a  male?  98.5  mm  SL,  Cueva  del  Nacimiento  del  Rio 
San  Antonio,  ca.  9  km  SW  of  Acatlan,  Oaxaca,  on  Atlantic  slope  of  eastern  Mexico; 
collected  by  James  R.  Reddell  and  Andy  G.  Grubbs,  2  January  1977. 

Paratypes.—W\  specimens  are  from  same  locality  as  holotype.  UMMZ  199016  (2 
specimens:  39  and  70  mm),  A.  G.  Grubbs,  M.  Cossey,  and  T.  Byrd,  8  January  1976, 
shipped  alive  to  Ann  Arbor  (larger  individual  karyotyped);  UMMZ  21 1 165  (6  speci- 
mens: 5 1 .5-94.2  mm),  taken  with  the  holotype;  UMMZ  2 1 1 1 66  (77. 1  mm),  R.  Mitchell 
and  L.  Faulkenberry,  7  January  1977.  AMNH  38216  (98.2  mm),  J.  Reddell,  D.  and 
M.  H.  McKenzie,  S.  Murphy,  26  December  1972;  AMNH  38217  (4  specimens:  36.7- 
90.5  mm),  same  collectors  and  date;  AMNH  38218  (2  specimens:  68.5,  69.5  mm), 
same  collectors.  9  March  1973. 

Diagnosis.  — A  species  of  Rhamdia  with  a  broad,  moderately  depressed  skull  in 
adult,  long  head,  very  weak  and  short  occipital  process,  almost  no  pigmentation,  and 
lacking  eyes  (a  tiny  eye  spot  occurs  in  a  37-mm  specimen).  Related  to  R.  laticaiida 
which  it  resembles  in  the  strong,  retrorse  serrae  on  the  posterior  edge  of  the  pectoral 
spine  and  in  the  shallowly  notched  caudal  fin.  From  R.  laticauda  and  its  closest  relatives 
{see  below)  it  differs  in  having:  (1)  a  much  longer  head  and  adipose  fin,  (2)  cavernous 
sensory  head  pores  (especially  on  chin),  and  (3)  longer  and  more  numerous  gill  rakers 


137 


Figure  2.     Rhamdia  reddelli.  Lateral  (A),  ventral  (B).  and  dorsal  (C)  views  of  paratype,  AMNH  38216.  98.2 
mm  SL.  Arrows  indicate  tips  of  pectoral  spines.  The  fork  of  the  caudal  fin  is  too  deep  as  drawn. 


(11-16  VS.  9-12  on  first  arch).  Head  enters  SL  3.25-3.75  times  (vs.  4.0-5.5)  and  the 
depressed  dorsal  fin  overlaps  the  adipose  fin. 

Description.  — Body  form  and  color  pattern  are  indicated  in  Figures  1-2.  Propor- 
tional measurements  are  presented  in  Table  1.  Meristic  data  (based  on  16  fish)  follow. 
Gill  rakers  were  counted  on  the  first  (right)  arch,  with  numbers  for  upper  and  lower 
limbs  recorded  separately  (raker  at  angle  included  in  lower-limb  count).  Vertebral 
counts  are  post-Weberian.  with  separation  of  precaudal  and  caudal  counts  where  pos- 
sible (5  vertebrae  comprise  the  Weberian  complex).  Dorsal  fin  invariably  1,6.  the  spine 
soft  and  flexible  as  typical  oi^  Rhamdia:  anal  rays  13  or  14  (anterior  rudiments  difiicult 
to  see);  pectoral  rays  1,10  or  1,1 1,  usually  1,10  (22  of  30  counts);  pelvic  rays  invariably 
6;  principal  caudal  rays  17-19  (16-18  branched).  Gill  rakers  long,  slender,  3  +  8  to  4+ 12, 
total  11(2),  12(0),  13(5),  14(6),  15(1).  16(1).  Vertebrae:  precaudal,  7  or  8,  caudal.  29- 
32,  total  37-39.  The  number  of  posterior  serrae  on  the  pectoral  spine  varies  with  size, 
from  6-6  in  a  36.7-mm  SL  specimen  to  15-14  in  a  90.5-mm  SL  specimen.  In  the 
larger  fish  the  serrae  are  triangular,  with  very  broad  bases,  and,  except  proximally.  there 
is  no  gap  between  individual  serrae  as  in  Rhamdia  laticauda  and  closest  relatives  {R. 
parryi.  R.  salvini,  R.  cabrcrai^).  The  pectoral  spine  is  gently  curved  in  adults  but  in 
specimens  less  than  70  mm  SL  it  is  straight  and  there  are  gaps  between  the  individual 
serrae.  There  are  no  serrae  on  the  anterior  edge  of  the  spine,  which  is  essentialh  smooth. 

The  maxillary  barbel  is  generally  longer  than  in  R.  laticauda  and  much  longer  than 
in  R.  parryi,  R.  salvini.  or  R.  cahrerai,  but  it  is  shorter  than  in  R.  guatemalensis  which 
belongs  to  a  diflferent  species  group.  It  may  extend  backward  almost  as  far  as  the  tip 
of  the  depressed  dorsal  fin  and  well  beyond  the  origin  of  the  adipose  fin.  but  in  some 


Meek  (1906)  named  this  fish  for  Senor  Cabrera  but  spelled  the  patronym  cabrerae.  It  is  here  corrected. 


138 


8S    ^   ^  SI   SS   ^  « 


^ 

u 

n 

^ 

1^  ^  ^  u  ^  ft  A. 


^    m^    ^  J^    {^    #^ 


FiGi  RE  3.     Somatic  chromosomes  at  metaphase  of  female  paratype,  UMMZ  199016:  2n  =  58.  FN  =  100±4 
(LeGrande,  1981:42). 


it  does  not  reach  beyond  the  middle  of  the  depressed  pectoral  fin.  The  insertion  of  the 
pelvic  fin  lies  before  the  end  of  the  dorsal-fin  base.  The  long  adipose  fin  is  well  developed, 
especially  posteriorly.  The  occipital  process  is  very  weak  and  short,  extending  less  than 
one-fourth  the  distance  to  the  dorsal-fin  origin. 

Alive  in  its  natural  habitat,  Rhamdia  reddelli  is  virtually  colorless  and  without 
visible  pigment.  In  ethyl  alcohol  (formalin-fixed),  the  holotype  (Fig.  1 )  has  fine,  scattered 
flecks  of  pigment  on  the  top  and  sides  of  the  head,  along  the  back  and  upper  sides,  and 
in  a  narrow  band  along  the  lateral  line.  The  lower  sides  and  entire  venter  are  immaculate, 
as  are  all  the  fins  except  the  caudal  which  has  the  interradial  membranes  largely  dusky. 
The  similar-sized  paratype  (Fig.  2)  also  essentially  lacks  pigment  except  for  fine  flecks 
along  and  below  the  base  of  the  adipose  fin  and  some  duskiness  on  the  interradial 
membranes  of  the  caudal  fin.  Other  adults  (between  69  and  9 1  mm  SL),  except  for  one 
mentioned  below,  resemble  either  the  holotype  or  the  above-described  paratype,  or  the 
caudal  fin  may  be  immaculate.  An  81.5-mm  specimen  (UMMZ  21 1 165)  is  more  pig- 
mented, with  fine  flecks  extending  downward  to  a  line  just  above  the  bases  of  the 
pectoral,  pelvic,  and  anal  fins  and  with  fine  pigment  grading  onto  the  ventral  surface 
of  the  caudal  peduncle;  the  venter  is  otherwise  immaculate.  Some  juveniles  (3  or  4  in 
UMMZ  21 1 165)  are  pigmented  as  fully  as  the  holotype,  whereas  others  virtually  lack 
pigment.  When  kept  alive  for  a  month  or  more  in  a  lighted  place,  fine  melanophores 
spread  over  much  of  the  body,  covering  the  venter  posterior  to  the  anus  and  encroaching 
anteriorly  onto  the  abdomen,  with  small  ones  developing  along  the  rays  of  the  paired 


139 


Table  1 .  Proportional  measurements  (in  permillage  of  SL)  of  1 2  specimens  (juv.-ad.)  of  the  types  of  Rhamdia 
reddelti.  The  data  for  the  holotype  are  included  in  the  summary.  Figures  in  parentheses  are  number  of 
specimens  when  fewer  than  12. 


Measurement 


Holotype 


Range 


Mean 


Standard  length,  mm 

Body  depth 

Predorsal  length 

Preanal  length 

Anal  origin  to  caudal  base 

Caudal-peduncle  length 

Caudal-peduncle  depth 

Head  length 

Head  depth 

Head  width  (11) 

Snout  length 

Mouth  width 

Interorbital  width 

Maxillary-barbel  length  (9) 

Outer  mental  barbel  length  (11) 

Inner  mental  barbel  length  (11) 

Adipose-fin  length 

Adipose-fin  maximum  height 

Anal-fin  basal  length 

Pectoral-fin  length  (11) 

Pectoral-spine  length 

Caudal-fin  length  (9) 

Caudal  fin,  to  notch  (8) 

Caudal  fin,  shortest  ray  length  (8) 


98.5 

50.3-98.5 

76.8 

198 

163-234 

194 

376 

366-414 

385 

682 

640-707 

675 

340 

300-356 

335 

201 

163-206 

186 

124 

99-130 

119 

286 

278-310 

295 

157 

138-171 

155 

213 

185-226 

205 

123 

101-137 

120 

145 

127-151 

142 

87 

79-100 

89 

434 

337-516 

466 

211 

175-287 

229 

128 

104-161 

127 

401 

337-417 

377 

42 

29-49 

42 

139 

131-158 

145 

172 

157-191 

171 

96 

91-121 

103 

211 

208-241 

223 

86 

76-111 

95 

124 

123-149 

134 

fins  and  the  anal  fin  as  well  as  on  the  interradial  membranes  of  the  dorsal  fin  (e.g.,  the 
69-mm  SL  specimen,  UMMZ  199016,  kept  alive  over  six  months).  The  one  individual 
with  a  tiny  pigmented  pupil  (AMNH  38217,  36.7  mm  SL)  appears  to  lack  pigment. 

From  the  recently  described  Lake  Nicaraguan  species  Rhamdia  hiigiana  Villa 
(1977).  the  new  species  differs  markedly  in  the  much  deeper  body,  shorter  pectoral 
spine  and  fin.  longer  head,  shorter  maxillary  barbel,  shallower  caudal-fin  notch,  and 
rounded  rather  than  pointed  caudal-fin  lobes.  It  also  has  fewer  post- Weberian  vertebrae 
(37-39  vs.  39-42  in  R.  luigiana).  Rhamdia  reddelli  is  compared  with  other  Mexican 
species  of  Rhamdia  in  the  Key. 

The  karyotype  (Fig.  3),  kindly  prepared  by  William  H.  LeGrande,  shows  a  diploid 
number  of  58  chromosomes  and  an  arm  number  {¥N)  of  100±4.  This  diploid  number 
may  be  the  ancestral  condition  for  pimelodids,  as  it  evidently  is  for  ictalurids  (LeGrande 
1981),  but  since  the  karyotypes  of  only  seven  species  of  pimelodids  have  been  published 
(2n  =  46,  56,  58,  62— see  LeGrande  1981:42)  this  tentative  conclusion  must  await 
further  karyological  studies  of  this  family. 

Habitat  and  associates.— Jho:  cave  is  at  Canada  San  Antonio,  approximately  9  km 
SW  of  Acatlan,  at  an  elevation  of  100  m.  The  stream  flowing  from  below  the  cave 
entrance  is  the  primary  source  of  the  Rio  San  Antonio  and  drains  south  into  Presa 
Miguel  Aleman,  a  dammed  portion  of  the  Rio  Tonto  that  is  tributary  to  Rio  Papaloapan. 
The  main  passage  of  the  cave  extends  for  about  120  m  to  a  deep  lake  containing  many 
blind  catfish  and  crayfish.  Beyond  this  lake  a  shallow  stream  extends  for  350  m  in  a 
passage  10  to  30  m  wide  and  up  to  1  1  m  high.  Several  major  side  passages  that  contain 
secondary  streams  occur  throughout  the  cave  and  bring  the  total  length  of  the  cave  to 
about  4.5  km.  The  main  stream  floor  is  generally  of  sand  and  gravel  with  areas  of 
flowstone  and  bedrock.  Both  air  and  water  temperatures  were  23. 5°C.  The  catfish  usually 
occurred  in  the  deeper  ponded  portions  of  the  streams,  especially  in  areas  over  which 
bats  roosted. 

A  rich  invertebrate  cave  fauna  is  associated  with  the  catfish.  Four  species  of  trog- 
lobitic  crustaceans  inhabit  the  cave:   Potamalpheops  stygicola  Hobbs  (Decapoda: 


140 


Alpheidae),  Macrobrachiu!)i  villalobosi  Hobbs  (Decapoda:  Palaemonidae).  Procam- 
banis  (Austrocaniharus)  oaxacae  rcddclli  Hobbs  (Decapoda:  Cambaridae),  and  Speleo- 
mysis  olivae  Bowman  (Mysidacea:  Lepidomysidae).  All  have  since  been  collected  in 
other  caves  in  the  vicinity  of  Acatlan.  A  second  species  of  mysid,  Antromy»is  (Antro- 
mysis)  reddelli  Bowman,  has  been  collected  from  a  nearby  cave  and  can  be  expected 
to  occur  in  Cueva  del  Nacimiento  del  Rio  San  Antonio.  A  specimen  of  the  alpheid 
shrimp  Potanialpheops  stygicola  was  disgorged  by  a  catfish  upon  preservation.  The 
rarity  of  shrimps  and  mysids  in  pools  containing  catfish  is  doubtless  related  to  predation 
by  the  fish  on  the  crustaceans.  The  cave  is  also  inhabited  by  a  possibly  troglobitic  clam, 
which  is  abundant  in  various  parts  of  the  cave  but  awaits  study. 

The  terrestrial  fauna  is  extremely  abundant  and  includes  troglobitic  trichoniscid 
isopods,  nicoletiid  thysanurans,  millipeds,  spiders,  and  opilionids. 

Etymology.  — \  am  pleased  to  name  this  distinctive  species  for  James  R.  Reddell, 
who  donated  all  of  the  type  specimens  and  has  pioneered  in  exploring  caves  in  Latin 
America. 

Nominal  or  Misidentified  Species  Referred 
TO  Mexican  Catfishes  of  the  Genus  Rhamdia 

Piinelodiis  laticaudus  Meckel  in  Kner.  1858  (Abtheilung  Sitzber.  Akad.  Wiss.  Wien,  26: 
420).  Type  locality  (on  label  in  jar):  "Rio  Xamapa,  Mexiko."  Oldest  available 
name  for  a  Middle  American  Rhamdia.  A  valid  species.  Three  syntypes,  Vienna 
Museum  50554  (166,  171,  203  mm  SL),  examined. 

Pi melodus  guatema/ensis  Gunthev,  1864  (Cat.  Fish.  British  Mus.,  5:122).  Type  locality: 
Huamuchal,  on  Pacific  coastal  plain,  Guatemala.  A  valid  species. 

Pimelodus  godmani  Giinther,  1 864  (ibid.:  1 24).  Type  locality:  Guatemala  (Rio  Motagua, 
lower  Vera  Paz)  and  Mexico.  Species  illustrated  by  Regan  (Biol.  Centrali-Ameri- 
cana,  8:pl.  21,  fig.  1).  A  synonym  o^ Rhamdia  guatemalensis  (Miller  1966:787). 

Pimelodus petenensis  Giinther,  1864  (ibid.:  126).  Type  locality:  Lake  Peten,  Guatemala. 
Listed  for  Mexico  by  Alvarez  1 950  (Sec.  de  Marina,  Dir.  Gen.  Pesca  e  Ind.  Conexas, 
Mexico:35)  with  the  remark  ''probably  only  in  Guatemala."  Illustrated  by  Regan 
(op.  cit.:pl.  22,  fig.  1).  A  subspecies  of/?,  guatemalensis  (Hubbs  1938:266). 

Pimelodus  hypselurus  Giinther,  1864  (ibid.:  126-1 27).  Type  locality  (on  label  in  jar): 
Orizaba  [but  listed  as  Cordova  in  cat.  book],  Mexico.  Holotype,  BMNH  1858- 
1 1.22.32  (103.5  mm  SL),  examined  by  R.  M.  Bailey;  illustrated  by  Regan  (op.  cit.: 
pi.  21,  fig.  3).  A  synonym  of  R.  laticauda. 

Pimelodus  motaguensis  Giinther,  1864  (ibid.:  127).  Type  locality:  Rio  Motagua,  Gua- 
temala. Holotype  illustrated  by  Regan  (op.  cit.:pl.  20,  fig.  1).  A  synonym  oi  R. 
laticauda  (see  comment  by  Miller  1976:4). 

Pi?nelodus  brachypterus  Cope.  1866  (Trans.  Amer.  Philos.  Soc,  13:404).  Type  locality: 
Orizaba,  Mexico.  Holotype,  ANSP  16471  (147  mm  SL),  examined.  A  synonym 
of  R.  laticauda. 

Rhamdia  parryi  Eigenmann  and  Eigenmann,  1888  (Proc.  Calif  Acad.  Sci.,  ser.  2,  vol. 
1:130).  Type  locality:  Rio  Zanaleneo  [=Sanatenco],  near  Tonala,  on  Pacific  slope 
of  Chiapas,  Mexico.  Five  syntypes,  MCZ  27273  (77-88  mm  SL),  examined.  A 
valid  species  (called  R.  hypselura  by  Miller  1966:787). 

Rhamdia  oaxacae  Meek,  1902  (Field  Col.  Mus.  Publ.  65:74,  pi.  14).  Type  locality:  Rio 
Quiotepec  at  Cuicatlan,  Oaxaca,  Mexico,  in  Rio  Papaloapan  basin.  A  synonym  of 
R.  guatemalensis  (Regan  op.  cit.:  132). 

Rhamdia  depressa  Barbour  and  Cole,  1906  (Bull.  Mus.  Comp.  Zool.  50(5):  155,  pi.  1). 
Type  locality:  Ikil  [=Ixil]  Cenote  near  Chichen-Itza,  Yucatan,  Mexico.  A  subspecies 
of  R.  guatemalensis  (Huhbs  1936:194). 

Rhamdia  sacrificii  Barbour  and  Cole,  1906  (ibid.:  156).  Type  locality:  Sacrificial  cenote 
near  Chichen-Itza,  Yucatan,  Mexico.  A  synonym  of  R.  guatemalensis  (Hubbs  1 936: 
200). 

Pimelodus  houcardi  Regan,   1907  (Ann.  Mag.  Nat.  Hist.,  ser.  7,  vol.   19:258).  Type 


141 


HjjHj^^ 

A 

ii 

-^   ^ 

1 

1 

1 

1 

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:y:!-  ■ 

^"1 

I^J 

■  -■■■in  l.iii^i     1   "T     " 

1 

■ 

FicH'RE  4.  Three  species  of  Rhanidia  from  Mexico.  A,  7?.  guatemalensis.  UMMZ  183901  (126.2  mm). 
Nacimiento  del  Rio  Cosalapa,  2.6  km  SE  of  Estacion  Tezonapa,  Veracruz;  B.  R.  laticauda.  UMMZ  196674 
(68.8  mm),  tributan,  to  Rio  Metlac.  Fortin.  Veracruz;  C,  R.  parryi.  UMMZ  184739  (68.0  mm),  headwaters 
of  Rio  Tapanatepec,  at  Hwy  190  bridge  E  of  Tapanalepec,  Oaxaca. 


locality:  Yucatan.  Mexico.  A  synonym  of  R.  guatemalensis  depressa  Barbour  and 
Cole  (Hubbs  1936:193,  195);  illustrated  by  Regan  (op.  cit.:pl.  20,  fig.  3). 
Pimehdiis  brachycephalus  Regan,  1 907  (op.  cit.:258).  Type  locality:  Rio  Nacasil,  Pacific 
slope  of  Guatemala.  Recorded  tentatively  from  Mexico  by  Alvarez  1950  (op.  cit.: 
37)  with  the  remark  "probably  only  in  Guatemala."  Illustrated  by  Regan  (op.  cit.: 
pi.  22,  fig.  2).  A  synonym  of  R.  cahrerai  (type  examined  by  R.  M.  Bailey).  Later. 
Alvarez  (1970:77)  listed  R.  hrachycephala  from  "southeastern  Mexico  near  the 


142 


Guatemalan  frontier,"  but  this  represents  a  misidentification  since  R.  cabrerai  is 
known  on  the  Atlantic  slope  of  Guatemala  only  from  the  upper  Rio  Motagua. 

Key  to  Mexican  Species  of  Rhamdia-  » 

R.  laticauda  and  R.  parryi  are  commonly  sympatric  with  R.  guatemalensis. 

la.  Anterior  and  posterior  edges  of  pectoral  spine  with  small,  numerous  serrae  of 
subequal  length,  developed  about  equally  or  those  on  posterior  edge  somewhat 
stronger  (especially  in  older  fish);  caudal  fin  deeply  notched  for  at  least  two- 
thirds  the  distance  from  tips  of  caudal  lobes  to  base  of  mid-caudal  rays. 
(Maxillary  barbel  long,  typically  extending  well  beyond  origin  of  adipose  fin; 
head  long,  3.5-3.8  in  SL;  adipose  fin  long,  ca.  one-third  SL;  occipital  process 
long,  extending  nearly  halfway  or  more  to  dorsal  origin.)  Atlantic  and  Pacific 
lowlands  from  just  NW  of  Veracruz  City  on  Atlantic  slope  and  Rio  Tehuan- 
tepec  basin  on  Pacific  versant  southward  to  Panama  (if  R.  wagneri  is  a  syn- 
onym—56'e'  Hubbs  1936:181);  typically  in  pools    

Rhamdia  guatemalensis  (Fig.  4) 

b.  Pectoral  spine  with  strong,  retrorse  to  nearly  straight  serrae  only  on  posterior 
edge  (anterior  edge  smooth  or  roughened);  caudal  fin  weakly  notched,  to  no 
more  than  half  distance  from  tips  of  caudal  lobes  to  base  of  mid-caudal  rays. 
Atlantic  and  Pacific  versants\  cavernicolous  and  in  rocky  streams  of  piedmont 

slopes  and  foothills    2 

2a.  Skull  depressed;  head  long,  3.25-3.75  in  SL;  blind  and  depigmented;  adipose 
fin  well  developed,  overlapped  by  depressed  dorsal  fin.  Cueva  del  Nacimiento 
del  Rio  San  Antonio,  Oaxaca;  cavernicolous   Rhamdia  reddelli  (Figs.  1-2) 

b.   Skull  domed;  head  short,  4.0-5.5  in  SL;  eyes  and  pigment  well  developed; 

adipose  fin  short,  not  (or  rarely)  overlapped  by  depressed  dorsal  fin    3 

3a.  A  prominent,  dark  lateral  stripe  on  midside,  from  behind  head  to  base  of 
caudal  fin,  becoming  broader  posteriorly;  post-Weberian  vertebrae  fewer,  35- 
38,  usually  36  or  37  (98%).  Pacific  slope  of  Oaxaca  and  Chiapas  southeastward 

into  Guatemala  (to  Dpto.  de  Santa  Rosa);  on  rocky  riffles    

Rhamdia  parryi  (Fig.  4) 

b.  Side  of  body  without  a  conspicuous  dark  stripe;  post-Weberian  vertebrae  more 
numerous,  37-41,  usually  38-40  (91%).  Atlantic  slope  from  Rio  Jamapa, 
Veracruz,  southeastward  to  western  Honduras;  on  rocky  riffles  and  in  current 
of  streams    Rhamdia  laticauda  (Fig.  4) 

Acknowledgments 

Robert  W.  Mitchell  called  my  attention  to  the  existence  of  the  new  species.  James 
R.  Reddell  provided  the  information  from  which  the  account  of  the  habitat  and  as- 
sociates was  written. 

I  am  grateful  to  the  following  for  information,  specimens,  radiographs,  and  other 
assistance,  and  for  loans  of  important  material  needed  for  the  completion  of  this  paper: 
P.  H.  Greenwood,  British  Museum  of  Natural  History  (BMNH),  Paul  Kahsbauer, 
Vienna  Museum,  Donn  E.  Rosen,  American  Museum  of  Natural  History  (AMNH), 
James  E.  and  Eugenia  B.  Bohlke,  Academy  of  Natural  Sciences  of  Philadelphia  (ANSP), 
Karel  Liem  and  William  L.  Fink,  Museum  of  Comparative  Zoology  at  Harvard  (MCZ), 
Robert  H.  Gibbs,  Stanley  H.  Weitzman,  and  Susan  L.  Jewett,  United  States  National 
Museum  of  Natural  History  (USNM),  Reeve  M.  Bailey,  Charles  E.  Dawson,  Andy  G. 
Grubbs,  William  H.  LeGrande,  Doyle  Mosier,  Marcia  K.  Dorsey,  and  Alexandra  Snyder 
Creighton.  Louis  P.  Martonyi  (Fig.  4B),  Edward  C.  Theriot  (Figs.  1,  4C),  and  William 
M.  Pelletier  (Fig.  4A)  are  responsible  for  the  photographs,  Anita  Cleary  for  the  drawings 
(Fig.  2).  Margaret  L.  Van  Bolt  mounted  Figure  2  and  added  heavier  arrows.  Mexican 


Much  of  this  key  was  derived  from  information  provided  by  R.  M.  Bailey. 


143 


officials  permitted  me  to  conduct  the  field  work,  and  the  National  Science  Foundation 
supported  field  and  laboratory  work  (most  recently  as  grant  DEB  80-02017).  Three 
anonymous  reviewers  improved  the  manuscript. 


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Nac.  Inv.  Biol.  Pesq.,  Ser.  Inv.  Pesq..  Estudio 
No.  1,  Mexico.  166  pp. 

Bailey.  R.  M.,  and  R.  R.  Miller.  1979.  Pimelodid 
catfishes  (genus  Rhamdia)  from  northern  Mid- 
dle America.  .Abst.  59th  .Ann.  Mtg.  Amer.  Soc. 
Ich.  &  Herp..  Orono.  Maine.  1  p. 

Banister.  K.  E..  and  M.  K.  Bunm.  1980.  A  new 
blind  cyprinid  fish  from  Iraq.  Bull.  Brit.  Mus. 
Nat.  Hist.  (Zool.)  38(3):  15  1-158. 

Carr.  A.  F.  Jr..  and  L.  Giovannoli.  1950.  The 
fishes  of  the  Choluteca  drainage  of  southern 
Honduras.  Occ.  Pap.  Mus.  Zool.  Univ.  Mich. 
523:1-38. 

Carranza.  J.  1954.  Descripcion  del  primer  bagre 
anoftalmo  y  depigmentado  encontrado  en  aguas 
mexicanas.  Ciencia  1 4(7-8):  129-1 36. 

Greenfield.  D.  W..  T.  A.  Greenfield,  and  R.  L. 
Woods.  1983.  A  new  subspecies  of  cave- 
dwelling  pimelodid  catfish,  Rhamdia  laticauda 
tvphla.  from  Belize.  Central  America.  Brenesia 
19/20(1982):563-576. 

Haseman,  J.  D.  1911.  Descriptions  of  some  new 
species  of  fishes  and  miscellaneous  notes  on 
others  obtained  during  the  expedition  of  the 
Carnegie  Museum  to  central  South  America. 
Ann.  Carnegie  Mus.  7(3-4):3 15-328. 

Hubbs,  C.  L.  1936.  Fishes  of  the  Yucatan  Pen- 
insula. Carnegie  Inst.  Wash.  Publ.  457:157- 
287. 

.    1938.    Fishes  from  the  caves  of  Yucatan. 

Carnegie  Inst.  Wash.  Publ.  491:261-295. 

LeGrande.  W.  H.  1981.  Chromosomal  evolution 
in  North  American  catfishes  (Siluriformes:  Ic- 
taluridae)  with  particular  emphasis  on  the 
madtoms,  Notunis.  Copeia  1981(l):33-52. 

Lundberg.  J.  G.    1982.   The  comparative  anatomy 


of  the  toothless  blindcat,  Trogloglams  paller- 

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of  the  ictalurid  catfishes.  Misc.  Publ.  Mus.  Zool. 

Univ.  Mich.  163:1-85. 
Meek,  S.  E.    1906.  Description  ofthree  new  species 

of  fishes  from  Middle  America.  Field  Col.  Mus. 

Publ.  1 16,  Zool.  Ser.  7(3):93-95. 
Mees,  G.  F.    1974.    The  Auchenipteridae  and  Pi- 

melodidae  of  Suriname  (Pisces,  Nematogna- 

thi).  Zoolog.  Verhand.  132:1-256  +  pis.  1-15. 
Miller,  R.  R.    1966.    Geographical  distribution  of 

Central  American   freshwater  fishes.   Copeia 

1966(4):773-802. 
.     1976.    An  evaluation  of  Seth  E.  Meek's 

contributions  to  Mexican  ichthyology.  Field- 

lana.  Zool.  69(1):  1-31. 
Nelson,  J.  S.    1984.    Fishes  of  the  World.  2nd  ed. 

John  Wiley  &  Sons,  N.  Y.  416  pp. 
Norman,  J.  R.     1926.    A  new  blind  catfish  from 

Trinidad,  with  a  list  of  the  blind  cave-fishes. 

Ann.  Mag.  Nat.  Hist.  ser.  9,  vol.  18:324-331. 
Reddell,  J.  R.    1981.    A  review  of  the  cavernicole 

fauna  of  Mexico,  Guatemala,  and  Belize.  Texas 

Mem.  Mus.,  Bull.  27:1-327. 
Roberts,  T.  R.,  and  D.  J.  Stewart.    1976.    An  eco- 
logical and  systematic  survey  of  fishes  in  the 

rapids  of  the  lower  Zaire  or  Congo  River.  Bull. 

Mus.  Comp.  Zool..  Harvard  Univ.  147(6):239- 

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gine,    taxonomie.    repartition,    geographique. 

comportement.  Ann.  Soc.  Roy.  Zool.  Belgique 

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o(oH^      y^  TRANSACTIONS 


MUS.  COMP.  ZOOL 

LfPPARY 


OF  THE  SAN  DIEGO 
SOCIETY  OF 
NATURAL  HISTORY 


[OCT  1 0 1984 

Volume  20  Number  9  pp.  145-150      20  June  1984 

A  complete  specimen  of  Peachella  brevispina  Palmer— an  unusual 
olenellid  trilobite  (Arthropoda:  Olenellida)  from  the 
lower  Cambrian  of  California 

James  H.  Stitt 

Department  of  Geology,  University  of  Missouri-Columbia,  Columbia,  MO  65211  USA 

R.  L.  Clark 

Department  of  Geology,  Paleontology  Section.  San  Diego  Natural  History  Museum, 
P.O.  Box  1390.  San  Diego,  C A  92112  USA 

Abstract.  The  thorax  and  possible  pygidium  of  Peachella  brevispina  are  described  for  the  first 
time  from  a  recently  collected  complete  specimen,  the  first  recovered  for  this  genus.  Unusual  features 
include  the  large  macropleural  third  segment  of  the  prothorax  that  distorts  the  adjacent  pleurae,  the 
rounded  distal  extremeties  of  the  other  prothoracic  segments,  the  rather  gradual  transition  from  the 
prothorax  to  the  opisthothorax,  and  the  large,  triangular  pygidium. 

Introduction 

Olenellid  trilobites  are  found  in  Lower  Cambrian  strata  in  various  parts  of  the 
world,  and  numerous  specimens  have  been  collected  from  localities  in  western  North 
America.  Olenellids  are  broad,  flat,  medium  to  large-sized  trilobites  characterized  by 
a  large  semicircular  cephalon,  prominent  ocular  lobes,  numerous  thoracic  segments 
that  terminate  laterally  in  spines  of  various  lengths,  and  a  small  pygidium.  The  absence 
of  specialized  morphologic  features  on  the  thorax  that  were  used  for  enrollment  in 
other  trilobites  (Bergstrom  1973:17)  indicates  that  trilobites  of  the  Family  Olenellidae 
were  unable  to  enroll  (Harrington  1959:102),  a  protective  feature  that  developed  in 
other  Lower  Cambrian  (Rasetti  1948:17-18,  pi.  4,  figs.  22-24)  and  later  trilobites. 

Initial  taxonomic  efforts  on  olenellid  trilobites  resulted  in  the  recognition  of  a 
number  of  closely  related  genera  (Walcott  1910)  and  species  (Resser  and  Howell  1938). 
More  recent  work  (Fritz  1972,  Palmer  and  Halley  1979)  has  resulted  in  recognition  of 
fewer  genera,  because  the  criteria  originally  used  to  discriminate  certain  genera  inter- 
grade  among  various  species.  Added  to  this  problem  is  the  fact  that  olenellids  frequently 
occur  in  the  same  beds  of  rock  in  pairs  of  closely  related  taxa  that  have  been  variously 
interpreted  as  paired  species  or  dimorphs  (see  Palmer  and  Halley  1974:66-67  for  a 
recent  discussion  of  these  taxonomic  problems). 

The  complete  specimen  of  Peachella  brevispina  Palmer  described  in  this  paper  was 
collected  by  R.  L.  Clark  from  the  lower  part  of  the  Carrara  Formation  at  Emigrant 
Pass  in  the  Nopah  Range,  Inyo  County,  California  (Fig.  1).  Palmer  and  Halley  (1979: 
13,  75;  fig.  1)  reported  that  P.  brevispina  is  moderately  rare  in  the  Thimble  Limestone 
Member  of  the  Carrara  in  the  nearby  Dublin  Hills  and  at  Eagle  Mountain.  Mount 
(1980:78-80;  fig.  1)  described  the  Emigrant  Pass  locality  in  the  Nopah  Range,  including 
a  detailed  columnar  section  and  range  chart  in  which  he  reported  P.  brevispina  from 
a  thin  limestone  bed  near  the  top  of  an  unnamed  lower  member  of  the  Carrara.  Mount's 
columnar  section  supports  Palmer  and  Halley's  observation  (p.  9,  13;  figs.  6A,  1 1)  that 
the  Thimble  Limestone  Member  of  the  Carrara  cannot  be  recognized  in  the  Nopah 
Range  because  the  Thimble  grades  southeastward  into  shale. 


146 


Peachella  is  an  unusual  genus  of  olenellid  trilobites  that  was  known  only  from  a 
small  number  of  cephala  before  Clark's  discovery  of  the  complete  specimen  described 
in  this  paper.  The  glabella,  ocular  lobes  and  cephalic  border  are  fainter  and  less  well 
defined  than  usual  for  an  olenellid.  The  most  unusual  feature  of  the  ceph^on  is  the 
genal  spines,  which  are  normally  long  and  pointed  in  other  olenellids  but  in  species  of 
Peachella  are  short,  wide  and  have  broadly  rounded  tips.  First  described  by  Walcott 
in  1910,  this  scarce  genus  included  only  the  type  species  Peachella  iddingsi  (Walcott) 
until  1979,  when  Palmer  described  a  second  species,  P.  brevispina,  from  the  Thimble 
Limestone.  Clark's  discovery  of  a  complete  specimen,  which  he  donated  to  the  San 
Diego  Natural  History  Museum  (SDSNH  locality  no.  3169),  allows  the  description  of 
this  unusual  trilobite  to  be  completed. 

Morphologic  terms  used  in  the  following  description  are  defined  in  Harrington 
(1959).  Suprageneric  classification  follows  Bergstrom  (1973)  and  Palmer  and  Halley 
(1979). 

Systematic  Paleontology 

Phylum  Arthropoda  Siebold  and  Stannius,  1 845 

Class  Trilobita  Walch,  1771 

Order  Olenellida  Resser,  1938 

Family  Olenellidae  Vogdes,  1893 

Genus  Peachella  Walcott,  1910 

Peachella  brevispina  Palmer,  1979 

Figures  2,  3 

Peachella  brevispina  PALMER  in  Palmer  and  Halley  (1979:75,  pi.  5,  figs  1-3). 

Material.—  1  complete  decalcified  specimen  (SDSNH  24548)  preserved  as  internal 
and  external  molds;  specimen  slightly  deformed,  especially  the  cephalon  and  pygidium. 

Description.  — 'LtngXh  of  specimen  33  mm  (excluding  macropleural  spines).  Ceph- 
alon semicircular  in  outline  with  short,  paddle-like  genal  spines.  Anterior  end  of  ce- 
phalon poorly  preserved.  Glabella  prominent,  elongate,  extends  close  to  anterior  edge 
of  cephalon,  set  off  by  rather  shallow  axial  furrows.  No  lateral  glabellar  or  occipital 
furrows  visible.  Occipital  ring  not  differentiated.  Ocular  lobes  poorly  preserved,  ap- 
parently short,  arcuate  and  located  close  to  the  glabella.  Anterior  border  and  border 
furrow  not  preserved.  Lateral  border  narrow  opposite  anterior  end  of  glabella,  gradually 
widens  posteriorly.  Lateral  border  furrow  shallow,  curved  slightly  adaxially  at  anterior 
end  of  genal  spine,  crossing  adaxial  base  of  genal  spine  and  continuing  to  posterior 
margin  of  cephalon.  Posterior  border  furrow  not  visible.  Characteristic  genal  spines 
short,  broad,  moderately  inflated,  posterolaterally  directed;  spine  termination  broadly 
rounded. 

Prothorax  of  1 5  segments.  Axis  prominent,  convex,  width  (trans.)  approximately 
one-fourth  of  prothoracic  width.  Most  articulating  half-rings  visible  on  outstretched 
specimen.  Axial  furrows  shallow.  Axial  ring  and  transverse  furrow  of  first  prothoracic 
segment  slope  steeply  anteriorly,  articulating  half-ring  apparently  still  connected  to 
posterior  ventral  edge  of  cephalon.  Axial  ring  of  second  prothoracic  segment  separated 
from  narrow  articulating  half-ring  by  prominent,  wide,  deep  transverse  furrow  that 
curves  slightly  anteriorly  abaxially  and  intersects  the  axial  furrows.  Remaining  axial 
rings  separated  from  articulating  half-rings  by  faint,  narrow  transverse  furrow  that 
curves  anteriorly  abaxially  and  terminates  near  anterior  edge  of  ring  before  reaching 
axial  furrows.  This  furrow  deepest  on  axial  rings  of  prothoracic  segments  12-14,  be- 
coming progressively  fainter  on  anterior  segments.  Small  axial  node  present  on  posterior 
edge  of  axial  rings  of  prothoracic  segments  10-14;  node  most  prominent  on  segment 
14,  progressively  diminishing  in  size  anteriorly.  Prothoracic  segment  15  broken  at  axis, 
undoubtedly  bears  axial  spine  whose  mold  can  be  seen  entering  the  surrounding  rock 
just  posterior  of  the  opisthothorax. 

Pleurae  of  prothorax  variable  in  appearance.  Pleurae  of  first  two  prothoracic  seg- 


147 


Figure  1.     Map  showing  location  of  Emigrant  Pass  locality  where  described  specimen  was  collected. 


ments  not  well  preserved,  appearing  flat  with  no  pleural  furrows  and  rounded  distal 
extremities.  Faint  transverse  ridge(s)  present  on  left  pleurae  of  segments  1  and  2  not 
present  on  right  pleurae,  and  interpreted  to  be  result  of  deformation  of  specimen.  Pleurae 
of  segment  3  expand  rapidly  abaxially,  especially  along  posterior  edge,  becoming  broad- 
ly oval  in  shape  and  approximately  seven  times  as  long  (exsag.)  as  the  pleurae  of  any 
other  segment.  Pleurae  of  segment  3  reach  maximum  length  (exsag.)  near  distal  margins 
of  thorax,  pleurae  narrow  abaxial  of  maximum  length  to  form  large,  long  macropleural 
spine  that  curves  posteriorly  and  extends  well  beyond  the  posterior  end  of  the  specimen. 
Unusual  ovoid  shape  of  pleurae  of  segment  3  distorts  shape  of  pleurae  of  segments  2 
and  4-6,  forcing  these  pleurae  to  compress  abaxially  and  to  taper  distally  (to  rounded 
extremeties)  in  order  to  accommodate  wider  pleurae  of  segment  3  without  leaving  gaps 
or  overlaps  between  pleurae.  Pleurae  of  prothoracic  segments  7-14  rather  plain,  nearly 
flat,  with  rounded  distal  extremeties.  All  pleurae  posterior  of  third  pleurae  divided  into 
narrow  anterior  and  posterior  bands  by  broad,  shallow  pleural  furrow.  Pleural  furrow 
narrower  and  deeper  in  segments  12-14.  Pleurae  of  prothoracic  segments  10-14  bend 
gently  to  the  posterior  abaxially,  with  posterior  bending  greatest  toward  posterior  end 
of  prothorax. 

Opisthothorax  consists  of  eight  or  nine  segments.  Axis  only  faintly  defined  by  very 
shallow  axial  furrows.  Pleurae  simple,  nearly  flat,  apparently  with  no  pleural  furrows 
and  rounded  distal  extremeties. 


148 


a 


b 


Figure  2.  Stereophotographs  of  internal  mold  of  specimen  of  Peachella  brevispina  Palmer  (SDSNH  24548) 
from  lower  part  of  Carrara  Formation,  Emigrant  Pass,  California;  a.  entire  specimen,  x2;  b.  enlargement  of 
thorax  and  pygidium,  x4. 


149 


■>^ 


N 


.V^. 


^ 


^^^^^^^HC| 

> 

if   ' 

4 

Figure  3.  Stereophotographs  ( x  1 6)  of  segments  1 3-1 5  of  the  prothorax,  the  opisthothorax,  and  the  possible 
pygidium  (p)  of  Peachella  brevispina  Palmer.  The  pygidium  is  poorly  preserved  and  twisted  underneath  and 
to  the  right  so  that  only  the  axis  and  right  pleural  region  are  visible.  Note  also  that  the  axial  ring  of  prothoracic 
segment  1 5  is  broken.  The  base  of  the  spine  that  extended  upward  and  backward  from  this  segment  is 
preserved  on  the  counterpart  to  this  specimen,  and  the  mold  of  this  spine  (s)  enters  the  rock  to  the  left  of 
the  possible  pygidium. 


Pygidium  possibly  present,  although  twisted  to  right  and  only  partly  preserved  on 
internal  and  external  molds.  Overall  shape  broadly  triangular,  with  convex  axis  tapering 
posteriorly  and  extending  almost  to  pygidial  margin.  Pleural  area  smooth,  slightly 
convex,  with  no  border  or  border  furrow. 

Remarks.  — Tht  cephalon  of  Peachella  brevispina  is  characterized  by  its  faintly 
defined  glabella  and  border  furrow,  short  ocular  lobes,  and  short,  moderately  inflated 
paddle-like  genal  spines.  The  cephala  illustrated  by  Palmer  and  Halley  (1979,  pi.  5, 
figs.  1,  2)  are  much  better  preserved  than  the  cephalon  on  this  complete  specimen. 

The  most  unusual  features  of  the  prothorax  are  the  pleurae  of  the  third  segment, 
which  expand  rapidly  away  from  the  axial  furrows  and  become  broadly  oval  in  shape, 
crowding  and  distorting  the  adjacent  pleurae.  Other  unusual  features  include  the  round- 
ed distal  extremeties  of  the  prothoracic  pleurae;  on  most  olenellid  trilobites.  the  pleurae 
terminate  laterally  in  spines.  The  prothoracic  segments  diminish  in  size  posteriorly  and 
somewhat  grade  into  the  segments  of  the  opisthothorax.  although  the  junction  between 
these  two  parts  of  the  thorax  is  not  perfectly  preserved.  Palmer  (in  Palmer  and  Halley 
1979:73)  described  a  partly  preserved  thorax  of  Olenelliis  multinodus  (pi.  4,  figs.  7,  8) 
that  has  enlarged  pleurae  on  the  third  prothoracic  segment  and  a  gradual  transition 
from  the  pleurae  of  the  prothorax  to  the  pleurae  of  the  opisthothorax.  He  suggests  that 
these  features  might  merit  placing  O.  multinodus  in  a  new  genus  (possibly  with  Olenellus 
arcuatus)  if  additional  specimens  prove  that  these  features  are  characteristic  of  the 
species.  These  two  species  also  have  short  ocular  lobes.  Similar  features  are  present  on 
P.  brevispina,  which  has  in  addition  the  unusual  and  generically  distinctive  paddle- 
shaped  genal  spines  not  present  on  O.  multinodus  or  O.  arcuatus.  This  mix  of  similar 


150 


unusual  features  on  species  that  clearly  belong  in  different  genera  illustrates  once  again 
some  of  the  problems  in  generic  level  taxonomy  in  olenellids. 

If  indeed  the  pygidium  of  P.  brevispina  is  preserved  in  a  twisted  position  at  the 
posterior  end  of  the  opisthothorax,  then  P.  brevispina  also  has  an  unusuaLpygidium 
for  an  olenellid  to  add  to  its  other  peculiarities.  Olenellid  pygidia  (when  preserved)  are 
very  small,  short  trapezoidal  plates  attached  to  the  end  of  the  narrow  opisthothorax. 
The  apparent  pygidium  of  P.  brevispina  is  relatively  large,  triangular  in  shape,  and  has 
broad,  smooth  pleural  areas,  a  combination  which  makes  the  pygidium  of  this  species 
unique  among  olenellids. 

Acknowledgments 

The  senior  author  would  like  to  thank  Frederick  Schram  of  the  San  Diego  Natural 
History  Museum  for  suggesting  the  project.  Three  anonymous  reviewers  are  thanked 
for  their  helpful  comments  and  suggestions.  Connie  Egerdahl  is  thanked  for  carefully 
typing  the  manuscript. 

Literature  Cited 


Bergstrom,  Jan.  1973.  Organization,  life,  and  sys- 
tematics  of  trilobites.  Fossils  and  Strata  2:1- 
69. 

Fritz,  W.  H.  1 972.  Lower  Cambrian  trilobites  from 
the  Sekwi  Formation  type  section.  Canadian 
Geological  Survey  Bulletin  212. 

Harrington,  H.  J.  1959.  General  description  of 
Trilobita,  p.  38-1 26  in  R.  C.  Moore  (ed.).  Trea- 
tise on  Invertebrate  Paleontology,  Part  0,  Ar- 
thropoda  1 .  Geological  Society  of  America  and 
the  University  of  Kansas  Press,  Lawrence, 
Kansas. 

Mount,  J.  D.  1980.  An  Early  Cambrian  fauna 
from  the  Carrara  Formation,  Emigrant  Pass, 
Nopah  Range,  Inyo  County,  California:  a  pre- 
liminary note,  p.  78-80  in  Paleontological  Tour 
of  the  Mojave  Desert,  California-Nevada. 
Southern  California  Paleontological  Society 
Special  Publication  No.  2. 


Rasetti,  Franco.  1948.  Lower  Cambrian  trilobites 
from  the  conglomerates  of  Quebec.  Journal  of 
Paleontology  22:1-24. 

Resser,  C.  E.,  and  B.  F.  Howell.  1938.  Lower 
Cambrian  Olenellus  Zone  of  the  Appalachians. 
Geological  Society  of  America  Bulletin  49:195- 
248. 

Palmer,  A.  R.,  and  R.  B.  Halley.  1979.  Physical 
stratigraphy  and  trilobite  biostratigraphy  of  the 
Carrara  Formation  (Lower  and  Middle  Cam- 
brian) in  the  southern  Great  Basin.  United 
States  Geological  Survey  Professional  Paper 
1047. 

Walcott,  CD.  1910.  Olenellus  and  other  genera 
of  the  Mesonacidae.  Smithsonian  Miscella- 
neous Collections  53:233-422. 


TRANSACTIONS 
OF  THE  SAN  DIEGO 
MU8.  COMP.  ZOOL  SOCIETY  OF 

LIBRARY  NATURAL  HISTORY 

JUL    91964 

HARVARD 

UNlVMSiTY, 

Volume  20  Number  1 0  pp.  1 51  -1 64     20  June  1 984 


Type  specimens  of  amphibians  and  reptiles  in  the 
San  Diego  Natural  History  Museum 

Gregory  K.  Pregill  and  James  E.  Berrian 

Department  of  Herpetology,  San  Diego  Natural  History  Museum,  San  Diego,  CA  92112  USA 

Publishing  a  list  of  type  specimens  in  a  collection  serves  several  purposes.  For 
those  who  compile  them,  the  inventory  is  a  propitious  means  of  revealing  the  taxonomic 
and  nomenclatural  errors  and  inconsistencies  that  routinely  creep  into  a  collection.  For 
users,  the  list  provides  a  comprehensive  reference  to  that  institution's  holdings  of 
primary  and  secondary  types,  and  serves  as  a  guide  to  the  original  literature,  perhaps 
even  yielding  insights  into  the  nomenclatural  history  of  the  taxa. 

In  generating  the  present  account,  we  made  use  of  the  only  previous  publication 
of  type  specimens  of  amphibians  and  reptiles  in  the  San  Diego  Natural  History  Museum, 
compiled  by  Allan  J.  Sloan  in  1965  (Transactions  San  Diego  Society  of  Natural  History 
(14(1):  1-8).  Sloan  listed  only  holotypes,  which  at  the  time  numbered  48,  all  of  reptiles. 
To  make  the  present  list  more  comprehensive  we  have  included  all  paratypic  material, 
and  revised  the  now  expanded  list  of  holotypes. 

The  collection  currently  includes  holotypes  of  1  frog,  14  lizards  and  41  snakes.  As 
well,  there  are  several  hundred  paratypes  of  24  additional  taxa.  Locating  all  secondary 
types  in  the  collection  was  a  difficult  task  because  many  of  these  specimens  had  not 
been  so  designated  in  the  early  years  of  cataloging,  nor  were  they  segregated  from  the 
main  collection.  Subspecies  comprise  the  majority  of  primary  and  secondary  types, 
many  of  which  have  been  synonymized  since  their  original  description.  Readers  familiar 
with  the  collections  of  amphibians  and  reptiles  at  the  San  Diego  Natural  History 
Museum  are  aware  that  the  geographic  emphasis  favors  the  Southwest,  Baja  California 
and  the  Gulf  of  California  Islands;  this  is  reflected  in  the  type  localities  that  follow. 

For  many  years  the  collections  of  amphibians  and  reptiles  at  the  San  Diego  Natural 
History  Museum  were  closely  tied  to  the  activities  of  Laurence  M.  Klauber,  although 
the  full  history  extends  beyond  the  KJauber  era  back  to  the  Society's  inception  in  1 874. 
Klauber  began  his  herpetological  studies  as  an  avocation  in  1920,  conducting  most  of 
his  research  and  maintaining  the  collections  in  the  basement  of  his  home.  By  1960  he 
had  amassed  35  000  specimens.  These,  as  well  as  his  vast  library  of  over  20  000  books 
and  reprints,  were  donated  to  the  San  Diego  Society  of  Natural  History,  whence  they 
were  housed  in  the  Natural  History  Museum  in  Balboa  Park.  At  the  time  of  Klauber's 
death  in  1968,  the  number  of  specimens  of  amphibians  and  reptiles  at  the  museum 
was  over  50  000. 

Over  the  years  the  collection  has  had  many  contributors,  among  the  earliest  being 
the  celebrated  naturalists  of  the  Southwest,  Charles  R.  Orcutt  and  Frank  Stephens. 
Later,  during  the  middle  third  of  this  century,  the  collection  benefited  considerably 
from  the  efforts  of  C.  B.  Perkins  and  Charles  E.  Shaw.  Klauber's  legendary  interest  in 
rattlesnakes  alone  resulted  in  an  enormous  series  of  Crotalus  and  a  substantial  repre- 
sentation of  reptiles  native  to  the  Far  West  and  northern  Mexico.  Indeed,  during  the 
1 10-year  history  of  the  Society,  the  museum  has  benefited  from  the  interests  and  travels 
of  many  individuals.  Hence,  the  collections  of  amphibians  and  reptiles  are  flavored 


152 


with  series  of  taxa  from,  for  example,  the  Galapagos  Islands,  the  Pacific,  and  the 
Caribbean. 

As  eariy  as  1928,  Klauber  foresaw  that  someday  his  collection  and  that  of  the 
museum  would  merge,  and  he  assigned  specimen  numbers  accordingly  so*  that  they 
would  not  have  to  be  recatalogued  at  a  future  date.  Consequently,  all  specimens  orig- 
inally in  the  LMK  series  have  the  same  numbers  in  the  herpetological  catalogue  of  the 
San  Diego  Natural  History  Museum.  The  collection  now  has  approximately  65  000 
specimens  which  may  be  broken  down  taxonomically  as  follows: 

Amphibians  (Apoda,  Caudata,  Anura)— 18  families,  60  genera,  175  species 

Turtles— 9  families,  34  genera,  80  species 

Lizards  (including  amphisbaenians  and  Sphenodon)—  1 7  families,  1 30  genera, 

360  species 
Snakes— 9  families,  195  genera,  400  species 

These  taxa  also  encompass  nearly  2000  skeletons  in  roughly  the  same  diversity,  and 
several  hundred  preserved  hemipenes  representing  70  species  of  snakes  (mostly  Cro- 
talus)  and  lizards. 

About  This  List 

Each  species  entry  begins  with  the  author,  date  and  publication  of  the  original 
description;  the  specific  page  number  where  the  description  appears  is  given  paren- 
thetically at  the  end  of  that  citation.  This  line  is  followed  by  the  SDSNH  number  of 
the  holotype,  sex,  type  locality  and  the  name  of  the  collector  and  date  that  the  specimen 
was  caught.  In  nearly  all  instances,  we  have  tried  to  preserve  the  original  wording  for 
the  type  locality.  Where  potential  confusion  exists  because  of  orthography  or  archaic 
and  vague  place  names,  we  have  clarified  them  as  needed,  inside  brackets  [  ]  or  at  the 
bottom  of  the  entry  under  Remarks.  When  only  the  paratype(s)  of  a  taxon  is  represented 
in  the  SDSNH  collection,  the  holotype  and  catalogue  number  of  the  holding  institution 
is  given  under  Remarks. 

Each  taxon  is  listed  alphabetically  by  genus,  species  and  subspecies  precisely  as  it 
was  originally  designated,  that  is,  irrespective  of  present  usage.  For  those  taxa  that  have 
been  synonymized  or  whose  nomenclature  is  currently  in  doubt,  this  is  so  noted  under 
Remarks  with  the  name  and  publication  of  the  first  authority  or  re  visor.  We  have  made 
no  nomenclatural  decisions  per  se,  and  a  particular  taxon  is  considered  valid  unless 
indicated. 

Abbreviations  of  museums  and  personal  collections  are  as  follows: 

AMNH:  American  Museum  of  Natural  History 

BYU:  Brigham  Young  University 

CAS:  California  Academy  of  Sciences 

CAS— SU:  California  Academy  of  Sciences— Stanford  University  collection 

CM:  Carnegie  Museum 

EHT:  Edward  H.  Taylor  collection* 

FMNH:  Field  Museum  of  Natural  History 

LACM:  Los  Angeles  County  Natural  History  Museum 

MCZ:  Museum  of  Comparative  Zoology,  Harvard  University 

SDSNH:  San  Diego  Society  of  Natural  History 

SU:  Stanford  University  Natural  History  Museum* 

T-S:  Edward  H.  Taylor  and  Hobart  M.  Smith,  University  of  Kansas* 

UIMNH:  University  of  Illinois,  Museum  of  Natural  History 

UMMZ:  University  of  Michigan,  Museum  of  Zoology 

USNM:  National  Museum  of  Natural  History 


*  Specimens  from  these  collections  have  been  dispersed  to  other  institutions. 


153 

Anura 

Bufo  exsul  Myers  (1942) 

Occ.  Papers  Mus.  Zool.  Univ.  Michigan  460:1-13  (p.  3). 

Paratypes. -SDSNH  29098  and  29099  Deep  Springs,  Inyo  Co.,  California. 
Remarks.  — =  Bufo  boreas  exsul  2iCcord\ng  to  Stebbins  (1962,  Amphibians  of  West- 
em  North  America.  Univ.  Calif.  Press).  Holotype:  UMMZ  83357  a  very  large  female. 

Hyla  regilla  cascadae  Jameson,  Mackay  and  Richmond  (1966) 
Proc.  California  Acad.  Sci.  4th  ser.  33(19):55 1-620  (p.  602). 

Paratypes. -SDS^H  44971-44987  '/2  mi  S  of  Bend,  Deschutes  Co.,  Oregon. 
Remarks.  — Ho\o\y^Q:  CAS  101038  an  adult  male. 

Hyla  regilla  deserticola  Jameson,  Mackey  and  Richmond  (1966) 
Proc.  California  Acad.  Sci.  4th  ser.  33(1 9):55 1-620  (p.  582). 

Holotype. —  SDSiNH  54176  an  adult  male;  San  Borjas,  Baja  California  Norte,  Mex- 
ico. By  David  L.  Jameson,  November  25,  1961. 

Paratypes. —  SY>SN\\  54166-54175  data  same  as  for  Holotype. 

Remarks.  — =Hyla  regilla  hypochondriaca  according  to  Duellman  (1970,  Univ. 
Kansas  Mus.  Nat.  Hist.  Monog.  No.  1:487).  San  Borjas  reads  San  Borja  on  recent  maps. 

Hyla  regilla  pacifica  Jameson,  Mackey  and  Richmond  (1966) 
Proc.  California  Acad.  Sci.  4th  ser.  33(19):55 1-620  (p.  591). 

Paratypes. -Sr>S>HH  53514-53529  4  mi  S  of  Waldport,  Lincoln  Co.,  Oregon. 
Remarks.  — HoXoXypt:  CAS  101007  an  adult  male. 

Hyla  regilla  palouse  Jameson,  Mackey  and  Richmond  (1966) 
Proc.  CaUfomia  Acad.  Sci.  4th  ser.  33(19):55 1-620  (p.  599). 

Paratypes. -SUSHH  44715-44718  6  mi  SE  of  La  Grande,  Union  Co.,  Oregon. 
Remarks.  — YioXoXypQ:  CAS  100982  an  adult  male. 

Hyla  regilla  sierrae  Jameson,  Mackey  and  Richmond  (1966) 

Proc.  California  Acad.  Sci.  4th  ser.  33(19):55 1-620  (p.  605). 

Paratypes. -^\:>S^H  53835-53841   I'/*  mi  SSE  of  Tioga  Pass  Ranger  Station.  E 
entrance  to  Yosemite  National  Park,  Mariposa  Co.,  California. 
Remarks.  — HoXoly^Q-.  CAS  100991  an  adult  male. 

Sauria 

Anniella  geronimensis  Shaw  (1940) 

Trans.  San  Diego  Soc.  Nat.  Hist.  9(24):225-228  (p.  225). 

Holotype. —  SUS^H  7543  an  adult  female;  San  Geronimo  Island,  Lower  California 
[Norte],  Mexico.  By  Margaret  [Mrs.  Griffing]  Bancroft,  March  28,  1932. 
Paratypes.  — 'S>r>'S>N\\  7542  data  same  as  for  Holotype. 
Remarks.  — =Anniella  pulchra  according  to  Hunt  (1983,  Copeia  (l):79-89). 

Anolis  rivalis  Williams  (1984) 
Breviora  478:1-22  (p.  7). 

Paratype. -SDSNH  31 163  "Port  Utria,"  [south  of  Punta  Solano,  Choco]  Colom- 
bia. 

Remarks.  —  Holotype:  LACM  42124  an  adult  male. 

Coleonyx  variegatus  abbotti  Klauber  (1945) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(1 1):  133-2 16  (p.  154). 

Holotype.  — SYySnW  34790  an  adult  male;  Proctor  Valley,  San  Diego  Co..  Cali- 
fornia. By  William  Moore,  February  28,  1942. 

Paratypes.- As  follows:  California:  Los  Angeles  Co.:  SDSNH  201  1  San  Francis- 
quito  Plant  2.  Riverside  Co.:  SDSNH  2725  Moreno.  San  Diego  Co.:  SDSNH  30  Cot- 


154 

tonwood;  SDSNH  843  Foster;  SDSNH  16702  Rincon;  SDSNH  16988,  32817,  34786 
Mission  Gorge;  SDSNH  1 6989  Jamui;  SDSNH  1 70 1 2  De  Luz  (says  "Sentenac  Canyon" 
in  catalogue);  SDSNH  21249  El  Capitan;  SDSNH  24050  San  Pasqual;  SDSNH  25303, 
34666  Jacumba;  SDSNH  27770  Foot  Agua  Tibia  Mt.;  SDSNH  32797  Pala;  SDSNH 
32821-32822  Black  Mtn.  near  La  Mesa.  Baja  California  Norte,  Mexico:  SDSNH  2593 
Ensenada;  SDSNH  5265-5266,  15970-15971,  27726,  30295  Cedros  Island;  SDSNH 
6553  65  mi  SE  of  Tecate;  SDSNH  24390  San  Jose  (lat.  32°). 

Coleonyx  variegatus  bogerti  Klauber  (1943) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(1 1):133-216  (p.  176). 

Ho/otvpe.  — SDSNH  32486  an  adult  male;  Xavier,  Pima  Co.,  Arizona.  By  Lee  W. 
Arnold,  July  17,  1939. 

Paratypes.— None  designated. 

Coleonyx  variegatus  utahensis  Klauber  (1943) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(1 1):133-216  (p.  171). 

Holotype.  — SDSNH  35792  an  adult  male;  Watercress  Spring,  Washington  Co., 
Utah.  By  Dr.  Ross  Hardy,  April  16,  1941. 

Paratypes.  — SDSNH  36021-36024  data  same  as  for  Holotype. 

Phyllodactylus  angelensis  Dixon  (1966) 

Proc.  Cahfomia  Acad.  Sci.  4th  ser.  33(13):4 15-452  (p.  444). 

Holotype.  — SDSNH  19996  an  adult  male;  north  end  of  Isla  Angel  de  la  Guarda, 
Baja  California  [Norte],  Mexico.  By  Allan  J.  Sloan,  March  22,  1963. 

Paratype.  — SDSNH  50851  Isla  Pond,  [Baja  California  Norte,  Mexico]. 

Phyllodactylus  apricus  Dixon  (1966) 

Proc.  California  Acad.  Sci.  4th  ser.  33(1 3):4 15-452  (p.  450). 

Holotype.  — SDSNH  44623  an  adult  male;  Isla  Las  Animas,  Baja  California  [Sur, 
Mexico].  By  Chris  Parrish  and  G.  E.  Lindsay,  June  27,  1964. 

Paratypes.-SDSNH  44620-44622,  44624,  50830-50842,  50844-50849  data  same 
as  for  Holotype. 

Phyllodactylus  bugastrolepis  Dixon  (1966) 

Proc.  Cahfomia  Acad.  Sci.  4th  ser.  33(13):4 15-452  (p.  447). 

Paratypes.-SDSNH  44604-44607,  44611-44613,  50792,  50793  Isla  Catalina, 
Baja  California  [Sur,  Mexico]. 

Remarks.  — HoXoXyTpe:  CAS  98485  an  adult  female. 

Phyllodactylus  homolepidurus  nolascoensis  Dixon  (1964) 
New  Mexico  State  Univ.  Sci.  BuH.  64(1):  1-1 39  (p.  42). 

Paratypes.-SDSNH  6840  and  6841  Isla  San  Pedro  Nolasco,  Sonora,  Mexico. 
Remarks.  — Ho\o\ype:  CAS  50552  an  adult  male. 

Phyllodactylus  partitus  Dixon  (1966) 

Proc.  California  Acad.  Sci.  4th  ser.  33(13):4 15-452  (p.  445). 

Paratypes.-SDSNH  6834-6836,  39258,  39649,  40508,  50820-50822  Isla  Partida 
(N),  Baja  California  [Norte,  Mexico]. 

Remarks.  — HoXoXype:  CAS  98429  an  adult  male. 

Phyllodactylus  santacruzensis  Dixon  (1966) 

Proc.  California  Acad.  Sci.  4th  ser.  33(13):4 15-452  (p.  449). 

Paratypes.-SDSNH  50872,  50873  Isla  Santa  Cruz,  Baja  California  [Sur,  Mexico]. 
Remarks. -Holotype:  CAS  98468  an  adult  female. 

Phyllodactylus  xanti  acorius  Dixon  (1966) 

Proc.  Cahfomia  Acad.  Sci.  4th  ser.  33(13):4 15-452  (p.  442). 

Paratypes.-SDSNH  50827-50829  Isla  San  Diego,  Baja  California  [Sur,  Mexico]. 


155 

Remarks.  — =Phyllodactyhis  nocticolus  acohiis  according  to  Murphy  (1983,  Occ. 
Papers  California  Acad.  Sci.  137:1-48).  Holotype:  CAS  98451  an  adult  male. 

Phyllodactylus  xanti  angulus  Dixon  (1966) 

Proc.  California  Acad.  Sci.  4th  ser.  33(13):4 15-452  (p.  433). 

Paratypes.-SX:>^NH  AAbll ,  44678,  50868,  50869  Isla  Salsipuedes,  Baja  California 
[Norte,  Mexico];  SDSNH  44238,  44239,  50833,  50856-50859  [Isla]  San  Lorenzo,  [Baja 
California  Norte,  Mexico]. 

Remarks.  — =PhyUodactylus  nocticolus  angulus  according  to  Murphy  (1983,  Occ. 
Papers  California  Acad.  Sci.'  137:1-48).  Holotype:  CAS  98477  an  adult  male. 

Phyllodactylus  xanti  circus  Dixon  (1966) 

Proc.  California  Acad.  Sci.  4th  ser.  33(13):4 15-452  (p.  439). 

Holotype. —  SUS^H  50814  an  adult  female;  Isla  Ildefonso  [Baja  California  Sur, 
Mexico].  By  Charles  E.  Shaw,  April  2,  1962. 

Paratypes.-^X:>Sn\\  50809-50812,  50815-50817  data  same  as  for  Holotype. 

Remarks.  — ^Phyllodactylus  nocticolus  circus  according  to  Murphy  (1983,  Occ. 
Papers  California  Acad.  Sci.  137:1-48). 

Phyllodactylus  xanti  estebanensis  Dixon  (1966) 

Proc.  California  Acad.  Sci.  4th  ser.  33(13):4 15-452  (p.  437). 

Paratypes.-SX:>S^H  50852  and  50853  Isla  Tiburon  [Gulf  of  California,  Mexico]; 
50870  and  50871  Isla  San  Esteban  [Gulf  of  California,  Mexico]. 

Remarks.  — =  Phyllodactylus  nocticolus  estebanensis  according  to  Murphy  (1983, 
Occ.  Papers  California  Acad.  Sci.  137:1-48).  Holotype:  CAS  98481  an  adult  male.  Isla 
Tiburon  and  Isla  Esteban  belong  to  the  state  of  Sonora, 

Phyllodactylus  xanti  nocticolus  Dixon  (1964) 

New  Mexico  State  Univ.  Sci.  Bull.  64(1):  1-1 39  (p.  55). 

Holotype.  SY^S^W  34824  an  adult  male;  Agua  Caliente  Hot  Springs,  San  Diego 
Co.,  California.  By  Laurence  M.  Klauber  [Charles  Shaw  and  Paul  Breese].  Preserved 
March  8,  1942. 

Paratypes.-¥rom  San  Diego  County,  as  follows:  SDSNH  2952,  18633,  32955, 
35476-35490,  40073  Yaqui  Well;  SDSNH  18632,  33247  Sentenac  Canyon;  SDSNH 
1 892 1  Yaqui  Pass;  SDSNH  33672-33674  Palm  Canyon;  SDSNH  34825-34827, 43786, 
43787  Topotypes;  SDSNH  38034,  38041  4  mi  W  of  Vallecitos  Stage  Station. 

Remarks.  —  =  Phyllodactylus  nocticolus  nocticolus  according  to  Murphy  (1983,  Occ. 
Papers  California  Acad.  Sci.  137:1-48). 

Phyllodactylus  xanti  sloani  Bostic  (1972) 

Trans.  San  Diego  Soc.  Nat.  Hist.  16(10):237-263  (p.  252). 

Holotype. -SDSNH  45895  an  adult  female;  23.5  mi  SE  of  El  Rosario,  Baja  Cal- 
ifornia Norte,  Mexico.  By  Dennis  L.  Bostic,  March  31,  1969. 

Paratypes.- AW  from  Baja  California  Norte  as  follows:  SDSNH  45896  1.8  mi  NW 
of  Puerto  de  San  Carlos:  SDSNH  45897  and  45898  5.4  mi  W  of  Punta  Canoas;  SDSNH 
45899  and  45900  Arroyo  San  Jose;  SDSNH  4590 1-45907  Las  Palomas;  SDSNH  45908 
EI  Cardon;  SDSNH  45909-45912  5  mi  N  of  San  Javier. 

Remarks.  — =  Phyllodactylus  nocticolus  sloani  according  to  Murphy  (1983,  Occ. 
Papers  California  Acad.  Sci.'  137:1-48).  SDSNH  45912  was  "sent  to  Mexico,  1971" 
[presumably  to  Fauna  Silvestre,  Mexico  City]. 

Sauromalus  australis  Shaw  (1945) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(15):269-306  (p.  286). 

Holotype.  — SDSNH  30170  an  adult  male;  San  Francisquito  Bay,  Baja  California, 
Mexico.  By  Robert  S.  Hoard,  July  30,  1938. 

Paratypes.  — W\  are  from  Baja  California  Sur  as  follows:  SDSNH  1 7707  Comondu; 
SDSNH  17708  La  Paz;  SDSNH  30168  Loreto;  SDSNH  30169  33  mi  N  of  Canipole. 


156 


Sauromalus  klauberi  Shaw  (1941) 

Trans.  San  Diego  Soc.  Nat.  Hist.  9(28):285-288  (p.  285). 

//o/ory;?£'.  — SDSNH  6859  an  adult  male;  Santa  Catalina  Island,  Gulf  of  California 
[Baja  California  Sur],  Mexico.  By  J.  R.  Pemberton,  December  14,  1931.       * 

Paratypes.  —  SDSHW  6860  and  6861  data  same  as  for  Holotype. 

Remarks.  — ^Sauromulus  ater  klauberi  according  to  Avery  and  Tanner  (1964, 
BYU  Science  Bull.  5(3):  1). 

Sauromalus  obesus  tumidus  Shaw  (1945) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(15):269-306  (p.  292). 

//o/or.vp^— SDSNH  27323  an  adult  male;  Telegraph  Pass,  Gila  Mountains,  Yuma 
Co.,  Arizona.  By  Laurence  M.  Klauber,  June  15,  1937. 

Paratypes.-ST:>S^H  8613,  27551,  33170-33175,  33224,  33225,  34141,  35090 
data  same  as  for  Holotype  (Gila  Mountains,  Arizona). 

Uma  notata  cowlesi  Heifetz  (1941). 
Copeia  1941(2):99-1 1 1  (p.  104). 

Paratypes.  —  'SDS^W  16460-16464  Punta  Penasco,  Sonora,  Mexico. 
Remarks.  — =  Uma  notata  rufopimctata  Cope  according  to  Norris  (1958,  Bull.  Amer. 
Mus.  Nat.  Hist.  1 14(3):25 1-326).  Holotype:  CAS  53370  an  adult  male. 

Uta  stansburiana  klauberi  Lowe  and  Norris  (1955) 
Hereptologica  ll(2):89-96  (p.  91). 

Holotype.  — SDS^H  6642  an  adult  male;  San  Esteban  Island,  Gulf  of  CaUfomia, 
Sonora,  Mexico.  By  J.  R.  Pemberton,  January  1 1,  1932. 

Paratypes.-Sr>%Nn  3968-3971,  6640,  6641  San  Esteban  Island,  Sonora,  Mexico. 

Remarks.  — =  Uta  stansburiana  tavlori  Smith  according  to  Ballinger  and  Tinkel 
(1972,  Misc.  Pub.  Mus.  Zool.  Univ.  Michigan  (145):  1-83). 

Xantusia  arizonae  Klauber  (1931) 

Trans.  San  Diego  Soc.  Nat.  Hist.  7(1):  1-1 6  (p.  3). 

Holotype.  SY^SNYi  5433  an  adult  female;  1  mi  S  of  Yamell,  Yavapai  Co.,  Arizona. 
By  Philip  M.  Klauber  and  Laurence  M.  Klauber,  August  21,  1931. 

Paratypes.-S\:>^^H  5434-5438  data  same  as  for  Holotype;  SDSNH  5451  and 
5452  bom  (dead)  of  Holotype  in  captivity;  SDSNH  5450  unborn  embryo  from  SDSNH 
5436:  SDSNH  5453  unborn  embryo  from  SDSNH  5434. 

Remarks.  — =Xantusia  vigilis  arizonae  according  to  Bezy  (1967  Copeia  (3):653- 
661).  SDSNH  5434  was  sent  to  J.  R.  Slevin  at  the  California  Academy  of  Sciences 
December  14,  1931. 

Serpentes 

Arizona  elegans  blanchardi  Klauber  (1946) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(17):31 1-398  (p.  328). 

Paratype.  — SDSNH  35343  Schramm,  Yuma  Co.,  Colorado. 

Remarks. -WoXoXy^Q:  SU  10393  a  young  adult  male;  now  CAS-SU  10393. 

Arizona  elegans  Candida  Klauber  (1946) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(17):31 1-398  (p.  364). 

Holotype.  —  SDSNH  34191  an  adult  male;  Kramer  Hills,  6  mi  S  of  Kramer  Junction 
on  US  395,  San  Bernardino  Co.,  California.  By  James  Deuel,  June  16,  1941. 

Paratvpes.  —  M\  are  from  near  the  type  locality  in  San  Bernardino  Co.,  California, 
as  follows:  Adelanto:  SDSNH  2725 1 ,  34 1 87,  34 1 90  1 6  mi  N  of;  SDSNH  28848,  33980 
8  mi  N  of;  SDSNH  3 1 9 1 7  1 2  mi  S  of;  SDSNH  3 1 94 1 ,  3 1 942,  35 109  3  mi  N  of;  SDSNH 
33323  6  mi  SW  of;  SDSNH  338 1 2  6  mi  N  of;  SDSNH  3398 1 ,  35093  4  mi  S  of;  SDSNH 
33982  14  mi  N  of;  SDSNH  34107  13  mi  N  of;  SDSNH  34150,  35594  2  mi  N  of 
Kramer  Hills:  SDSNH  28846,  31700,  33795,  35149.  Kramer  Junction:  SDSNH  31766 


157 

5  mi  N  of;  SDSNH  31940,  33832,  33983,  34019  8  mi  S  of;  SDSNH  31959  20  mi  S 
of;  SDSNH  33826,  34189  3  mi  S  of;  SDSNH  33888,  34164,  34192  5  mi  S  of;  SDSNH 
33975,  34017,  35536  7  mi  S  of;  SDSNH  33977,  33978  10  mi  S  of;  SDSNH  33979  12 
mi  S  of;  SDSNH  34165  6  mi  of;  SDSNH  34184  1  mi  W  of;  SDSNH  34185  1  mi  E  of; 
SDSNH  34186  1 1  mi  S  of;  SDSNH  35151,  35537  9  mi  S  of  SDSNH  35654  6.5  mi 
Sof 

Arizona  elegans  eburnata  Klauber  (1946) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(17):31 1-393  (p.  350). 

Holotype.  —  ^Y^SNH  33094  a  young  male;  Benson's  Dry  Lake,  San  Diego  Co., 
California.  By  James  Deuel,  June  5,  1940  [preserved]. 

Paratvpes.  —  M\  are  from  near  the  type  locality  in  San  Diego  Co.,  California,  as 
follows:  SDSNH  4454,  23854,  23914,  23915,  25437,  25438,  26914,  26939-26942, 
27308,  27383,  27405,  29300,  33094,  33095,  17026  Topotypes;  SDSNH  4862  Beatty 
Ranch,  Borrego  Valley;  SDSNH  5136,  21 108,  21 121  Borrego  Valley;  SDSNH  23024, 
23774-23776,  23852,  23853,  26814,  29301,  29487,  32035  The  Narrows;  SDSNH 
23773  5  mi  E  of  The  Narrows;  SDSNH  26732  2  mi  S  of  Borrego  Post  Office;  SDSNH 
27331  3  mi  W  of  Benson's  Dry  Lake. 

Remarks. -SiDS^H  4454,  17026,  21 108  were  exchanged  with  H.  M.  Smith,  Jan- 
uary 14,  1957  and  are  presumed  to  be  at  the  University  of  Illinois  Museum  of  Natural 
History.  KJauber  also  gave  as  paratypes  SDSNH  26056  and  26057,  but  these  are  actually 
specimens  of  Pituophis  melanoleucus  deserticola  and  Crotalus  viridis  oreganus,  respec- 
tively; the  source  of  this  error  is  unknown. 

Arizona  elegans  noctivaga  Klauber  (1946) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(17):31 1-398  (p.  343). 

Holotype. —  SDSNH  34188  a  young  adult  male;  8  mi  N  of  Owlshead,  Pinal  Co., 
Arizona.  By  Charles  E.  Shaw  and  Laurence  M.  Klauber,  May  31,  1941. 

Paratypes.— AW  are  from  Arizona,  as  follows:  Gila  Co.:  SDSNH  34438  7  mi  SE 
of  Globe.  Maricopa  Co.:  SDSNH  15835,  23925,  23926  Sentinel;  SDSNH  25829,  260 1 0 
Mesa;  SDSNH  26913  5  mi  S  of  Wickenburg;  SDSNH  26943  Cactus  Gardens;  SDSNH 
32781  Gila  Bend;  SDSNH  34331  Stanwix.  Pima  Co.:  SDSNH  13724,  29222  Tucson: 
SDSNH  1 7949  1 1  mi  N  of  Tucson;  SDSNH  1 7950,  17951  4V2  mi  N  of  Tucson;  SDSNH 
17952  2  mi  N  of  Tucson;  SDSNH  27177  13  mi  N  of  Tucson;  SDSNH  27178  4  mi 
N  of  Sahuarita;  SDSNH  32293  Martinez  Hill;  SDSNH  32521  1 1  mi  S  of  Tucson; 
SDSNH  32714  2  mi  NE  of  Tanque  Verde  Ranch;  SDSNH  34018  14  mi  N  of  Tucson. 
Pinal  Co.:  SDSNH  21492,  21493  Picacho;  SDSNH  27180  Rorence;  SDSNH  32323  8 
mi  W  of  Casa  Grande;  SDSNH  34104  3  mi  W  of  Superior;  SDSNH  34332  Oracle 
Junction.  Yavapai  Co.:  SDSNH  1 7623  1  mi  S  of  Congress  Junction.  Yuma  Co.:  SDSNH 
34526  5  mi  E  of  Salome. 

Arizona  elegans  pacata  Klauber  (1946) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(1 7):31 1-398  (p.  379). 

Holotype. -SDSNH  17652  an  adult  male;  Santo  Domingo  (lat.  25°30'N),  Baja 
California  Sur,  Mexico.  By  Frank  F.  Gander,  November  16,  1941. 

Arizona  elegans  philipi  Klauber  (1946) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(1 7):31 1-398  (p.  333). 

Holotype.  — SDSNH  34456  an  adolescent  male;  10  mi  E  of  Winslow,  Navajo  Co., 
Arizona.  By  Charles  E.  Shaw  and  Carl  Engler,  July  29,  1941. 

Paratypes. -SDSNH  20990  Two  Guns,  Coconino  Co.,  Arizona;  SDSNH  34426 
data  same  as  for  Holotype. 

Charina  bottae  umbratica  Klauber  (1943) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(7):83-90  (p.  83). 

Holotype.  — SDSNH  12101  an  immature  male;  Fern  Valley,  near  Idyllwild,  Riv- 
erside Co.,  California.  By  Clyde  Searl,  July  1,  1929. 


158 

Remarks.— The  validity  of  C.  b.  umbratica  as  a  subspecies  has  been  questioned 
on  several  occasions,  whereas  in  other  instances  full  species  status  has  been  proposed. 
The  taxonomic  history  is  treated  by  Stewart  (1977,  SSAR  Cat.  Amer.  Amph.  Rept. 
205.2). 

Chilomeniscus  stramineus  esterensis  Hoard  (1939) 

Pomona  College  Jour.  Ent.  and  Zool.  31(4):45-46  (p.  45). 

//o/o/V/?^— SDSNH  30368  an  adult  male;  Estero[s]  Salina,  Lower  [Baja]  California, 
Mexico  [24°36'N,  1 1 1°49'W].  By  R.  S.  Hoard,  July  10,  1938. 

/'ararv/?6'5.-SDSNH  30364-30367,  30369,  30370  data  same  as  for  Holotype. 

Chionactis  occipitalis  talpina  Klauber  (1951) 

Trans.  San  Diego  Soc.  Nat.  Hist.  1 1(9):  141-204  (p.  172). 

Paratypes.-SY:>S^\l  39520,  39521  10  mi  N  of  Goldfield,  Esmerelda  Co.,  Nevada. 
Remarks.  — WoXoXype:  CAS  81364  an  adult  male. 

Chionactis  palarostris  organica  Klauber  (1951) 

Trans.  San  Diego  Soc.  Nat.  Hist.  1 1(9):  141-204  (p.  178). 

Holotype. —SD^^W  40673  an  adult  male;  on  the  Sonoyta-Ajo  road,  9  mi  N  of  the 
U.S. -Mexican  border,  in  Organ  Pipe  Cactus  National  Monument,  Pima  Co.,  Arizona. 
By  William  R.  Supernaugh  and  Grover  E.  Steele,  May  22,  1950. 

Crotalus  cerastes  cercobombus  Savage  and  Cliff  (1953) 
Nat.  Hist.  Misc.  1 19:1-7  (p.  2). 

Paratvpes.—M\  from  Arizona,  as  follows:  Maricopa  Co.:  SDSNH  979-981  18  mi 
W  of  Phoenix;  SDSNH  17072  10  mi  E  of  Gila  Bend;  SDSNH  22410-22412  Desert 
Wells;  SDSNH  23879,  23888-23890,  25553-25555,  25854-25857  near  Mesa;  SDSNH 
26915  1  mi  S  of  Morristown;  SDSNH  39082  26  mi  N  of  Ajo;  SDSNH  39088  1  mi  W 
of  Tartron  [sic]  (see  Remarks);  SDSNH  40893  vicinity  of  Phoenix.  Pima  Co.:  SDSNH 
2324,  2325  Sells;  SDSNH  38660  Sonoita  (see  Remarks).  Pinal  Co.:  SDSNH  17068- 
17071,  17073  3  mi  SE  of  Picacho;  SDSNH  25499  5  mi  W  of  Casa  Grande;  SDSNH 
41110-41116,41 362,  4 1 363  4  mi  E  of  Coolidge;  SDSNH  4 1 24 1  4  mi  S  of  Coolidge. 

Remarks.  — WoXoXyi^e:  SU  7287  an  adult  male;  now  CAS-SU  7287.  The  correct 
spelling  is  Tarton.  Sonoita  is  in  Santa  Cruz  Co. 

Crotalus  cerastes  laterorepens  Klauber  ( 1 944) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(8):9 1-126  (p.  94). 

Holotype.  —  SDSNH  34074  an  adult  male;  The  Narrows,  San  Diego  Co.,  California. 
By  Cyrus  B.  Perkins  and  Charles  Shaw,  June  6,  1941. 

Par  at  vpes.— All  are  from  within  a  13-mile  radius  of  the  type  locality  as  follows: 
SDSNH  1762,  1858,  2209,  2210,  4571,  4572,  4645-4647,  4827,  4875,  4931,  4958, 
5052-5055,  5173-5175,  9507,  21098,  21426,  22275,  22357,  23009,  23233,  23640, 
23858,  23860,  23917,  23954-23956,  23998,  24006,  24019,  24020,  25423,  25445, 
26729-26731,  26823,  26847,  26848,  26857,  26865,  26866,  26937,  26938,  27240, 
28113,  28228,  28682,  28683,  28728,  28750,  29085,  29118,  29119,  29271,  29277, 
29898,  30719,  31930,  31999,  32307,  32977,  32978,  33044,  33058,  33059,  33123, 
33333-33335,  33342,  34035,  34176-34179,  34351,  34570,  35179,  35187,  35188,  35305, 
35557-35559,  35597,  35634,  35635. 

Crotalus  confluentus  abyssus  Klauber  (1930) 

Trans.  San  Diego  Soc.  Nat.  Hist.  6(3):95-144  (p.  1 14). 

Holotype.  — SDSNH  2216  an  adult  male;  Tanner  Trail,  300  feet  below  south  rim 
of  the  Grand  Canyon,  Coconino  Co.,  Airzona.  By  E.  D.  McKee,  September  15,  1929. 

Paratypes.— None  designated. 

Remarks.  —=Crotalus  viridis  abvssus  according  to  Klauber  (1936,  Trans.  San  Diego 
Soc.  Nat.  Hist.  8(20):  185-276). 


159 


Crotalus  confluentus  kellyi  Amaral  (1929) 

Bull.  Antivenin  Inst.  Amer.  2(4):86-97  (p.  91). 

Holotvpe.  —  SX^S^H  194  a  male;  Needles  [San  Bernardino  Co.],  California.  By  Mr. 
O.  R.  West,  July  11,  1926. 

Paratype.  —  SV>S,N\\  195  data  same  as  for  Holotype. 

Remarks.  — =Cwtalus  s.  scutulatus  according  to  Klauber  (1930.  Trans.  San  Diego 
Soc.  Nat.  Hist.  6(3):95-144. 

Crotalus  confluentus  lutosus  Klauber  (1930) 

Trans.  San  Diego  Soc.  Nat.  Hist.  6(3):95-144  (p.  100). 

Holotvpe.  —  SDSNH  1814a  young  adult  male;  10  mi  NW  of  Abraham  on  the  road 
to  Joy,  Millard  Co.,  Utah.  By  Cyrus  B.  Perkins,  May  12,  1929. 

Paratvpes.-SDSnYl  1800-1813,  1815,  1816  20  mi  NW  of  Delta,  Millard  Co., 
Utah. 

Remarks.  — =Crotalus  viridis  lutosus  according  to  Klauber  (1936,  Trans.  San  Diego 
Soc.  Nat.  Hist.  8(20):  195-276). 

Crotalus  confluentus  nuntius  Klauber  (1935) 

Trans.  San  Diego  Soc.  Nat.  Hist.  8(13):75-90  (p.  78). 

//c»/o/y/7£'.  — SDSNH  3105  an  adult  male;  Canyon  Diablo,  Coconino  Co.,  Arizona. 
By  R.  L.  Borden,  August  9,  1930. 

Remarks.  —=CrotaIus  viridis  nuntius  according  to  Klauber  (1936,  Trans.  San  Diego 
Soc.  Nat.  Hist.  8(20):  185-276). 

Crotalus  durissus  culminatus  Klauber  (1952). 
Bull.  Zool.  Soc.  San  Diego  26:1-143  (p.  67). 

Para/yp^.— SDSNH  43403  Hacienda  El  Sabino,  near  Uruapan,  Michoacan,  Mex- 
ico. 

Remarks.  — YioXoXyTpQ-.  FMNH  126616  (formerly  EHT  5224)  a  juvenile  female. 
SDSNH  43403  was  formerly  EHT  5233. 

Crotalus  mitchelli  angelensis  Klauber  (1963) 

Trans.  San  Diego  Soc.  Nat.  Hist.  13(5):73-80  (p.  75). 

//o/oO'P^.- SDSNH  51994  an  adult  male;  4  mi  SE  of  Refugio  Bay,  Isla  Angel  de 
la  Guarda,  Gulf  of  California,  Mexico.  By  Dr.  Reid  Moran,  March  22,  1963. 

Paratypes.-^U^NH  19717,  19718,  19990-19995,  44358,  51991-51993,  51995, 
51996.  All  are  from  Isla  Angel  de  la  Guarda. 
Remarks.  — Ende:rx\\c  to  Isla  Angel  de  la  Guarda. 

Crotalus  mitchellii  [sic]  muertensis  Klauber  (1949) 

Trans.  San  Diego  Soc.  Nat.  Hist.  1 1(6):61-1 16  (p.  97). 

//o/c»0'P^— SDSNH  37447  an  adult  male;  El  Muerto  Island,  Gulf  of  California, 
Mexico.  By  Charles  H.  Lowe,  Jr.,  June  6  or  7,  1946. 

Paratypes.-SDS^H  37442-37444,  37446,  37448,  37449,  38040  El  Muerto  Island. 

Remarks.  — E[  Muerto  Island  =  Isla  Miramar  in  the  San  Luis  group,  Baja  California 
Norte.  The  correct  spelling  is  mitchelli. 

Crotalus  molossus  estebanensis  Klauber  ( 1 949) 

Trans.  San  Diego  Soc.  Nat.  Hist.  1  1(6):61-1  16  (p.  104). 

//o/o/V'/?^.— SDSNH  26792  an  adult  female;  San  Esteban  Island,  Gulf  of  California, 
Mexico.  By  an  expedition  under  Capt.  G.  Allan  Hancock,  April  17.  1937  (preserved). 
Remarks.  — Endemic  to  Isla  San  Esteban. 

Crotalus  ruber  lorenzoensis  Radcliffe  and  Maslin  (1975) 
Copeia  1975(3):490-493  (p.  490). 

//o/o/v/?^— SDSNH  46009  an  adult  male;  San  Lorenzo  Sur  Island,  Gulf  of  Cali- 
fornia, Baja  California  Norte,  Mexico.  By  Charles  E.  Shaw,  May  23,  1966. 


160 


Paratypes. -SUSNH  6605,  45052,  45053,  all  from  San  Lorenzo  Island  Sur. 
Remarks.  — Endemic  to  San  Lorenzo. 

Crotalus  triseriatus  anahuacas  Gloyd  ( 1 940) 

Chicago  Acad.  Sci.  Special  Publ.  No.  4:1-270  (p.  91).  * 

Paratvpes.  —  ^DSHYi  43404  43  km  N  of  Tres  Cumbres,  Morelos,  Mexico. 
Remarks. -HoXoXyipQ:  MCZ  33681  a  female.  SDSNH  43404  was  formerly  T-S 
5492. 

Crotalus  triseriatus  aquilus  Klauber  (1952) 

Bull.  Zool.  Soc.  San  Diego  26:1-143  (p.  24). 

Paratypes.-SUS^H  3496-3501,  6575-6577  vicinity  of  Alvarez,  San  Luis  Potosi, 
Mexico  (Topotypes). 

Remarks.  — =^Crotallus  aquilus  according  to  Harris  and  Simmons  (1978,  Bull. 
Maryland  Herp.  Soc.  14(3):105-21 1).  Holotype:  MCZ  27843  an  adult  female. 

Crotalus  vegrandis  Klauber  (1941) 

Trans.  San  Diego  Soc.  Nat.  Hist.  9(30):333-335  (p.  334). 

Paratypes.SDS^H  34607  Maturin  Savannah,  near  Uracoa  Monagas,  Venezuela. 
Remarks. -HoXoXype:  CM  17384  an  adult  male.  SDSNH  34607  was  formerly  CM 

17385. 

Crotalus  viridis  caliginis  Klauber  (1949) 

Trans.  San  Diego  Soc.  Nat.  Hist.  1 1(6):61-1 16  (p.  90). 

//o/o/.vp^.  —  SDSNH  2800  an  adult  male;  South  Coronado  Island  off  the  northwest 
coast  of  Baja  California,  Mexico.  By  E.  H.  Quayle,  June  2,  1930. 

Paratypes.-SDS^U  2801-2804,  4924-4926,  7538-7540,  1 1 177,  1 1 178,  1371 1- 
13715,  20077,  20078  "All  probably  came  from  South  Coronado  Island,  Baja  California 
Norte,  Mexico." 

Remarks.  — ¥jno^n  only  from  the  type  locality. 

Crotalus  willardi  meridionalis  Klauber  (1949) 

Trans.  San  Diego  Soc.  Nat.  Hist.  1 1(8):  12 1-1 40  (p.  131). 

Holotype.  — SDSNW  6569  an  adult  female;  Coyotes  ["on  the  railroad  to  El  Sato"], 
elevation  8000',  Durango,  Mexico.  By  Edmund  Heller  and  Charles  M.  Barber,  August 
1904. 

Remarks. —  S>r>SHY{  6569  was  one  of  two  specimens  obtained  from  the  Field  Mu- 
seum, both  of  which  were  numbered  FMNH  1493. 

Hypsiglena  nuchalatus  W.  Tanner  (1943) 
Great  Basin  Nat.  4(1  &  2):49-54  (p.  49). 

Paratypes.-ST:>SNH  20233,  20293,  22501  Visalia,  Tulare  Co.,  California. 

Remarks.  —  =Hvpsiglena  torquata  nuchulata  according  to  Bogert  and  Oliver  ( 1 945, 
Bull.  Amer.  Mus.  Nat.  Hist.  83(6):297-426  (p.  381)).  Holotype:  BYU  3008  a  small 
female. 

Hypsiglena  ochrorhynchus  [sic]  klauberi  W.  Tanner  (1944) 
Great  Basin  Nat.  5(3  &  4):25-92  (p.  71). 

//o/ory/7^.  — SDSNH  20228  a  male;  South  Cornoado  [=Coronado]  Island,  Lower 
California' [Baja  California  Norte].  By  Philip  M.  Klauber,  June  11,  1933. 

Remarks.  — =Hypsiglena  torquata  klauberi  according  to  Bogert  and  Oliver  (1945, 
Bull.  Amer.  Mus.  Nat.  Hist.  83(6):297-426).  A  review  of  the  nomenclatural  problem 
surrounding  ochrorhvncha  Cope  vs.  torquata  Giinther  is  given  by  Hardy  and  Mc- 
Diarmid  (1969:169,  Univ.  Kansas  Publ.  Mus.  Nat.  Hist.  18(3):39-252). 

Hypsiglena  ochrorhynchus  [sic]  tortugaensis  W.  Tanner  (1944) 
Great  Basin  Nat.  5(3-4):25-92  (p.  69). 

Paratypes.—^DS^H  4074  Tortuga  Island  [Baja  California  Sur,  Mexico]. 


161 

Remarks.  —  =Hypsiglena  torquata  tortugaensis  according  to  Bogert  and  Oliver  ( 1 945, 
Bull.  Amer.  Mus.  Nat.  Hist.  83(6):297-426).  Holotype:  CAS  51460  a  female  (by  in- 
ference). See  also  Remarks  under  H.  o.  klauberi. 

Hypsiglena  torquata  catalinae  W.  Tanner  (1966) 

Trans.  San  Diego  Soc.  Nat.  Hist.  14(1 5):  189-1 96  (p.  192). 

//(9/o/\'/7£'.  —  SDSNH  44680  an  adult  male;  Santa  Catalina  Island,  approximately 
25°38'N,  ilO°47'W,  Gulf  of  California,  Baja  California  [Sur],  Mexico.  By  George  E. 
Lindsay,  June  25,  1964. 

Paratypes.-SU^^H  44376,  44681  Topotypes. 

Lampropeltus  zonata  pulchra  Zweifel  (1952) 
Copeia  1952(3):  152-1 68  (p.  162). 

Holotype. —  S,r>S>N\\  38667  an  adult  male;  near  Crater  Camp  [450  ft],  Santa  Monica 
Mountains,  Los  Angeles  Co.,  California. 

Remarks.  — Date  and  collector  unknown.  Apparently  received  from  the  San  Diego 
Zoo,  preserved  August  2,  1947. 

Leptotyphlops  humilis  cahuilae  Klauber  (1931) 

Trans.  San  Diego  Soc.  Nat.  Hist.  6(23):333-352  (p.  339). 

//o/o/yp6'.  — SDSNH  2637  an  adult;  Yaqui  Well  by  the  County  Road  Camp,  San 
Diego  Co.,  California.  By  Laurence  M.  Klauber,  May  15,  1930. 
Paratypes.—^onQ  designated. 
Remarks. —  Sex  of  Holotype  not  given. 

Leptotyphlops  humilis  lindsayi  Murphy  (1975) 

Proc.  California  Acad.  Sci.  ser.  4,  40(5):93-107  (p.  96). 

Holotype.— SUSNH  44386  an  adult  female;  Marquer  Bay,  Isla  Carmen,  Baja 
California  (Sur),  Mexico.  By  Charles  E.  Shaw  and  George  E.  Lindsay,  April  4,  1962. 
Remarks.  — Yjivo^jn  only  from  Holotype. 

Lichanura  roseofusca  gracia  Klauber  (1931) 

Trans.  San  Diego  Soc.  Nat.  Hist.  6(20):305-318  (p.  307). 

Holotype. —  SDSNH  2995  a  young  female;  Randsburg,  Kern  Co.,  California.  By 
Lucile  Rector,  June,  1930. 

Remarks.  — =Lichamira  trivirgata  gracia  according  to  Miller  and  Stebbins  (1964: 
189  The  Lives  of  Desert  Animals  in  Joshua  Tree  Nat'l  Monument,  Univ.  Calif.  Press 
vi  +  452  p.)  and  others  since,  all  without  comment.  For  a  discussion  see  Yingling 
(1982,  SSAR  Cat.  Amer.  Amph.  Rept.  294.1). 

Masticophis  bilineatus  lineolatus  Hensley  (1950) 

Trans.  Kansas  Acad.  Sci.  53(2):270-288  (p.  272). 

//o/oryp<?.-SDSNH  43402  an  adult  male;  12.9  mi  S  and  5  mi  E  of  Ajo,  Pima  Co., 
Arizona. 

Remarks. -SDS^H  43402  was  formerly  UIMNH  561 1. 

Masticophis  bilineatus  slevini  Lowe  and  Norris  (1955) 
Herpetological  1 1(2):89-96  (p.  93). 

Holotype. —  SDSNH  3826  an  adult  female;  San  Esteban  Island,  Gulf  of  California, 
Sonora,  Mexico.  By  Mrs.  Griffing  [Margaret]  Bancroft,  April  18.  1930. 

Paratype. —  SDSNH  41571  San  Esteban  Island,  Gulf  of  California,  Sonora,  Mexico. 

Phyllorhynchus  browni  lucidus  Klauber  (1940) 

Trans.  San  Diego  Soc.  Nat.  Hist.  9(20):  1 95-2 14  (p.  202). 

Holotype.  — SDSNH  28819  a  juvenile  female;  Enchanto  Valley,  7  mi  W  of  Cave 
Creek,  Maricopa  Co.,  Arizona.  By  V.  Housholder,  May  21,  1938. 

Phyllorhynchus  decurtatus  nubilus  Klauber  ( 1 940) 

Trans.  San  Diego  Soc.  Nat.  Hist.  9(20):  195-2 14  (p.  197). 


162 


Holotype.  —  SDSNH  32493  an  adult  male;  Xavier  (Weisner's  Ranch),  Pima  Co., 
Arizona.  By  Lee  Arnold,  July  16,  1939. 

Paratvpes.  — From  Arizona  as  follows:  Pima  Co.:  SDSNH  29216  2  mi  N  of  San 
Xavier  Mission;  SDSNH  29287  4  mi  N  of  San  Xavier  Mission;  SDSNH  3^2273  Ajo 
Junction  (Escuela);  SDSNH  32289  4  mi  S  of  Ajo  Junction;  SDSNH  32290  2  mi  S  of 
Ajo  Junction  near  Tucson;  SDSNH  32468  east  base  of ''A"  Mountain;  SDSNH  32494 
5  mi  N  of  Tucson.  Pinal  Co.:  SDSNH  32274  26  mi  N  of  Tucson. 

Remarks.  —  ¥Aauber  noted  in  his  catalogue  that  SDSNH  29287  and  32468  were 
"traded  to  Slevin,  1941";  these  are  now  CAS  specimens. 

Phyllorhynchus  decurtatus  perkinsi  Klauber  (1935) 
Bull.  Zool.  Soc.  San  Diego  12:1-31  (p.  11). 

//o/oO'P^.  — SDSNH  23757  an  adult  male;  [Benson's]  Dry  Lake  3  mi  W  of  Imperial 
Co.  Line  on  Narrows,  Kane  Springs  Road,  San  Diego  Co.,  California.  By  Laurence  M. 
Klauber,  May  4,  1935. 

Paratvpes— ^D^^U  22260,  22261,  22295,  22297,  22298,  22741,  22954,  22955, 
23027,  23386,  23387,  23750-23756, 23758-23761,  23815,  23846-23848, 2391 1-23913, 
23918-23923,  23951,  23995-23997  all  from  very  near  the  type  locality. 

Pituophis  catenifer  bimaris  Klauber  ( 1 946) 

Trans.  San  Diego  Soc.  Nat.  Hist.  1 1(1):  1-40  (p.  7). 

//o/on^/?^'.  —  SDSNH  3262 1  an  adult  male;  Santa  Gertrudis,  near  El  Arco  (lat.  28°N), 
Baja  California,  Mexico.  By  Robert  S.  Hoard,  August,  1939. 

Paratvpes.- A\\  from  Baja  California,  Mexico,  as  follows:  SDSNH  2934,  2935  El 
Refugio  (northeast  of  Magdalena  Bay);  SDSNH  32523  18  mi  N  of  Punta  Prieta  (lat. 
27°N);  SDSNH  3813,  11553  San  Ignacio;  SDSNH  31032  Bahia  Thurtoe  (1  mi  S  of 
Bahia  Tortuga);  SDSNH  17562  and  17563  Rancho  Las  Flores  (12  mi  E  of  El  Arco); 
SDSNH  1129,  1181  El  Marmol. 

Remarks.  — =Pituophis  melanoleucus  bimaris  according  to  Smith  and  Kennedy 
(1951,  Hereptologica  7(3):93-96). 

Pituophis  catenifer  coronalis  Klauber  (1946) 

Trans.  San  Diego  Soc.  Nat.  Hist.  1 1(1):  1-40  (p.  19). 

Holotype.-SUSNW  20229  an  adult  female;  South  Coronado  Island,  Baja  Cali- 
fornia Norte,  Mexico.  By  Philip  M.  Klauber,  June  11,  1933. 

Paratype.  —  SD^^Yi  1 1365  Coronado  Islands,  Baja  California  Norte,  Mexico  (re- 
stricted to  South  Coronado  Island  by  Klauber,  ibid  p.  20). 

Remarks.  — =Pituophis  melanoleucus  coronalis  according  to  Smith  and  Kennedy 
(1951,  Herpetologica  7(3):93-96). 

Pituophis  catenifer  fuliginatus  Klauber  (1946) 

Trans.  San  Diego  Soc.  Nat.  Hist.  11(1):  1-40  (p.  14). 

//o/o/vp£'.  — SDSNH  17449  a  young  adult  female;  San  Martin  Island,  off  the  west 
coast  of  Baja  California,  Mexico.  By  Lewis  W.  Walker,  July  1 1,  1939. 

Paratypes.-^DS^H  17463,  17464  Topotypes. 

Remarks.  — =Pituophis  melanoleucus  fuliginatus  according  to  Smith  and  Kennedy 
(1951,  Herpetologica  7(3):93-96). 

Pituophis  catenifer  pumilus  Klauber  ( 1 946) 

Trans.  Soc.  Diego  Soc.  Nat.  Hist.  ll(2):41-48  (p.  41). 

Holotype.  —  SDS^H  17238  a  young  adult  male;  Santa  Cruz  Island,  Santa  Barbara 
Co.,  California.  By  Norman  Bilderback,  May  5,  1938. 

Remarks.  — =Pituophis  melanoleucus  pumilis  according  to  Smith  and  Kennedy 
(1951,  Herpetologica  7(3):93-96). 

Rhinochelus  lecontei  clarus  Klauber  (1941) 

Trans.  San  Diego  Soc.  Nat.  Hist.  9(29):289-332  (p.  308). 


163 


Holotvpe.  —  SDSNH  3 1440  an  adult  male;  Borrego  Valley,  2  mi  N  of  The  Narrows, 
San  Diego  Co.,  California.  By  Richard  Neil,  May  7,  1939. 

Paratvpes.- As  follows:  San  Diego  County:  SDSNH  2631,  1 1288,  26815,  28650, 
3 1 49 1  Yaqui  Well:  SDSNH  1 1 349  La  Puerta:  SDSNH  1 6998.  268  1 6  San  Felipe  Vallev; 
SDSNH  17009,  25632,  20849,  20850,  28755,  33022  The  Narrows;  SDSNH  3 1445  "5 
mi  W  of  The  Narrows:  SDSNH  23442  Benson's  Dry  Lake;  SDSNH  29078,  32993 
Borrego  Valley:  SDSNH  29110,  29514,  33398  Sentenac  Canyon  (vicinity);  SDSNH 
33021,  33049,  33060  Scissors  Crossing  (vicinity).  Riverside  County:  SDSNH  28670 
Cathedral  City:  SDSNH  31503  Indio;  SDSNH  32137  Palm  Springs  R.R.  Station. 

Remarks.  — =Rhinochelus  lecontei  lecontei  according  to  Shannon  and  Humphrey 
(1963,  Herpetological  19(3):153-160).  See  also  Medica  (1975,  SSAR  Cat.  Amer.  Am- 
phib.  Rept.  175.2). 

Salvadora  grahamiae  virgultea  Bogert  (1935) 

Bull.  So.  California  Acad.  Sci.  34(l):88-94  (p.  89). 

Holotvpe.  — SDSNH  12025  a  young  adult  male;  Deerhorn  Flat,  San  Diego  Co., 
California'.  By  F.  E.  Walker,  June  29,  1929. 

Remarks.  — =Salvadora  hexalepis  virgultea  according  to  Schmidt  (1940,  Zool.  Ser. 
Field  Mus.  Nat.  Hist.  24:143-150). 

Salvadora  hexalepis  klauberi  Bogert  (1945) 
Am.  Mus.  Novit.  1285:1-14  (p.  2). 

Holotvpe.  — SDSNH  20912  an  adult  male;  Cape  San  Lucas,  Baja  California,  Mex- 
ico. By  Fred  Lewis,  August  6,  1933  (preserved). 

Paratvpes.— W\  from  Baja  California  as  follows:  SDSNH  3827  San  Ignacio:  SDSNH 
20466,  20511,  20858  Cape  San  Lucas;  SDSNH  30385  3  mi  S  of  Canipole,  NW  of 
Loreto;  SDSNH  30386  5  mi  N  of  San  Xavier  Mission;  SDSNH  30387  Loreto. 

Salvadora  hexalepis  mojavensis  Bogert  (1945) 
Am.  Mus.  Novit.  1285:1-14  (p.  6). 

Paratvpes.— \s  follows:  Arizona:  Mojave  Co.:  SDSNH  17315  Lucky  Star  Mine, 
Chemehuevis  Mountains;  SDSNH  25385  White  Hills,  28  mi  N  of  Chloride.  Coconino 
Co.:  SDSNH  34439  9  mi  W  of  Cameron.  California:  Inyo  Co.:  SDSNH  25384  Towne's 
Pass,  Panamint  Mountains;  SDSNH  28578  Linnie;  SDSNH  32827  Daylight  Pass, 
Funeral  Mountains;  SDSNH  34100  3  mi  SW  of  Wildrose  Station.  Kern  Co.:  SDSNH 
25864  2  mi  N  of  Grapevine  Station;  SDSNH  26 1 30  20  mi  S  of  Inyokem.  San  Bernardino 
Co.:  SDSNH  4400,  4401,  33446  Mountain  Pass,  Ivanpah  Mountains;  SDSNH  8503 
Twenty-nine  Palms;  SDSNH  10692  Klinefelter;  SDSNH  33951  Hawes;  SDSNH  35896 
7  mi  W  of  Red  Pass,  NE  of  Barstow.  Nevada:  Clark  Co.:  SDSNH  25357  13  mi  W  of 
Indian  Springs;  SDSNH  25386  19  mi  SE  of  Indian  Springs  near  Com  Creek. 

Remarks.  — HoXoXy^Q-.  AMNH  63000  an  adult  male. 

Sonora  bancroftae  Klauber  (1943) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(4):69-70  (p.  69). 

Holotvpe.  — SDSNH  35077  a  female;  2  mi  E  of  San  Jorge,  Lower  California,  Mexico. 
By  Mrs.  Griffing  [Margaret]  Bancroft,  April  10,  1942. 

Remarks.  — "^Sonora  semiannulata  according  to  Frost  (1983,  Trans.  Kansas  Acad. 
Sci.  86(l):31-37). 

Sonora  occipitalis  klauberi  Stickel  (1941) 

Bull.  Chicago  Acad.  Sci.  6(7):  135-140  (p.  138). 

Holotvpe.  — SDSNH  29647  an  adult  male;  Tucson,  Pima  Co.,  Arizona.  By  C.  T. 
Vorhies,  June  3.  1938. 

Paratype.  — SDSNH  171 15  3  mi  SE  of  Picacho,  Pinal  Co.,  Arizona. 

Remarks.  — ^Chionactus  occipitalis  klauberi  according  to  Stickel  (1943,  Proc.  Biol. 
Soc.  Wash.  56:109-128). 


164 


Sonora  palarostris  KJauber  (1937) 

Trans.  San  Diego  Soc.  Nat.  Hist.  8(27):363-366  (p.  363). 

Holotype.  —  S>DSNY\  26771  an  adult  male;  5  mi  S  of  Magdalena,  Sonora,  Mexico. 
By  George  Lindsay,  April,  1937. 

Remarks.  —=Chionactis  palarostris  palarostris  according  to  Klauber  (1951,  Trans. 
San  Diego  Soc.  Nat.  Hist.  1 1(9):141-204). 

Tantilla  eiseni  transmontana  Klauber  (1943) 

Trans.  San  Diego  Soc.  Nat.  Hist.  10(5):71-74  (p.  71). 

Holotype.  —  SiySNW  29273  an  adult  male;  1  mi  E  of  Yaqui  Well,  San  Diego  Co., 
California.  By  Charles  Shaw  and  Cyrus  Perkins,  June  6,  1938. 

Paratvpes.  —  ¥vom  California  as  follows:  Riverside  Co.:  SDSNH  33760,  Palm 
Springs.  San  Diego  Co.:  SDSNH  2633,  2634  Yaqui  Well;  SDSNH  11260  La  Puerta; 
SDSNH  32419,  33997  Sentenac  Canyon. 

Remarks.  — =  Tantilla  planiceps  transmontana  according  to  Tanner  (1966,  Her- 
petologica  22(2):  134-1 52)  and  Tantilla  planiceps  according  to  Cole  and  Hardy  (1981: 
268,  Bull.  Amer.  Mus.  Nat.  Hist.  1 7 1  (3):  1 99-284),  who  regard  the  species  as  monotypic. 

Trimorphodon  biscutatus  semimtus  Smith  (1943) 

Proc.  U.S.  Natl.  Mus.  93(3169):393-504  (p.  492). 

Paratype. —  SDSNH  43401  10  mi  N  of  Tafetan,  Michoacan,  Mexico. 

Remarks.  — =  Trimorphodon  biscutatus  biscutatus  according  to  Duellman  (1954, 
Occ.  Pap.  Mus.  Zool.  Univ.  Michigan  560:1-24).  Holotype:  USNM  110410  a  male 
(cited  as  EHT-HMS  23619  in  Smith  1943). 

Acknowledgments 

We  are  very  grateful  to  James  R.  Dixon,  Thomas  H.  Fritts,  Darrel  Frost,  and  Jay 
M.  Savage  for  their  reviews  of  earlier  drafts.  Marjorie  Rea  expertly  typed  the  manuscript. 


Caudata 

Plethodon  richmondi  Netting  and  Mittleman  (1938) 
Ann.  Carnegie  Mus.  27:287-293  (p.  288). 

Paratype.—SDSNW  32680  Oglebay  Park  near  Wheeling,  Ohio  Co.,  West  Virginia. 
Remarks. -Ho\oXypQ\  CM  14189  an  adult  male.  SDSNH  32680  was  formerly  CM 
5296. 


TRANSACTIONS 
^p^ZOO^  OF  THE  SAN  DIEGO 

MOS.  ^pp^^f?y  SOCIETY  OF 


50^ 


^i^ 


NATURAL  HISTORY 


#^ 


Volume  20  Number  11  pp.  165-168     20  June  1984 


Imocaris  tuberculata,  n.  gen.,  n.  sp.  (Crustacea:  Decapoda) 
from  the  upper  Mississippian  Imo  Formation,  Arkansas 

Frederick  R.  Schram 

Department  of  Geology,  San  Diego  Natural  History  Museum,  San  Diego,  CA  92112  USA 

Royal  H.  Mapes 

Department  of  Geology,  Ohio  University,  Athens,  OH  45701  USA 

Abstract.  A  new  genus  and  species  of  decapod  crustacean  is  described  from  the  Upper  Missis- 
sippian Imo  Formation,  near  Leslie,  Arkansas.  The  exceptionally  well-preserved  specimen  is  a  single 
carapace,  associated  with  a  moUuscan  dominated  fauna  including  ammonoids,  gastropods,  and  bivalves. 
Imocaris  tuberculata  serves  to  help  fill  the  stratigraphic  gap  between  the  oldest  decapod,  Palaeopalaemon 
newberryi  in  the  Upper  Devonian  of  Ohio  and  Iowa,  and  the  diverse  Triassic  decapod  faunas  of  Europe. 


Introduction 

Schram  et  al.  (1978)  redescribed  Palaeopalaemon  newberryi  Whitfield,  1880,  and 
recognized  it  as  the  earliest  decapod  crustacean.  However,  until  now,  there  has  been  a 
significant  gap  in  the  decapod  fossil  record  from  this  Upper  Devonian  form  to  the 
better  known  Triassic  decapods  of  central  Europe  (see  e.g.,  Forster,  1 967).  The  specimen 
described  herein  is  a  single  well-preserved  carapace  from  the  Upper  Mississipian- 
Chesterian  (Naurian-Amsbergian;  ammonoid  zone  E2b-c)  of  Arkansas  interpreted  to 
be  a  decapod  crustacean. 

This  specimen  was  collected  from  a  road  cut  along  Peyton  Creek  just  south  of  the 
Van  Buren  Co.  hne,  Arkansas,  along  U.S.  Hwy.  65  approximately  6.4  km  (4  mi) 
southeast  of  Leslie,  Arkansas.  The  fossil  was  found  as  the  result  of  washing  and  con- 
centrating some  817  kgs  of  matrix  from  a  0.6  m  (2  ft)  concretionary  shale  layer  ap- 
proximately 32  m  (105  ft)  above  the  contact  of  the  Imo  Formation  with  the  underlying 
Pitkin  Formation  (in  the  middle  of  "bed  21,"  of  Sutherland  and  Manger  1977,  fig.  1). 


Systematic  paleontology 

Order  Decapoda  Latreille,  1803 

Suborder  Pleocyemata  Burkenroad,  1963 

Infraorder  Reptantia  Boas,  1 800 

Section  Brachyura  Latreille,  1803 

Subsection  Dromiacea  de  Haan,  1833 

Genus  Imocaris  n.  gen. 

Z)/a^«05/5.— Carapace  cylindrical,  with  subrectangular  outline  in  both  dorsal  and 
lateral  views.  Surface  tuberculate.  Cervical  and  branchiocardiac  grooves  prominent. 
Rostrum  small  to  non-existent.  Antero-lateral  margin  with  prominent  denticles. 

Etymology.— ^3.mQd.  after  the  Imo  Formation,  gender  feminine. 

Type  species.  — Imocaris  tuberculata  n.  sp. 


166 


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Figure  1.     Imocaris  tuberculata  n.  sp.,  holotype  SDSNH  25139,  stereo  pair,  scale  =  5  mm. 


Imocaris  tuberculata  n.  sp. 
Figure  1 

Diagnosis.  — ^QcaM^t  there  is  but  one  species,  the  diagnosis  is  the  same  as  that  of 
the  genus. 

//o/orw?^.- SDSNH  25139  (Fig.  1). 

Loca//7y.- SDSNH  loc.  3191;  NE  Va,  sec.  11,  T.13N.,  R.15W.;  along  road  cut  in 
U.S.  Hwy.  65,  south  of  Van  Buren  Co.  line,  Arkansas,  on  Peyton  Creek. 

Stratum.  — Imo  Formation,  Chesterian,  Upper  Mississippian. 

Etymology.— After  the  tuberculate  nature  of  the  carapace. 

Z)^^^/?//^^.— Anterior  margin  of  carapace  straight,  without  orbits,  with  only  slight 
suggestion  of  rostrum.  Lateral  margin  anterior  of  branchiocardiac  groove  with  at  least 
5  prominent  denticles,  and  posterior  of  branchiocardiac  groove  marked  by  submarginal 
furrow.  Posterior  margins  slightly  concave,  with  faint  submarginal  furrow.  Prominent, 
deep  cervical  and  branchiocardiac  grooves,  cervical  grooves  extend  laterally  from  mid- 
line and  turn  anteriad  as  short  antennar  grooves,  branchiocardiac  grooves  continuous 
laterally  with  prominent  inferior  grooves.  Carapace  surface  tuberculate.  Mid-dorsal  line 
marked  by  row  of  4  large  tubercles  between  rostral  area  and  cervical  groove,  by  slight 
groove  between  the  cervical  and  branchiocardiac  grooves,  and  by  clear  ridge  sur- 
mounted by  row  of  5  (?6)  tubercles  between  branchiocardiac  groove  and  posterior 
margin.  Pair  of  tubercle  rows  located  laterally  between  branchiocardiac  groove  and 
posterior  margin  about  '/2-way  between  median  ridge  and  lateral  margins,  forming 
planes  at  which  curvature  of  posterior  carapace  surface  changes  from  horizontal  to 
more  vertical  orientation.  Carapace  mid-line  length—  1 1  mm,  approximate  lateral  mar- 
gin length— 12  mm,  width  — 9.5  mm. 

Remarks.  — The  fossil  preserves  an  excellent  view  of  the  dorsal  surface  of  the 
carapace.  Attempts  at  preparation  of  the  underside  and  margins  of  the  specimen, 
however,  seem  to  indicate  that  only  the  carapace  is  fossilized,  i.e.,  no  ventral  thoracic 
structures  appear  to  be  present. 


167 


Discussion 

The  higher  taxonomic  affinity  of  any  Paleozoic  decapod  is  of  interest  because  of 
the  understanding  it  might  lend  to  elucidating  the  sequence  of  events  in  the  early 
radiation  of  Decapoda.  Paleozoic  decapods  have  been  rare  and  widely  scattered  geo- 
graphically. The  type  specimens  of  the  supposed  Permian  decapod  carapaces  from 
Sicily,  Palaeopemphix  Gemmellaro,  1890,  have  been  lost  and,  thus,  their  suggested 
decapod  versus  cumacean  affinities  can  never  be  verified.  The  Devonian  Palaeopalae- 
mon  newberryi,  preserved  in  great  detail,  possesses  several  features  in  a  combination 
of  both  astacidean  and  palinuran  reptants  (Schram  et  al.  1978);  and,  in  addition, 
Felgenhauer  and  Abele  (1983)  have  noted  a  characteristic  natatian  feature  in  that  form 
as  well,  viz.,  the  large  scaphocerite.  Bachmeyer  and  Malzahn  (1983)  have  recorded 
from  the  Upper  Permian  Zechstein  of  Germany  a  free  decapod  cheliped  (given  the 
name  Erymastacus  ?  hoerstgenensis),  as  well  as  leg  and  carapace  fragments  attributed 
to  decapods  in  the  stomach  contents  of  the  chimaeriform  fish  Janassa  hituminosa.  The 
fragmentary  nature  of  the  Zechstein  fossils,  however,  makes  them  difficult  to  compare 
to  the  other  known  Paleozoic  decapod  whole  body  fossils.  Finally,  little  can  be  said 
about  the  Soviet  Permian  form  Protoclytiopsis  antiqua  Birshtein,  1958,  except  that  it 
appears  to  be  taxonomically  close  to  a  large,  reptant,  glypheoid  type  (Schram  1980). 

Several  characters  mark  Imocahs  tuberculata  as  unusual,  viz.,  the  cylindrical  form 
and  rectangular  proportions  of  the  carapace,  the  presence  of  only  a  branchiocardiac 
groove  posterior  to  the  cervical  (or  alternatively  a  fusion  of  the  post-cervical  and 
branchiocardiac  grooves),  the  lack  of  orbits,  and  the  small  or  absent  rostrum.  Unfor- 
tunately, the  lack  of  preservation  of  any  eyes,  antennae,  mouthparts,  thoracopods,  and 
abdomen  makes  a  definitive  higher  taxonomic  assignment  difficult. 

Quite  unexpectedly,  the  closest  analogs  to  the  derived  features  of  Imocaris  tuber- 
culata listed  above  are  to  be  found  among  the  dromiacean  brachyurans.  These  living 
forms  tend  to  possess  markedly  cylindrical  and  somewhat  subrectangular  carapaces, 
strong  cervical  and  branchiocardiac  grooves,  absent  or  poorly  developed  orbits,  fre- 
quently weakly  developed  rostra,  and  (in  at  least  the  prosopids,  some  dynomenids,  and 
the  homoloids)  prominent  carapace  sculpturing.  The  assignment  of  Imocahs  to  the 
dromiaceans  on  the  basis  of  this  single  specimen  considerably  extends  the  range  of  that 
taxon  back  from  the  Lower  Jurassic  into  the  middle  of  the  Carboniferous. 

The  recognition  of  Imocaris  as  a  decapod  begins  to  fulfill  the  expectations  of  Schram 
et  al.  (1978)  that  an  extensive  Paleozoic  radiation  of  Decapoda  is  to  be  discovered  in 
upper  Paleozoic  strata. 

ACICNOWLEDGMENTS 

This  research  is  supported  in  part  by  NSF  grant  BSR  82-12335  (FRS)  and  a  grant 
from  the  Ohio  University  Research  Council  (RHM). 


Literature  Cited 


Bachmeyer,  F.,  and  E.  Malzahn.  1983.  Der  erste 
Nachweis  eines  decapoden  Krebses  im  nie- 
derrheinischen  Kupferschiefer.  Annalen  des 
Naturhistorischen  Museums  in  Wien.  85/A:99- 
106. 

Birshtein,  Ya.  A.  1958.  Drevneishii  predsta  vitel 
otryada  desyatinogikh  rakoobraznikh,  Proto- 
clytiopsis antiqua,  in  Permakikh  otiozhenii  za- 
padnoi  Sibiri.  Doklady  Akademii  Nauk  SSSR 
122:477-480. 

Felgenhauer,  B.  E.,  and  L.  G.  Abele.  1983.  Phy- 
logenetic  relationships  among  shrimp-like 
decapods.  Crustacean  Issues  1:291-31 1. 

Forster,  R.    1967.    Die  reptanten  Dekapoden  der 


Trias.  Neues  Jahrbuch  fur  Geologic  und  Pa- 
leontologie  Abhandhungen  128:136-194. 

Gemmellaro,  G.  G.  1890.  Crostacei  dei  calcari 
con  Fusulina  della  Valle  del  Fiume  Sosio,  nella 
provicia  di  Palermo  in  Sicilia.  Societa  Italiana 
di  Scienze  Naturali,  Mcmorie  (3)8:1-40. 

Schram,  F.  R.  1980.  Notes  on  miscellaneous  crus- 
taceans from  the  Late  Paleozoic  of  the  Soviet 
Union.  Journal  of  Paleontology  54:542-547. 

,  R.  M.  Feldmann,  and  M.  J.  Copcland.  1978. 

The  Late  Devonian  Palaeopalaemonidae  and 
the  earliest  decapod  crustaceans.  Journal  of  Pa- 
leontology 52:1375-1387. 

Sutherland,  P.  K.,  and  W.  L.  Manger.  1977.  Up- 
per Chesterian-Morrowan  Stratigraphy  and  the 


168 


Mississippian-Pennsylvanian  Boundary  in  Whitfield,  R.  P.    1880.    Notice  of  new  forms  o. 

Northeastern    Oklahoma   and    Northwestern  fossil  crustaceans  from  the  Upper  Devonian 

Arkansas.    Oklahoma   Geological    Survey  rocks  of  Ohio,  with  description  of  new  genera 

Guidebook  1 8.  University  of  Oklahoma,  Nor-  and  species.  American  Journal  of  Science  (3)19: 

man.  33-42.                                         » 


(^^"^     j^  TRANSACTIONS 

%  OF  THE  SAN  DIEGO 

^  SOCIETY  OF 

^   ^^J^^  NATURAL  HISTORY 

Volume'^  Number  12  pp.  169-188       20  November  1984 


New  material  of  Hydrodamalis  cuestae  (Mammalia:  Dugongidae)  from 
the  Miocene  and  Pliocene  of  San  Diego  County,  California 


Daryl  P.  Domning 

Department  of  Anatomy,  Howard  University.  Washington,  D.C.  20059 

Thomas  A.  Demere 

Department  of  Geology,  Natural  History  Museum,  P.O.  Box  J  390,  San  Diego,  CA  92112 


Abstract.  The  geology,  faunal  content,  and  age  of  sirenian-bearing  marine  rocks  in  San  Diego 
County,  southern  California,  are  reviewed  and  reevaluated.  The  San  Mateo  Formation  comprises  two 
members,  respectively  of  Late  Miocene  (late  Clarendonian  or,  more  likely,  early  Hemphillian)  and 
Early  Pliocene  (late  Hemphillian)  age.  The  San  Diego  Formation  also  includes  two  members,  both 
probably  of  Late  Pliocene  (Blancan)  age,  though  the  upper  member  may  extend  into  the  Early  Pleis- 
tocene. All  four  members  have  yielded  new  specimens  of  sirenians,  most  or  all  of  which  represent 
Hydrodamalis  cuestae  Domning  1978.  These  specimens  extend  our  knowledge  of  the  osteology  of  this 
species,  and  confirm  the  supposition  that  the  holotype  is  abnormal  in  several  respects.  During  the 
Hemphillian,  the  juveniles  of  the  species  had  an  upper  dentition  probably  consisting  of  DP^-^;  it  is  still 
unknown  whether  later  Pliocene  juveniles  retained  teeth.  A  braincase  from  the  San  Diego  Formation 
is  the  largest  known  of  any  sirenian,  probably  representing  an  individual  over  10  m  long  and  confirming 
the  previous  observation  that  Hydrodamalis  grew  larger  in  California  than  at  Bering  Island. 


Introduction 

Domning  ( 1 978)  reviewed  the  fossil  record  of  North  Pacific  sirenians  and  described 
the  new  species  Hydrodamalis  cuestae,  a  form  intermediate  between  the  Late  Miocene 
Dusisiren  jordani  (Kellogg  1925)  and  the  Late  Pleistocene-Recent  H.  gigas  (Zimmer- 
mann  1780)  (Steller's  sea  cow).  The  holotype  of  H.  cuestae  came  from  the  Upper 
Pliocene  (Blancan)  Pismo  Formation  in  San  Luis  Obispo  County,  California,  and  re- 
ferred specimens  came  from  the  Hemphillian  San  Mateo  Formation  in  San  Diego 
County,  the  Hemphillian  Capistrano  Formation  in  Orange  County,  and  an  unnamed 
Pliocene  unit  in  Baja  California,  Mexico.  A  single  thoracic  vertebra  from  the  Blancan 
San  Diego  Formation  near  Tijuana,  Mexico,  was  identified  only  as  Hydrodamalis 
species. 

Since  1978,  a  number  of  additional  specimens  o^  Hydrodamalis  cuestae  have  been 
obtained  by  the  San  Diego  Natural  History  Museum  from  the  San  Mateo  and  San 
Diego  Formations  in  San  Diego  County.  These  clarify  some  of  the  problems  posed  by 
the  previously  known  material;  in  particular,  they  show  that  certain  anomalous  features 
of  the  holotype,  thought  by  Domning  (1978)  to  be  pathological,  are  indeed  atypical  of 
the  species. 

Abbreviations  used  are  as  follows:  SDSNH,  San  Diego  Society  of  Natural  History; 
UCMP,  University  of  California  Museum  of  Paleontology,  Berkeley;  USNM,  U.S. 
National  Museum  of  Natural  History,  Washington,  D.C. 


170 


Geology  and  Age  of  the  Deposits 

Remains  of  fossil  hydrodamalines  from  San  Diego  County,  California,  have  been 
recovered  from  both  the  Upper  Miocene-Lower  Pliocene  San  Mateo  Formation  (Hemp- 
hillian)  at  Oceanside  and  the  Upper  Pliocene  San  Diego  Formation  (Blanc^n)  at  San 
Diego.  These  rock  units  accumulated  in  separate  sedimentary  basins  and  contain  dis- 
tinctly different  marine  vertebrate  assemblages. 

San  Mateo  Formation 

Barnes  (1976)  and  Barnes  et  al.  (1981)  have  tentatively  assigned  the  vertebrate- 
producing  marine  beds  at  Oceanside  to  the  San  Mateo  Formation  of  Woodford  (1925). 
This  rather  poorly  defined  rock  unit  has  been  mapped  from  the  type  area  near  San 
Clemente  throughout  the  coastal  portion  of  Camp  Pendleton  Marine  Corps  Base  as  far 
south  as  the  city  of  Oceanside  (Moyle  1973,  Young  and  Berry  1981).  The  best  exposures 
of  the  San  Mateo  Formation  at  Oceanside  are  in  the  Lawrence  Canyon  area  adjacent 
to  the  San  Luis  Rey  River  (Fig.  1 ).  Here  erosion,  grading,  and  quarry  operations  have 
combined  to  produce  a  number  of  natural  and  artificial  outcrops. 

Lawrence  Canyon  is  aligned  in  part  along  a  north-south  striking,  eastward-dipping, 
high  angle  normal  fault.  The  San  Mateo  Formation  is  confined  to  the  east  side  of  this 
fault  where  it  rests  unconformably  on  westward-dipping  strata  of  the  Middle  Miocene 
San  Onofre  Breccia  and  in  turn  is  overlain  unconformably  by  flat-lying  Upper  Pleis- 
tocene, nonmarine  terrace  deposits.  In  Lawrence  Canyon  the  San  Mateo  Formation  is 
approximately  24  m  thick  and,  as  noted  by  Barnes  et  al.  (198 1),  is  divisible  into  a  lower 
and  an  upper  unit.  The  lower  unit  consists  of  white,  fine-grained,  massive,  friable, 
micaceous  sandstones  with  occasional  green  or  black  claystone  lenses  and  locally  com- 
mon pebbles  and  cobbles.  The  upper  unit  is  a  complexly  bedded  sequence  of  gravels, 
pebble  to  cobble  conglomerates  and  friable  sandstones.  A  sharp  unconformity  marked 
in  places  by  scour  and  fill  features  separates  the  upper  gravel  conglomerate  unit  from 
the  lower  white  sandstones.  Marine  vertebrate  fossils  have  been  collected  from  both 
the  upper  and  the  lower  unit  (Barnes  1976,  Domning  1978,  Barnes  et  al.  1981,  Howard 
1982)  whereas  invertebrate  fossils  are  conspicuously  absent.  Barnes  et  al.  (1981)  have 
assigned  the  vertebrate  fossil  assemblages  from  the  lower  and  upper  units  to  the  San 
Luis  Rey  River  Local  Fauna  and  the  Lawrence  Canyon  Local  Fauna,  resnectively. 

Fossils  in  the  lower  white  sandstones  generally  occur  as  single  isolated  elements 
and  occasionally  as  associated  partial  skeletons.  In  contrast,  fossils  from  the  upper 
gravel  conglomerate  unit  are  often  concentrated  in  distinct  "bone  horizons."  One  par- 
ticular horizon  (SDSNH  locality  3161)  is  a  1.2  m  thick,  fining-upward  sequence 
divisible  into  four  lithologic  units:  1)  a  basal  clast-supported  pebble  to  cobble  con- 
glomerate; 2)  a  matrix-supported  pebble  conglomerate;  3)  an  interval  of  poorly  bedded 
sandstones  and  gravels  with  laminated  fine-grained  sandstone  lenses;  and  4)  thickly 
laminated  fine-  and  coarse-grained  sandstones.  Fossils  are  concentrated  in  the  matrix- 
supported  pebble  conglomerate  and  generally  occur  as  broken  and  abraded  fragments 
of  large  bones  or  as  complete  and  differentially  preserved  smaller  resistant  elements. 
Particularly  common  are  shark  teeth,  cetacean  earbones,  and  bird  humeri  and  ulnae. 
Larger  bones  generally  occur  in  association  with  the  larger  pebbles  or  cobbles,  suggesting 
size-sorting  by  currents.  Apparently  the  fossils  were  transported  and  deposited  as  bio- 
genic clasts  within  the  traction  load. 

In  terms  of  general  depositional  environments  it  appears  that  the  lower  white 
sandstone  unit  was  deposited  under  "normal"  marine  conditions,  perhaps  at  middle 
to  inner  shelf  depths.  The  dominance  of  sandstones  over  finer-grained  lithologies  would 
support  this  idea  of  shallow  water  deposition,  although  without  a  preserved  benthonic 
assemblage  (e.g.,  mollusks)  it  is  not  possible  to  make  any  definite  paleobathymetry 
estimates.  Clearly,  however,  because  of  the  rich  marine  vertebrate  assemblage,  this  is 
a  marine  unit. 

The  upper  gravel  conglomerate  unit,  in  contrast  to  the  lower  white  sandstones,  is 
characterized  by  complex  bedding  and  very  coarse-grained  lithologies.  The  bedding 


171 


33    00 


5    m  I 


10    km 


—   32   30 


I    1  1   7      00 


Figure  1.  Index  map  of  western  San  Diego  County  showing  the  principal  outcrop  areas  of  rocks  containing 
Hydrodamalis  cuestae.  1,  Hemphillian  San  Mateo  Formation,  Lawrence  Canyon;  2,  Blancan  San  Diego 
Formation,  northern  San  Diego  Mesa;  3,  Blancan  San  Diego  Formation,  Chula  Vista/National  City  area. 


features  are  interpreted  as  representing  anastomosing  channel-fill  deposits  and,  together 
with  the  coarse-grained  lithologies,  suggest  a  fluvial  environment.  However,  the  oc- 
currence of  common  marine  vertebrate  fossils  (not  reworked  from  the  lower  unit)  points 
instead  to  a  marine  environment.  The  model  herein  proposed  to  accommodate  these 


172 

apparently  conflicting  environmental  settings  involves  deposition  of  the  upper  gravel 
conglomerate  unit  at  the  distal,  submarine  margin  of  a  river-dominated  gravel-cobble 
delta. 

Fossil  remains  of  Hydwdamalis  cuestae  occur  in  both  the  San  Luis  Rey  River 
Local  Fauna  (lower  white  sandstone  unit)  and  the  Lawrence  Canyon  Local  Fauna  (upper 
gravel  conglomerate  unit).  Both  local  faunas  contain  a  variety  of  taxa  including  sharks, 
rays,  bony  fishes,  sea  birds,  fur  seals,  walrus,  and  toothed  and  baleen  whales  (Barnes 
et  al.  1981).  Recent  work  by  personnel  at  the  San  Diego  Natural  History  Museum  has 
added  additional  taxa,  especially  species  of  birds  (R.  M.  Chandler,  personal  commu- 
nication), to  the  published  faunal  lists  of  Barnes  et  al.  (1981)  and  Howard  (1982).  To 
the  San  Luis  Rey  River  Loca  Fauna  can  be  added  the  white  shark  Carcharodon  me- 
galodon  Agassiz,  a  new  large  species  of  the  flightless  auk  Praemancalla,  a  shearwater 
(Puffinus  species),  an  eagle  (Accipitriformes),  a  mysticete  (Balaenopteridae),  and  the 
camel  Aepycamelus  species  (S.  D.  Webb,  personal  communication).  To  the  Lawrence 
Canyon  Local  Fauna  can  be  added  a  billfish  (Istiophoridae),  a  cormorant  (Phalacrocorax 
species),  a  shearwater  {Puffinus  species),  the  loon  Gavia  concinna  Wetmore,  an  albatross 
(Diomedea  species),  a  mysticete  (Balaenopteridae),  a  llama  (cf  Hemiauchenia  species; 
S.  D.  Webb,  personal  communication),  and  a  peccary  (Tayassuidae).  These  last  two 
taxa  and  the  camel  {Aepycamelus  species)  are  part  of  a  small  terrestrial  mammal  as- 
semblage from  the  San  Mateo  Formation  which  also  includes  horses  of  the  genus 
Dinohippus  (or  possibly  Pliohippus;  B.  J.  MacFadden,  personal  communication). 

Barnes  et  al.  (1981)  have  used  the  joint  occurrence  of  the  horse  (Pliohippus  or 
Dinohippus)  and  the  sabertoothed  salmon  Smilodonichthyes  rastrosus  Cavender  and 
Miller  to  correlate  both  the  upper  and  lower  unit  of  the  San  Mateo  Formation  at 
Oceanside  with  the  Hemphillian  North  American  Land  Mammal  Age.  The  recent 
recovery  of  an  isolated  M-  belonging  to  the  camelid  Aepycamelus  species  from  the 
lower  white  sandstone  unit  offers  a  refinement  of  this  correlation  and  suggests  that  the 
San  Luis  Rey  River  Local  Fauna  is  of  late  Clarendonian  or  early  Hemphillian  age 
(Webbet  al.  1981). 

As  suggested  by  Howard  (1982),  the  avifauna  of  the  San  Mateo  Formation  (es- 
pecially the  mancalline  taxa)  may  offer  a  means  for  finer  biostratigraphic  resolution  in 
the  section.  The  occurrence  of  the  genus  Praemancalla  in  the  lower  unit  and  the  genus 
Mancalla  in  the  upper  unit  is  the  basis  for  this  biostratigraphy.  Previously,  the  genus 
Praemancalla  was  known  only  from  Clarendonian-aged  rocks  (Monterey  Formation) 
in  Orange  County  (Howard  1976),  with  two  species  assigned  to  this  genus.  The  older 
species  P.  lagunensis  Howard  is  recorded  from  the  lower  part  of  the  Monterey  For- 
mation in  Orange  County,  while  P.  wetmorei  Howard  is  recorded  from  the  upper  part 
of  the  formation.  Howard  (1982)  referred  material  from  the  lower  white  sandstone  unit 
at  Oceanside  to  P.  species  cf  P.  wetmorei,  noting  that  this  was  the  first  Hemphillian 
record  of  Praemancalla.  In  the  upper  gravel  conglomerate  unit  Praemancalla  is  replaced 
by  the  genus  Mancalla,  represented  here  by  three  species,  M.  cedrosensis  Howard,  M. 
milleri  Howard,  and  M.  diegensis  (Miller).  The  co-occurrence  of  these  three  species 
appears  to  define  a  useful  assemblage  zone.  Howard  (1982:13)  has  suggested  that  a 
considerable  hiatus  is  represented  by  the  unconformity  within  the  San  Mateo  For- 
mation, one  that  allowed  ".  .  .  time  for  the  demise  of  the  Praemancalla  and  the  evolution 
of  Mancalla.'^  It  seems  then,  that  the  range  of  Praemancalla  defines  a  biostratigraphic 
interval  and  that  the  hiatus  in  the  San  Mateo  Formation  may  be  represented  elsewhere 
by  an  additional  and  younger  biozone.  From  discussions  with  R.  M.  Chandler  {personal 
communication)  it  seems  that  such  a  biozone  is  preserved  in  mudstones  of  the  Cap- 
istrano  Formation  at  San  Clemente.  Here  an  avifauna  was  recovered  which  contains 
Mancalla  californiensis  Lucas,  M.  cedrosensis,  and  the  auklet  Cerorhinca  minor  How- 
ard. (Unfortunately  it  has  not  been  possible  to  place  the  stratigraphic  position  of  this 
fossil  locality  within  a  composite  section  for  the  very  thick  and  long-ranging  Capistrano 
Formation.)  Apparently  this  assemblage  zone  is  also  preserved  within  the  basal  portion 
of  the  Almejas  Formation  on  Cedros  Island,  Baja  California,  Mexico,  where  Howard 
(1971)  recorded  M.  cedrosensis  and  C  m/nor  (although  without  M.  californiensis). 


173 


To  summarize  these  assemblage  zones,  we  then  have:  Praemancalla  lagunensis  in 
the  Monterey  Formation  (Clarendonian);  P.  wetmorei  in  the  Monterey  Formation 
(Clarendonian)  and  "lower"  San  Mateo  Formation  (late  Clarendonian  or  early  Hemp- 
hillian);  Mancalla  cedrosensis  and  Cerorhinca  minor  with  or  without  M.  californiensis 
in  the  Upper  Miocene  portions  of  the  Almejas  Formation  (Hemphillian)  and  Capistrano 
Formation  (Hemphillian);  M.  cedrosensis,  M.  milleri,  and  M.  diegensis  in  the  "upper" 
San  Mateo  Formation  (late  Hemphillian);  and  to  complete  the  zonation.  M.  milleri, 
M.  diegensis,  and  M.  emlongi  Olson  in  the  San  Diego  Formation  (Blancan).  (These 
correlations  and  age  assignments  are  based  in  part  on  the  work  of  Barnes  1976,  Re- 
penning  and  Tedford  1977,  Howard  1978,  1982,  and  R.  M.  Chandler,  personal  com- 
munication.) Whether  the  "lower"  San  Mateo  Formation  is  late  Clarendonian  or  early 
Hemphillian  in  age  cannot  at  this  time  be  resolved  biostratigraphically.  However, 
physical  stratigraphy  tends  to  support  an  early  Hemphillian  age  assignment.  Vedder 
(1972)  has  suggested  that  the  San  Mateo  Formation  (at  least  in  the  type  area)  may  be 
a  channel  facies  within  the  lower  part  of  the  Capistrano  Formation.  Similarly,  Ehlig 
(1979)  has  mapped  the  San  Mateo  Formation  near  San  Onofre  as  a  member  of  the 
Capistrano  Formation.  Although  the  nomenclatural  standing  of  the  San  Mateo  For- 
mation is  questionable  (it  is  retained  here  pending  completion  of  ongoing  field  work 
at  Camp  Pendleton),  its  correlation  with  the  Hemphillian  Capistrano  Formation  (and 
not  the  Clarendonian-aged  Monterey  Formation)  appears  to  be  certain.  Thus  the  lower 
white  sandstones  at  Oceanside  probably  are  correlative  with  the  basal  portion  of  the 
Capistrano  Formation  (Upper  Miocene,  early  Hemphillian),  while  the  upper  gravel 
conglomerate  unit  at  Oceanside  probably  correlates  with  some  horizon  near  the  top  of 
the  Capistrano  Formation  (Lower  Pliocene,  late  Hemphillian).  Howard  (1982)  has 
proposed  a  similar  Late  Miocene-Early  Pliocene  correlation  for  the  San  Mateo  For- 
mation at  Oceanside,  in  contrast  to  the  strictly  Miocene  age  assignment  of  Barnes  et 
al.  (1981). 

San  Diego  Formation 

The  geology  of  the  marine  Pliocene  San  Diego  Formation  has  recently  been  sum- 
marized by  Demere  ( 1 983).  This  rock  unit  extends  in  a  broad  area  of  outcrop  throughout 
much  of  the  southwestern  portion  of  San  Diego  County  (Fig.  1).  The  formation  is 
fossiliferous  throughout  this  area  of  outcrop  and  contains  a  diverse  and  well-preserved 
assemblage  of  both  marine  invertebrate  and  vertebrate  taxa. 

The  San  Diego  Formation  was  deposited  during  a  marine  transgression  of  the 
Neogene  San  Diego  Basin,  which  like  other  onshore  sedimentary  basins  in  southern 
California  (e.g.,  Ventura  Basin,  Los  Angeles  Basin)  is  structurally  related  to  the  wrench 
and  extensional  tectonics  of  the  continental  borderland.  Deposition  began  during  the 
Late  Pliocene  and  possibly  continued  into  Early  Pleistocene  time,  accumulating  at  least 
75  m  of  marine  and  9  m  of  nonmarine  sedimentary  rocks.  The  overall  stratigraphic 
sequence  suggests  a  successive  filling  and  shallowing  of  this  basin.  It  is  now  apparent 
that  extensional  tectonics  have  controlled  both  the  initial  deposition  as  well  as  the 
present  outcrop  distribution  of  this  rock  unit.  Numerous  high-angle  normal  faults 
striking  north  to  northwest  cut  the  area  into  a  series  of  fault  blocks  which  expose 
different  portions  of  the  Pliocene  section. 

In  an  attempt  to  correlate  these  various  fault  blocks.  Demere  ( 1 983)  has  informally 
subdivided  the  San  Diego  Formation  into  a  "lower"  and  an  "upper"  member  using 
both  lithologic  and  paleontologic  criteria.  The  "lower"  member  is  characterized  by 
yellowish,  very  fine-grained,  massive,  friable,  micaceous  sandstones  with  locally  well- 
bedded  sequences  of  laminated  and  cross-bedded  sandstones,  pebble  to  cobble  con- 
glomerates and  well-cemented  shell  beds.  This  "lower"  member  is  richly  fossiliferous 
and  has  produced  the  bulk  of  the  marine  invertebrate  fauna  so  well  known  through 
the  work  of  Grant  and  Gale  (1931)  and  Hertlein  and  Grant  (1944,  1960,  1972).  In 
addition,  the  diverse  avifaunas  (Howard  1949,  Miller  1956)  and  cetacean  assemblages 


174 


(Barnes  1973,  1976)  reported  from  the  San  Diego  Formation  have  been  largely  collected 
from  the  "lower"  member. 

Lithologically  the  "upper"  member  is  characterized  by  well-bedded  sequences  of 
pebble  to  cobble  conglomerate,  well-cemented  fossiliferous  sandstones,  and^medium- 
to  coarse-grained  friable  sandstones.  Marine  invertebrate  fossils  are  locally  common 
in  this  member,  which  to  date  has  produced  only  a  few  vertebrate  remains. 

Vertebrate  fossils  in  the  San  Diego  Formation  generally  occur  as  single  isolated 
skeletal  elements,  although  occasionally  partial  or  complete  skeletons  are  found.  A  few 
rare  "bone  beds"  containing  concentrated,  unassociated  elements  have  recently  been 
discovered  in  the  formation.  To  date,  the  most  productive  fossil  sites  have  been  in  the 
Mission  Hills  area  near  downtown  San  Diego  and  in  the  eastern  portions  of  National 
City  and  Chula  Vista.  In  these  areas  large-scale  grading  operations  have  provided  very 
extensive  exposures  of  the  San  Diego  Formation,  although  in  most  cases  these  new 
exposures  are  accessible  for  only  a  short  period  of  time  because  of  development  and 
construction. 

Found  in  association  with  the  vertebrates  are  rich  assemblages  of  marine  macroin- 
vertebrates,  primarily  mollusks,  which  provide  both  biostratigraphic  and  paleoenvi- 
ronmental  control.  The  "lower"  member  contains  a  middle  to  outer  shelf  molluscan 
fauna  characterized  by  Patinopecten  healeyi  (Arnold),  Pecten  stearnsii  Dall,  Lucinoma 
annulata  (Reeve),  and  Opalia  varicostata  (Steams).  In  contrast,  mollusks  from  the 
"upper"  member  indicate  deposition  in  littoral  to  inner  shelf  depths.  Characteristic 
species  include  Pecten  bellus  (Conrad),  Argopecten  hakei  (Hertlein),  and  Nucella  la- 
mellosa  Gmelin  along  with  the  echinoid  Dendraster  ashleyi  (Arnold).  Both  members 
reflect  normal  marine  deposition  in  a  broad  coastal  embayment  probably  similar  to 
present-day  Monterey  Bay  along  the  central  California  coast. 

Fossil  remains  of  Hydrodamalis  cuestae  have  been  recovered  from  both  the  "low- 
er" and  "upper"  members  of  the  San  Diego  Formation.  As  presently  understood,  the 
aggregate  vertebrate  faunule  from  the  "lower"  member  consists  of  7  species  of  sharks 
and  rays,  more  than  50  species  of  bony  fishes  (most  of  which  are  known  only  from 
otoliths),  24  species  of  sea  birds,  an  otariid  and  a  dusignathine  (odobenid)  pinniped, 
7  odontocetes,  10  mysticetes,  and  the  sirenian.  A  few  terrestrial  mammal  taxa  including 
horse  {Equus  species),  camel  (cf.  Titanotylopus  species),  peccary  (cf.  Platygonus  species; 
M.  O.  Woodbume,  personal  communication),  and  gomphothere  {Stegomastodon  cf.  S. 
rexroadensis  Woodbume)  occur  in  association  with  the  diverse  marine  vertebrate  as- 
semblage. The  "upper"  member  has  produced  only  three  vertebrate  taxa  to  date:  an 
albatross  {Diomedea  species),  a  mysticete  (cf.  Balaenopteridae),  and  the  sirenian. 

Bames  (1976)  has  correlated  the  San  Diego  Formation  ("lower"  member)  with  the 
Blancan  North  American  Land  Mammal  Age.  This  correlation,  based  on  the  occurrence 
of  the  horse  Equus,  is  supported  by  the  recent  discovery  of  teeth  referable  to  the 
gomphothere  Stegomastodon  cf.  S.  rexroadensis  (M.  O.  Woodbume,  personal  com- 
munication). Relying  on  the  stratigraphic  ranges  of  molluscan  species,  the  "lower" 
member  is  correlative  with  the  San  Joaquin  Formation  in  the  San  Joaquin  Basin,  the 
Careaga  Formation  in  the  Santa  Maria  Basin,  and  the  Niguel  Formation  and  the  upper 
Femando  Formation  in  the  southeastem  Los  Angeles  Basin,  all  Late  Pliocene  in  age 
(Woodring  and  Bramlette  1950,  Vedder  1972).  In  tum,  the  "upper"  member  of  the 
San  Diego  Formation  correlates  with  the  lower  part  of  the  Santa  Barbara  Formation 
in  the  Ventura  Basin,  which  is  considered  to  be  Late  Pliocene  to  Early  Pleistocene  in 
age  (Keen  and  Bentson  1944).  The  meager  microfossil  evidence  available  (Ingle  1967, 
Mandel  1973)  suggests  that  the  San  Diego  Formation  is  apparently  no  older  than 
planktonic  foraminiferal  zone  N.21  (approximately  3.0  million  years  B.P.,  Late  Plio- 
cene) and  is  perhaps  as  young  as  the  Emiliania  annula  calcareous  nannoplankton 
subzone  (approximately  1.5  million  years  B.P.,  Early  Pleistocene;  E.  D.  Milow,  personal 
communication).  This  Pleistocene  correlation  is  tenuous  at  the  moment  and  must  await 
completion  of  additional  field  work  for  confirmation.  It  should  not  be  interpreted  to 
mean  that  the  entire  "upper"  San  Diego  Formation  is  Pleistocene. 


175 


Systematics 

Order  Sirenia 

Family  Dugongidae 

Subfamily  Hydrodamalinae 

Hydrodamalis  cuestae  Domning,  1978 

Material 

San  Mateo  Formation 

San  Luis  Rev  River  Local  Fauna  (of  Barnes  et  al.  1981).— 
SDSNH  locality  2957  (=UCMP  loc.  V68144),  Loretta  St.,  Oceanside: 

SDSNH  22655:     Two  rib  fragments.  Coll.  J.  W.  Tobiska,  I  1979. 
SDSNH  locality  3004  (=UCMP  loc.  V68145),  Loretta  St.,  Oceanside: 

SDSNH  21076:     Juvenile  left  maxilla.  Coll.  T.  A.  Demere,  15  III  1980. 
SDSNH  locality  3134  (=UCMP  loc.  V68147),  Lawrence  Canyon,  Oceanside: 

SDSNH  23384:     Proximal  end  of  rib.  Coll.  R.  A.  Cerutti,  22  III  1981. 

Lawrence  Canyon  Local  Fauna  (of  Barnes  et  al.  1981).— 
SDSNH  locality  3161  (=UCMP  loc.  V68106),  Lawrence  Canyon,  Oceanside: 
SDSNH  24413 


SDSNH  24454 
SDSNH  24685 


Fragment  of  juvenile  left  maxilla.  Coll.  SDSNH  party,  VII  1982. 
Rib  fragment.  Coll.  SDSNH  party,  VII  1982. 
Immature  right  humerus.  Coll.  B.  O.  Riney  and  R.  A.  Cerutti,  26 
VI  1982. 
SDSNH  24686:     Proximal  end  of  juvenile  ?right  ulna.  Coll.  SDSNH  party,  VII 

1982. 

San  Diego  Formation 

'  'Lo  wer ' '  Mem  ber.  — 

SDSNH  locality  2970-B,  Washington  St.,  San  Diego: 

SDSNH  21685:     Rib  fragment.  Coll.  R.  A.  Cerutti,  IX  1980. 

SDSNH  locality  3148,  California  St.,  San  Diego: 

SDSNH  23719:     Vertebrae  CI -2,  C6-7,  Tl-?6.  Coll.  R.  A.  Cerutti  and  T.  A.  De- 
mere, 16  II  1981. 

SDSNH  locality  3172,  Florida  Canyon,  San  Diego: 

SDSNH   24679:     Two  immature  thoracic  vertebrae.  Coll.  R.  A.  Cerutti  and  T.  A. 

Demere,  2  XII  1981. 

SDSNH  locality  3 1 74,  H  Street  extension,  Chula  Vista: 

SDSNH  24683:     AduU  right  rib.  Coll.  R.  A.  Cerutti  and  B.  O.  Riney,  24  VIII  1982. 

SDSNH  locality  3175,  Adams  Ave.,  San  Diego: 

SDSNH  24684:     Anterior  end  of  immature  left  mandible.  Coll.  R.  H.  Norwood, 

1975. 
SDSNH  24687:     Rib  fragment.  Coll.  R.  H.  Norwood,  1975. 

SDSNH  locality  3158,  Hidden  Vista,  Chula  Vista: 

SDSNH  23726:     Adult  braincase.  Coll.  R.  A.  Cerutti,  15  IX  1981. 


176 


SDSNH  2468 1 :     Distal  ends  of  immature  ?right  radius  and  ulna.  Coll.  R.  A.  Cerutti, 

20  VII  1981. 
SDSNH  24682:     Seven  rib  fragments.  Coll.  R.  A.  Cerutti,  22  VII  1981. 

"Upper"  Member.  — 

SDSNH  locality  3173,  38th  and  Beech  Streets,  San  Diego: 

SDSNH  24680:     Neural  arch  of  immature  thoracic  vertebra  {Hvdrodamalis  species 

indet.).  Coll.  T.  A.  Demere,  21  I  1983. 

Description  of  Specimens  from  the  San  Mateo  Formation 

A/axz7/a.  — A  juvenile  left  maxilla  (SDSNH  21076,  Fig.  2a-e),  as  noted  by  Barnes 
et  al.  (1981),  is  almost  identical  to  the  right  maxilla  (UCMP  86345)  previously  reported 
from  the  same  locality  (Domning  1978:  tab.  24;  pi.  17,  fig.  a).  It  is  exactly  the  same 
size  and  represents  the  same  growth  stage,  but  is  more  complete,  measuring  129  mm 
in  overall  length  and  58  mm  in  height.  (It  may  even  represent  the  same  individual.) 
The  first  alveolus,  for  a  single-rooted  tooth,  lies  slightly  posterior  to  the  zygomatic- 
orbital  bridge  and  is  7.5  mm  long.  The  second  alveolus,  for  a  two-rooted  tooth,  is  8.5 
mm  long,  and  the  third,  for  a  three-rooted  tooth  with  apparently  coalesced  roots,  is 
about  9  mm  long  and  7  mm  wide.  The  combined  length  of  these  three  alveoli  is  27 
mm.  Posterior  and  dorsal  to  the  third  alveolus  is  a  large  broken  dental  capsule  (1 1  mm 
in  dorsoventral  height)  for  an  unerupted  tooth.  The  narrowest  part  of  the  palatal  surface 
(20  mm  wide,  left  half  only)  lies  just  forward  of  the  zygomatic-orbital  bridge,  and 
consists  of  a  flat  surface  bordered  medially  by  a  shallow  palatal  gutter  (containing  a 
foramen  anteriorly  and  ending  posteriorly  in  another  foramen)  and  laterally  by  a  sharp 
edge  where  it  meets  the  lateral  surface  perpendicularly.  The  zygomatic-orbital  bridge 
is  elevated  19  mm  above  the  palatal  surface,  and  is  30  mm  long  anteroposteriorly  and 
10  mm  thick.  The  vertical  plate  of  the  bridge,  which  articulated  with  the  jugal,  is  55 
mm  in  height  and  contacts  the  main  body  of  the  maxilla  above  the  infraorbital  foramen; 
the  latter  is  invisible  in  ventral  view.  On  the  ventromedial  wall  of  the  foramen  are  the 
anterior  and  posterior  openings  of  a  short,  horizontal  canal  (a  continuation  of  the 
premaxillary  canal);  medial  to  the  posterior  opening  is  a  larger  pit  or  canal  opening 
posteriorly.  The  palate  is  a  maximum  of  19  mm  thick. 

A  fragmentary  left  maxilla  of  a  larger  juvenile  (SDSNH  2441 3)  has  a  palatal  surface 
which  slopes  dorsolaterad  just  forward  of  the  zygomatic-orbital  bridge,  as  in  more 
mature  individuals  of  Hydrodamalis.  At  the  level  of  the  front  side  of  the  bridge,  this 
surface  is  only  18  mm  wide,  proportionately  narrower  than  in  the  above  specimen. 
The  lateral  surface  bears  a  prominent  groove  (continuation  of  the  premaxillary  canal) 
anterior  to  the  infraorbital  foramen.  The  palate  reaches  a  thickness  of  31  mm,  and  the 
intermaxillary  suture  bears  numerous  deep  vertical  interdigitations.  No  dental  alveoli 
are  preserved. 

Dentition.  — TttXh  o^  Hydrodamalis  cuestae  are  still  unknown,  but  the  essentially 
complete  maxilla  described  above  (SDSNH  21076)  allows  us  to  speculate  on  the  ho- 
mologies of  the  teeth  present  in  the  juvenile.  The  pattern  of  roots  of  the  three  fully 
erupted  teeth  matches  that  of  the  teeth  identified  as  DP-""  in  Dusisiren  (Domning  1 978); 
this  implies  that  the  dental  capsule  contained  DP^  In  view  of  the  immediate  descent 
of  H.  cuestae  from  Dusisiren  (Domning  1978),  this  seems  the  most  parsimonious 
interpretation.  However,  in  living  Dugong  (Marsh  1980)  and  Trichechus  (Domning 
1 982),  the  three  most  anterior  cheek  teeth  of  the  juvenile  appear  to  be  the  only  premolars 
(DP'-5).  A  fuller  growth  series  of//,  cuestae  is  needed  to  settle  this  question  conclusively, 
and  also  to  determine  whether  Blancan  as  well  as  Hemphillian  juveniles  of  the  species 
still  possessed  teeth. 

Humerus.— \n  immature  right  humerus  (SDSNH  24685,  Fig.  5)  is  complete  except 
for  the  unfused  epiphyses.  In  size,  it  falls  between  the  two  immature  humeri  of  Hy- 
drodamalis gi  gas  described  by  Whitmore  and  Gard  ( 1 977);  in  shape,  it  closely  resembles 


77 


Figure  2.  Hydrodamalis  cuestae.  a-e,  juvenile  left  maxilla.  SDSNH  21076.  a,  lateral  view;  b,  medial  view, 
c,  dorsal  view;  d,  ventral  view;  e,  posterior  view,  f-h,  atlas.  SDSNH  23719.  f,  dorsal  view;  g,  anterior  view; 
h,  posterior  view.  Scales  =  4  cm. 


178 


Table  1.  Skull  measurements  of  Hydrodamalis  cuestae  (SDSNH  23726)  from  the  San  Diego  Formation, 
in  millimeters.  Letters  in  parentheses  denote  measurements  used  by  Domning  (1978:  tab.  2,  fig.  7);  e  = 
estimated. 

Rear  of  occipital  condyles  to  anterior  end  of  interfrontal  suture  (BI)  »       4 1 8e 

Length  of  interfrontal  suture  143e 

Length  of  skull  roof  to  rear  of  external  occipital  protuberance  310  + 

Top  of  supraoccipital  to  ventral  side  of  occipital  condyle  (de)  223 

Breadth  across  occipital  condyles  (fF)  276e 

Width  of  supraoccipital  250 

Height  of  supraoccipital  to  top  of  external  occipital  protuberance  120e 

Breadth  of  cranium  at  frontoparietal  suture  (GC)  123 

Width  of  foramen  magnum  (gg')  125e 

Height  of  foramen  magnum  (hi)  64 

Minimum  width  of  basioccipital  78 

Length  of  right  zygomatic  process  of  squamosal  (O'P')  229  + 

Anterior  tip  of  zygomatic  process  to  rear  edge  of  squamosal  below  mastoid  foramen  (O'T')  293  + 

Frontoparietal  suture  to  rear  of  external  occipital  protuberance  (P)  169e 

Anteroposterior  length  of  root  of  zygomatic  process  (Q'R')  103 

Length  of  cranial  portion  of  squamosal  (S'T')  196  + 

Posterior  height  of  cranial  portion  of  squamosal  (U'V)  182 

Dorsoventral  breadth  of  zygomatic  process  (W'X')  1 17 


the  smaller  of  these  (USNM  186807),  differing  from  the  other  (USNM  170761)  in 
lacking  a  pronounced  "shoulder"  above  the  ectepicondyle.  The  deltoid  crest  is  massive 
and  rounded,  without  a  recurved  flange.  The  total  length  (without  epiphyses)  is  291 
mm;  the  proximal  breadth,  130  mm;  and  the  distal  breadth,  156  mm. 

Ulna.— T\iQ  proximal  end  of  a  juvenile  ?right  ulna  (SDSNH  24686)  has  a  width 
of  65  mm;  the  shaft  is  thicker  anteroposteriorly  than  mediolaterally.  It  compares  well 
in  proportions  with  the  ulna  of  a  much  larger  immature  H.  gigas  (USNM  170761) 
from  Amchitka  (Whitmore  and  Gard  1977:pl.  8,  figs.  3-5). 

Ribs.—ThQ  proximal  end  of  a  subaduh  right  rib  (SDSNH  22655)  is  very  flat  (65  x 
34  mm)  and  has  a  tiny  tubercle  whose  lateral  edge  is  74  mm  from  the  tip  of  the 
capitulum.  The  distal  end  of  an  immature  rib  (SDSNH  24454)  measures  59  x  54  mm 
in  diameter  and  has  a  broad,  concave,  rugose  distal  surface  for  cartilage  attachment. 
A  zone  of  cancellous  bone  is  partly  exposed  by  bone  resorption  on  its  medial  side.  The 
proximal  end  of  a  larger,  more  posterior  left  rib  (SDSNH  23384)  has  the  capitulum 
and  tubercle  coalesced.  Both  it  and  SDSNH  22655  are  completely  dense  where  broken. 

Description  of  Specimens  from  the  San  Diego  Formation 

Skull.— A  nearly  complete  adult  braincase  (SDSNH  23726,  Fig.  6,  Table  1),  in- 
cluding the  right  zygomatic  process  of  the  squamosal  but  lacking  the  pterygoid  processes, 
expands  our  knowledge  of  the  occipital  region  of  Hydrodamalis  cuestae.  The  interior, 
however,  is  still  filled  with  very  hard  matrix,  precluding  examination  of  the  internal 
structures.  The  dimensions  of  the  preserved  portions  indicate  that  this  individual  was 
larger  than  any  other  Hydrodamalis  or  other  sirenian  ever  found,  and  in  life  no  doubt 
measured  well  over  10  m  in  body  length. 

Frontal:  The  supraorbital  processes  are  missing,  but  the  anterior  border  seems  to 
be  complete.  It  is  arched  upward  somewhat  at  the  midline,  due  to  a  broad,  prominent 
median  boss  on  the  dorsal  surface.  Lateral  to  this  are  depressions  bordered  by  upraised 
forward  extensions  of  the  temporal  crests.  The  lateral  surfaces  drop  almost  perpendic- 
ularly from  these  crests,  with  little  or  no  overhang. 

Parietal:  The  cranial  vault  is  square  anteriorly  with  no  overhang  of  the  temporal 
crests,  which  are  nearly  parallel  anteriorly  but  posteriorly  form  sharp  ridges  which 
diverge  and  descend  to  meet  the  dorsal  ends  of  the  sigmoid  ridges  of  the  squamosals. 
There  is  only  a  slight  indentation  in  the  dorsal  surface  where  it  meets  the  top  of  the 


179 

squamosal,  much  as  in  H.  gigas.  The  parietal  roof  is  slightly  concave  anteriorly  and 
flat  posteriorly. 

Supraoccipital:  The  width/height  ratio  is  2.08,  being  within  the  previously  reported 
range  of  variation.  The  supraoccipital  forms  an  angle  of  124°  with  the  after  part  of  the 
parietal  roof  and  is  within  the  range  of  variation  of  the  USNM  sample  of  Bering  Island 
H.  gigas.  The  external  occipital  protuberance  is  low  and  indistinct,  with  no  median 
ridge  extending  below  it.  The  lateral  borders  are  rounded,  rugose,  and  extremely  thick 
and  massive. 

Exoccipital:  The  sutures  with  the  supraoccipital  are  indistinct,  but  the  exoccipitals 
apparently  fail  to  meet  in  the  dorsal  midline.  The  dorsal  border  of  the  foramen  magnum 
is  very  gently  arched  with  almost  no  dorsal  peak,  as  in  H.  gigas.  The  dorsolateral  border 
of  the  exoccipital  is  very  thick,  rounded,  and  rugose,  and  overhangs  posteriorly,  but  is 
not  expanded  to  the  degree  seen  in  H.  gigas.  However,  it  does  confirm  the  supposition 
(Domning  1978)  that  the  thin  exoccipital  border  seen  in  the  holotype  of//,  cuestae 
(UCMP  86433)  is  abnormal  for  the  species.  The  paroccipital  process  is  massive  and 
rugose.  The  occipital  condyle  is  very  large  and  broad,  and  projects  well  abaft  the  occiput. 
Its  upper  part  curves  forward  only  slightly  and  there  is  no  supracondylar  fossa,  again 
confirming  the  abnormality  of  the  holotype.  The  condyle  measures  78  mm  in  width 
and  1 17  mm  front  to  back,  a  ratio  of  0.67  (intermediate  between  values  reported  for 
Dusisiren  and  H.  gigas  by  Domning  1978). 

Basioccipital:  Completely  fused  with  basisphenoid  and  exoccipitals,  indicating 
adulthood. 

Basisphenoid:  Largely  removed  by  bulldozer,  but  originally  much  thicker  than  in 
the  (abnormal)  holotype. 

Squamosal:  The  sigmoid  ridge  is  about  as  reduced  as  in  H.  gigas,  and  is  invisible 
in  posterior  view.  The  cranial  portion  dorsal  to  the  zygomatic  root  is  very  slightly 
concave,  as  in  the  holotype.  There  is  a  protuberance  about  1  cm  high  just  above  the 
external  auditory  meatus,  an  individual  peculiarity  not  observed  in  the  related  forms. 
The  temporal  condyle  is  broad,  smooth,  and  not  distinctly  demarcated;  the  postarticular 
fossa  is  broad  and  well  developed;  and  the  postglenoid  process  is  about  2  cm  high  and 
very  robust,  unlike  H.  gigas.  The  processus  retroversus  is  straight  and  very  prominent, 
with  a  deep  posterior  indentation  as  in  the  holotype.  The  posterodorsal  edge  of  the 
zygomatic  process  is  damaged,  giving  the  process  a  more  lozenge-shaped  outline  than 
it  originally  had;  it  may  have  approached  that  of  H.  gigas  in  convexity.  The  forward 
end  of  the  process  is  missing;  its  lateral  edge  was  sharper  than  its  medial.  Its  underside 
bears  a  clear  impression  for  the  zygomatic  process  of  the  jugal,  which  extended  back 
almost  to  the  level  of  the  forward  edge  of  the  zygomatic  root.  Just  posterior  to  the  tip 
of  the  jugal  is  a  large  smooth  convexity  not  so  prominent  in  other  Hydrodamalis 
specimens. 

Periotic:  Present  on  right  side  but  not  prepared. 

Tympanic:  A  fragment  is  present  on  the  right  side,  just  posterior  to  the  auditory 
meatus. 

Mandible.  — ThQ  anterior  end  of  an  immature  left  mandible  (SDSNH  24684).  badly 
worn,  shows  a  H.  gigas-\\\iQ  convex  outline  of  the  anteroventral  border.  The  mental 
foramen  appears  to  have  lain  well  forward  (at  the  level  of  the  symphysis),  as  assumed 
for  the  holotype  of  H.  cuestae. 

Vertebrae.— Ttn  of  the  anterior  vertebrae  are  known  from  SDSNH  23719,  an 
individual  smaller  than  SDSNH  23726.  Partial  thoracics  from  other  animals  (SDSNH 
24679,  24680)  are  also  available. 

Atlas  (Fig.  2f-h,  Table  2):  The  upper  arch  lacks  a  keel  and  articular  surface  for 
the  axis,  but  has  an  anterior  median  notch  flanked  by  a  pair  of  rugose  protuberances, 
and  is  penetrated  posteriorly  by  a  vertical  canal  about  4  mm  wide.  The  canal  for  the 
first  cervical  nerve  atop  each  cotyle  is  deep  but  not  bridged  by  bone.  A  possible  vestige 
of  a  vertebral  arterial  canal  on  each  side,  filled  with  matrix,  may  not  have  been  patent; 
in  any  case  there  is  no  distinct  notch  on  the  transverse  process  as  in  Dusisiren  jordani. 

Axis  (Fig.  3a-c,  Table  3):  The  odontoid  process  bears  a  broad,  smooth,  saddle- 


180 


Table  2.     Measurements  of  atlas  of  Hydrodamal is  cuestae  (SDSNH  237 1 9)  from  the  San  Diego  Formation, 
in  millimeters;  e  =  estimated. 


External  height 

Internal  height 

Total  breadth 

Width  between  tips  of  processes  for  transverse  ligament 

Breadth  across  anterior  cotyles 

Breadth  across  posterior  cotyles 

Length  in  dorsal  midline 

Length  in  ventral  midline 


181 
U5 

292e 

78 

219 

190 

42 

49 


Table  3.     Measurements  of  axis  of  Hydrodamalis  cuestae  (SDSNH  237 19)  from  the  San  Diego  Formation, 
in  millimeters;  e  =  estimated. 


Total  height 

Tip  of  odontoid  process  to  rear  of  centrum 

Breadth  across  cotyles 

Breadth  of  cotyle 

Height  of  cotyle 

Posterior  breadth  of  centrum 

Posterior  height  of  centrum 

Width  of  neural  canal 

Height  of  neural  canal 

Breadth  across  postzygapophyses 


196  + 

102 

182 

54 

85 
116 

71 

88 

95e 
167 


Table  4.     Measurements  of  cervical  and  thoracic  vertebrae  of  Hydrodamalis  cuestae  (SDSNH  23719)  from 
the  San  Diego  Formation,  in  millimeters;  e  =  estimated;  a  =  asymmetrical. 


C6 


C7 


Tl   ?T2   ?T3 


7X4   ?T5   ?T6 


Total  height 

Breadth  across  transverse  processes 
Anterior  breadth  of  centrum 
Posterior  breadth  of  centrum 
Height  of  centrum  in  midline 
Thickness  of  centrum  in  midline 
Width  of  neural  canal 
Height  of  neural  canal 
Breadth  across  prezygapophyses 
Breadth  across  postzygapophyses 
Length  from  front  of  prezygapophysis 
to  rear  of  postzygapophysis 


180 

206 

219  + 

306e 

355a 

344  + 

345  + 

354 

126 

135 

132 

125 

137 

153 

160 

131 

123 

128 

137 

156 

161e 

165 

82 

81 

76 

81 

79 

95 

97 

100 

33 

38 

56 

69 

69 

71 

75 

77 

124 

120 

131 

119 

116 

116 

112 

81 

81 

91 

81 

81 

93 

249 

240e 
218 

240e 

173e 
169 

170e 
178 

::: 

58 


72 


76 


82+      105e       llOe 


shaped  articular  surface  ventrally.  The  vertebral  arterial  canal  is  broadly  open  as  in  the 
holotype.  The  neural  spine  is  not  preserved. 

Cervicals  6-7  (Fig.  3d-e,  Table  4):  The  vertebral  arterial  canal  is  large  in  C6  and 
on  the  right  side  of  C7,  but  open  laterally  on  the  left  side,  which  lacks  the  fused  cervical 
rib;  an  anterior  demifacet  and  a  tubercular  articulation,  apparently  for  the  cervical  rib, 
are  present.  A  similar  asymmetry  was  observed  in  one  D.  Jordan!  vertebra  (Domning 
1978:pl.  9,  fig.  h).  The  transverse  process  of  C6  juts  farther  laterad  than  in  the  corre- 
sponding vertebra  of  D.  jordani. 

Thoracics  l-?6  (Fig.  3f-g,  4,  Table  4):  The  anterior  thoracic  centra  have  square 
sagittal  sections  like  H.  gigas  and  unlike  the  holotype,  and  sieve-like  epiphyses  as  in 
the  Recent  species.  The  neural  arch  of  Tl  does  not  have  "shoulders"  like  D.  jordani 


181 


Figure  3.  Hydrodamalis  cuestae.  Vertebrae.  SDSNH  23719.  a-c,  axis,  a,  anterior  view;  b,  lateral  view;  c, 
posterior  view;  d,  C6,  posterior  view;  e,  C7,  posterior  view,  f-g,  Tl.  f,  posterior  view;  g,  lateral  view.  Scale  = 
4  cm. 


182 


•^ 


.-^'^ 


-^t*. 


-  ri.r 


"•^ 


Figure  4.     Hydrodamalis  cuestae.  thoracic  vertebrae.  SDSNH  23719.  Posterior  views:  a,  ?T2;  b,  ?T3;  c, 
?T4;  d,  ?T5;  e,  ?T6.  f,  ?T6,  lateral  view.  Scale  =  4  cm. 


183 


Figure  5.     Hydrodamalis  cuestae.  Immature  right  humerus.  SDSNH  24685.  a,  anterior  view;  b,  medial 
view.  Scale  =  4  cm. 


or  the  holotype.  The  protuberances  for  semispinaUs  tendons  are  indistinct.  The  neural 
spines  are  incUned  backward  and  are  much  thicker  posteriorly  than  at  their  anterior 
edges,  in  the  manner  of//,  gigas  rather  than  Dusisiren.  The  posterior  sides  of  the  spines 
are  concave  with  distinct  median  ridges.  The  apices  of  the  neural  canals  are  not  slit- 
like. SDSNH  24680,  a  partial  neural  arch,  represents  an  immature  animal  and  was 
apparently  not  fused  to  its  centrum.  A  poorly  preserved  pair  of  thoracic  vertebrae  with 
arches  and  centra  unfused  (SDSNH  24679)  represents  a  still  younger  animal;  the  max- 
imum breadth  of  one  of  these  centra  is  127  mm,  and  its  thickness  is  48  mm. 

Ribs.  — One  nearly  complete  right  adult  rib  (SDSNH  24683,  Fig.  7a)  and  several 
fragmentary  ones  (SDSNH  21685,  24682,  24687)  have  been  recovered  from  the  San 
Diego  Formation.  SDSNH  24683  is  from  near  the  middle  of  the  thorax;  it  lacks  the 
capitulum,  but  would  have  measured  about  1002  mm  in  total  straight-line  length.  At 


184 


Figure  6.     Hydrodamalis  cuestae.  Adult  braincase,  SDSNH  23726.  a,  dorsal  view;  b,  ventral  view;  c,  lateral 
view;  d,  posterior  view.  Left  occipital  condyle  restored.  Scale  =  10  cm. 


185 


Figure  7.     Hydrodamalis  cuestae.  a,  adult  right  rib.  SDSNH  24683.  Scale  =  10  cm.  b,  immature  distal  ?right 
radius  and  ulna,  ?medial  view.  SDSNH  24681.  Scale  in  cm. 


the  middle  of  the  shaft  it  measures  88  x  68  mm;  the  distal  end  is  swollen  to  98  x  78 
mm,  and  slightly  swept  back.  The  distal  tip  bears  an  oval  rugose  concavity,  and  there 
is  no  well-marked  angle.  There  is  also  no  zone  of  cancellous  bone  visible  on  the  medial 
surface.  SDSNH  24682  (seven  associated  adult  rib  fragments)  includes  a  damaged 
proximal  end  and  two  complete  distal  ends;  the  latter  taper  more  gradually  than  SDSNH 
24683  and  are  more  flattened  mediolaterally  (90  x  39  mm,  25  cm  from  distal  end  of 
one  fragment).  Except  for  the  proximal  end  and  another  fragment  of  a  distal  tip,  these 
fragments  are  all  composed  wholly  of  dense  bone.  Another  fragment  (SDSNH  24687) 
from  the  middle  of  a  large  rib  (92  x  72  mm)  has  a  large  zone  of  cancellous  bone  just 
beginning  to  crop  out  on  its  medial  side. 


186 


Radius- ulna.  — SDSNH  24681  (Fig.  7b)  comprises  the  distal  ?right  radius  and  ulna 
of  an  immature  animal,  lacking  the  epiphyses.  The  shaft  of  the  radius  is  flattened 
anteroposteriorly  (about  60  x  40  mm)  and  is  concave  both  mediolaterally  and  prox- 
imodistally  on  its  posterior  surface;  it  is  also  slightly  bowed  (concave  ?medially).  Its 
distal  end  is  expanded  to  a  width  of  78  mm  and  an  anteroposterior  thickness  of  75 
mm,  with  a  flat  posterior  surface.  The  anterior  surface  bears  three  low,  irregular  knobs 
about  1.5,  4,  and  10  cm,  respectively,  from  the  distal  end.  The  shaft  of  the  ulna  is 
compressed  mediolaterally  and  markedly  bowed  (concave  posteriorly);  its  expanded 
distal  end  measures  71  mm  mediolaterally  and  an  estimated  75-80  mm  anteroposte- 
riorly, with  a  flat  anterior  surface.  The  distal  thickness  from  the  anterior  side  of  the 
radius  to  the  posterior  side  of  the  ulna,  allowing  for  the  interosseous  space  whose  width 
is  indicated  by  fragments  of  matrix,  was  about  155  mm. 

Discussion 

The  specimens  herein  described  are  distinguished  from  Dusisiren  Domning  1978 
by  the  following  derived  characters:  In  the  case  of  those  from  the  San  Mateo  Formation, 
their  large  adult  size,  reduced  juvenile  dentition,  and  invisibility  of  the  infraorbital 
foramen  in  ventral  view;  in  the  case  of  those  from  the  San  Diego  Formation,  their 
large  size,  lack  of  a  dorsal  peak  to  the  foramen  magnum,  reduced  sigmoid  ridge  on  the 
squamosal,  lack  of  a  dorsal  articulation  between  the  atlas  and  axis,  stouter  and  more 
posteriorly  inclined  thoracic  neural  spines,  presence  of  a  core  of  cancellous  bone  in 
immature  ribs,  and  greater  curvature  of  the  ulna.  Primitive  characters  of  these  speci- 
mens separating  them  from  Hydrodamalis  gigas  include:  A  square  cranial  vault  with 
more  distinct  temporal  crests,  less  expanded  occipital  borders,  a  posteriorly  notched 
zygomatic  root,  a  large  postglenoid  process,  a  more  anteriorly  located  mental  foramen, 
and  retention  of  teeth  in  the  juvenile  (at  least  in  the  Hemphillian  specimens).  In  some 
details  (zygomatic  process  possibly  with  rounded  posterior  end,  thoracic  centra  square 
in  sagittal  section,  first  thoracic  neural  arch  without  "shoulders")  the  new  specimens 
from  the  San  Diego  Formation  are  more  derived  than  previously  known  H.  cuestae, 
while  in  another  respect  (well-developed  postarticular  fossa  of  squamosal)  they  are 
more  primitive.  However,  such  characters  may  be  expected  to  vary  from  more  "prim- 
itive" to  more  "derived"  conditions  in  a  single  population  at  any  given  time.  In  most 
respects  the  new  specimens  most  closely  resemble  the  previously  described  specimens 
of//,  cuestae.  They  are,  accordingly,  referred  to  that  species,  with  the  possible  exception 
of  the  fragmentary  vertebra  (SDSNH  24680)  from  the  highest  unit,  which  is  best 
regarded  as  Hydrodamalis  species  indet. 

Conclusions  reached  earlier  by  Domning  (1 978)  are  supported  by  the  new  material: 

1.  The  holotype  of//,  cuestae  is  abnormal  in  degree  of  development  of  the  exoccipital 
border,  supracondylar  fossa,  and  basisphenoid.  The  skull  from  the  San  Diego  For- 
mation shows  the  conditions  expected  in  a  form  phylogenetically  intermediate  be- 
tween D.  jordani  and  H.  gigas. 

2.  Hydrodamalis  grew  larger  in  the  southern  parts  of  its  range  than  in  the  marginal 
habitat  of  the  Commander  Islands.  The  San  Diego  skull  is  the  largest  of  any  indi- 
vidual Hydrodamalis  or  other  sirenian  ever  discovered,  and  several  postcranial 
elements  also  appear  to  set  new  size  records. 

Additional  conclusions  are  permitted  by  the  new  material: 

1 .  The  sirenian  present  in  the  San  Diego  Formation  is  indeed  //.  cuestae,  as  would  be 
predicted  from  the  age  of  the  unit. 

2.  Hemphillian-aged  juveniles  of  H.  cuestae  possessed  at  least  four  upper  teeth,  prob- 
ably DP2-5. 

Acknowledgments 

The  junior  author  benefited  greatly  from  discussions  with  Robert  M.  Chandler 
(San  Diego  Natural  History  Museum)  concerning  West  Coast  Tertiary  avian  assem- 
blages. 


187 


The  cooperation  of  the  following  companies  is  gratefully  acknowledged  in  allowing 
and  supporting  the  collection  of  fossils  on  their  properties:  Vance  Johnson,  Inc.,  Es- 
condido,  California;  Watt  Industries,  Rancho  Santa  Fe,  California;  Financial  Scene, 
San  Diego,  California;  and  The  Gersten  Companies,  Chula  Vista,  California.  In  addition 
the  cities  of  Oceanside,  San  Diego,  and  Chula  Vista  are  commended  for  their  recognition 
of  the  importance  of  salvaging  and  preserving  paleontological  resources  within  their 
respective  jurisdictions. 

The  senior  author's  work  on  this  project  was  supported  by  National  Science  Foun- 
dation grant  #DEB  80-20265.  The  junior  author's  work  was  supported  in  part  by  grants 
from  the  Parker  Foundation,  the  Scripps  Foundation,  and  the  J.  W.  Sefton  Foundation. 
The  additional  support  of  Joseph  and  Joanne  Parker  is  also  gratefully  acknowledged. 

We  thank  L.  G.  Barnes,  C.  A.  Repenning,  and  an  anonymous  reviewer  for  their 
helpful  comments  on  the  manuscript. 


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(^^^     ^^^  TRANSACTIONS 

OF  THE  SAN  DIEGO 
SOCIETY  OF 
NATURAL  HISTORY 


Volume  20  Number  13  pp.  189-24Qj^/^  20  November  1984 


Fossil  Syncarida  ju^     ^A/^y^^O(^ 


**«^!^o 


Frederick  R.  Schram 

Department  of  Geology.  San  Diego  Natural  History  Museum,  San  Diego, 

Abstract.  All  known  fossil  syncaridans  are  reviewed,  and  their  family  level  ta^oRbmy  revised  to 
form  a  more  natural  system.  One  anaspidid  anaspidacean  is  known,  Anaspidites  antiquus  (Chilton), 
from  the  Triassic  of  Australia.  The  northern  hemisphere  Paleozoic  Palaeocaridacea  are  sorted  into  four 
families:  Minicarididae  (Minicaris  brandi  Schram,  Erythrogaulos  carrizoensis  new  genus,  new  species), 
Acanthotelsonidae  {Acanthotelson  stimpsoni  Meek  &  Worthen,  A.  kentuckiensis  new  species,  Uronectes 
fimbriatus  (Jordan),  U.  kinniensis  Schram  &  Schram,  Palaeosyncaris  dakotensis  Brooks,  P.  micra  new 
species),  Palaeocarididae  (Palaeocaris  typus  Meek  &  Worthen,  P.  retractata  Caiman,  P.  secretanae  new 
species),  and  Squillitidae  (Squillites  spinosus  Scott,  Praeanaspides  praecursor  Woodward,  Nectotelson 
krejcii  (Fritsch)).  Several  taxa  are  too  incompletely  known  to  be  placed  with  certainty  at  this  time  within 
these  families:  Pleurocaris  annulatus  Caiman,  Williamocalmania  vandergrachti  (Pruvost),  Brooksyn- 
caris  canadensis  (Brooks),  Palaeorchestia  parellela  (Fritsch),  and  Clarkecaris  brasilicus  (Clarke).  An 
analysis  of  phylogenetic  relationships  of  syncaridan  families  is  presented. 


Introduction 

It  is  a  historical  curiosity  that  syncarids  were  known  as  fossils  45  years  before  they 
were  discovered  living  in  Tasmania.  The  understanding  of  their  relationship  to  other 
eumalacostracans  has  unfolded  only  gradually,  and  is  still  not  completely  resolved  (see 
e.g.,Dahl  1983;  Hessler  1983;  Schram  1981c.  1984;  Watling  1981,  1983).  The  syncarids 
remain  one  of  the  most  singularly  interesting  groups  within  the  Eumalacostraca. 

The  first  syncarid,  a  Permian  fossil,  was  described  by  Jordan  (1 847)  as  Gampsonyx 
fimbriatus  (now  known  as  Uronectes  fimbriatus).  The  species  was  immediately  recog- 
nized by  Jordan  as  unusual,  though  he  compared  it  to  amphipods  in  terms  of  its  general 
form.  The  question  of  its  exact  systematic  position,  however,  could  not  be  definitively 
resolved,  as  evinced  by  Burmeister  (1 855)  who,  in  a  detailed  consideration  of  the  beast, 
made  passing  mention  of  possible  stomatopod  a/7^amphipod  similarities.  Burmeister 
remarked  that  its  closest  affinities  seemed  to  be  with  schizopods,  yet  concluded  it  was 
an  example  of  a  singular  group  ("sie  ist  vielmehr  der  Reprasentiert  einer  besondem 
Gruppe,"  p.  200).  Roemer  (1856)  had  no  such  reservations  and  placed  this  species 
within  the  Stomatopoda. 

Subsequently,  Meek  and  Worthen  (1865)  described  2  more  "syncarid"  species, 
Acanthotelson  stimpsoni  and  Palaeocaris  typus,  and  placed  them  within  the  Isopoda. 

Fritsch  (1870)  described  what  he  thought  was  a  species  related  to  G.  fijnbriatus, 
which  he  called  Gampsonychus  krejcii.  Fritsch  (1876)  also  described  what  he  thought 
was  yet  another  species  o^  ""Gampsonychus,''''  which  was  later  placed  by  Zittel  (1885) 
in  a  separate  genus,  Palaeorchestia  parallela. 

It  was  Packard  (1885,  1886a)  who  finally  recognized  in  part  the  separate  status  of 
these  fossils,  and  erected  the  taxon  Syncarida.  However,  he  placed  only  A.  stimpsoni 
within  this  new  group.  He  then  proceeded  to  compare  ""Gampsonyx''  with  Palaeocaris 
typus,  and  concluded  that  these  latter  taxa  served  ".  .  .  to  bridge  over  the  chasm  existing 
between  the  thoracostracous  suborders,  Syncarida  and  Schizopoda  .  .  .,"  (Packard  1886a: 
129). 


190 

When  Thompson  (1893,  1894)  described  the  living  species  Anaspides  tasmaniae 
he  placed  it  in  a  separate  family  of  the  Schizopoda.  However,  it  was  Caiman  (1896) 
who  realized  the  relationship  oi  Anaspides  to  the  various  fossil  forms  and  united  them 
altogether  in  the  Syncarida,  which  he  later  (1904)  elevated  to  superorder  status  within 
the  Eumalacostraca.  This  arrangement  completely  overshadowed  Grobbea's  (1919) 
attempt  to  erect  a  subdivision  Anomostraca  within  the  Malacostraca  for  Anaspides. 

As  if  to  celebrate  this  apparent  resolution  of  syncarid  affinities,  a  whole  host  of 
new  fossil  species  soon  entered  the  literature:  Praeanaspides  praecursor  Woodward, 
1908;  Pleurocaris  annulatus  Caiman,  1911;  Anaspides  brasilicus  Clarke,  1920;  Palaeo- 
caris  vandergrachti  Pruvost,  1922;  Anaspides  antiquus  Chilton,  1929;  Palaeocaris  re- 
tractata  Caiman,  1932  (actually  known  since  1911);  and  Squillites  spinosus  Scott,  1938 
(a  name  which  mistakenly  resurrected  the  idea  of  supposed  affinities  to  stomatopods). 

The  taxonomy  of  the  group  then  achieved  a  certain  degree  of  stability  until  Brooks 
(1962(2,  b)  recognized  distinct  generic  status  for  Anaspidites  antiquus  and  Clarkecaris 
brasilicus  from  Anaspides,  and  also  recognized  at  that  time  the  separate  status  of  the 
Paleozoic  taxa  with  his  order  Palaeocaridacea.  Brooks  {\962b)  went  on  to  describe  a 
new  species,  Palaeosyncaris  dakotensis,  but  mistakenly  synonymized  (Brooks  1969) 
most  of  the,  until  then  separate.  Palaeozoic  genera  with  the  genus  Palaeocaris. 

A  major  revision  of  the  fossil  syncarids  began  with  a  redescription  of  Squillites 
spinosus  by  Schram  and  Schram  (1974).  Schram  (1979(2)  continued  this  review  by 
reestablishing  the  separate  generic  status  of  several  of  the  Paleozoic  taxa,  at  least  for 
the  British  Carboniferous  fauna,  as  well  as  describing  the  earliest  syncarid,  Minicaris 
brandi.  A  second  species  of  Uronectes,  U.  kiniensis,  was  described  by  Schram  and 
Schram  (1979).  The  work  herein  completes  this  revision,  and  examines  all  the  known 
fossil  syncarids.  In  addition  to  reestablishing  as  valid  some  old  generic  names,  4  new 
species  are  described,  and  3  new  genera  are  recognized.  The  artificial  familial  arrange- 
ment of  Brooks  (1962(3)  is  essentially  discarded  and  a  new  classification  of  the  Paleozoic 
families  is  put  forth,  one  which  is  felt  to  be  more  natural. 


Abbreviations 

Prefixes  of  catalog  numbers  for  various  institutions  are  as  follows: 

AM  Museum  d'Histoire  Naturelle,  Autun,  France 

B  Museum  d'Histoire  Naturelle,  Paris,  France 

BS  Bayerisches  Staatssamlungen  fur  Palaontologie  und  historisches  Geologic, 

Munich,  West  Germany 

CGH  Narodni  Museum,  Prague,  Czechoslovakia 

F  Australian  Museum,  Sydney,  New  South  Wales 

GSE  Institute  of  Geological  Sciences,  Edinburgh,  Scotland 

GSL  Institute  of  Geological  Sciences,  Leeds,  England 

I,  In  British  Museum  (Natural  History),  London,  England 

ISGS  Illinois  State  Geological  Survey,  Urbana,  Illinois 

Jk  Museum  fur  Naturkunde  (Janensch  Catalog),  Berlin,  East  Germany 

M,  Me  Narodni  Museum,  Prague,  Czechoslovakia 

NB  Rijks  Geololgische  Dienst,  Heerlen,  The  Netherlands 

NYSM  New  York  State  Museum,  Albany,  New  York 

PE  Field  Museum  of  Natural  History,  Chicago,  Illinois 

PMB  Museum  fiir  Naturkunde  (Paleontologisches  Museum  Catalog),  Berlin,  East 

Germany 

SDSNH  San  Diego  Natural  History  Museum,  San  Diego,  California 

US  University  of  Sydney,  Paleontology  Collection,  Sydney,  New  South  Wales 

USNM  National  Museum  of  Natural  History,  Smithsonian  Institution,  Washington, 

D.C. 

X  University  of  Illinois,  Paleontology  Collection,  Urbana,  Illinois 

YPM  Yale  Peabody  Museum  of  Natural  History,  New  Haven,  Connecticut 


191 


Living  Anaspidacea 

In  1980  I  was  able  to  collect  and  study  several  species  of  living  anaspidaceans  in 
Tasmania  with  the  assistance  of  the  staff  of  the  University  of  Tasmania.  Several  of 
these  observations  have  not  been  recorded  before,  and  offer  some  insights  into  the 
biology  of  the  fossil  syncarids. 

The  most  widely  dispersed  anaspidacean  is  Anaspides  tasmaniae  (a  rather  variable 
taxon  which  will  probably  contain  several  subspecies— R.  Swain,  personal  communi- 
cation) which  occur  in  streams,  lakes,  and  caves  widely  scattered  about  the  island  of 
Tasmania.  The  animals  are  in  constant  motion,  somewhat  less  so  in  the  wild  than  in 
the  laboratory.  They  tend  to  engage  in  a  constant  and  random  patrol  of  their  pools. 
They  seem  to  ignore  each  other,  and  in  the  laboratory  they  climb  over  each  other  in 
the  course  of  their  wanderings  like  any  other  obstacle  in  their  path.  The  exopods 
constantly  vibrate  anteriorly  to  posteriorly,  moving  the  epipodites  in  the  process.  The 
annulate  pleopods  are  directed  ventro-laterally,  and  push  the  body  along  the  bottom 
in  metachronal  rhythm  with  the  thoracic  endopods.  While  collecting  Anaspides,  some 
inadvertently  fell  out  of  the  dip  net  onto  the  ground,  whereupon  they  righted  themselves 
and  commenced  to  explore  their  terrestrial  environs  with  ease.  Richardson  {personal 
communication)  relates  that  occasionally,  albeit  rarely,  they  are  naturally  encountered 
out  of  their  pools  on  land.  This  species  does  not  swim  at  all  well.  When  startled  they 
will  execute  a  single  caridoid  flexure  that  propels  them  up  into  the  water  column. 
However,  they  then  drift  passively  until  gravity  returns  them  to  the  bottom.  This 
probably  accounts  for  their  inability  to  survive  in  areas  where  European  sport  fish  have 
been  introduced,  and  also  makes  it  extremely  easy  to  collect  them.  Overall,  A.  tasmaniae 
is  a  very  alert  animal.  The  flagellae  of  both  sets  of  antennae  orient  in  different  directions 
and  constantly  sweep  about.  They  are  omnivorous,  prefering  to  scavenge,  and  are  also 
known  to  pick  up  large  sand  grains  and  manipulate  them  with  their  mouthparts,  ap- 
parently to  scrape  them  of  organics. 

Paranaspides  lacustris  is  a  smaller  animal  than  A.  tasmaniae,  and  exhibits  some 
distinctly  different  behavior.  Their  pleopods  also  function  like  those  o{ Anaspides  when 
they  walk  on  the  bottom.  However,  Paranaspides  seems  to  be  much  more  versatile  in 
its  locomotion.  When  startled  they  may  execute  the  single  caridoid  flexure  already 
mentioned,  or  they  may  dart  away  in  some  direction  parallel  to  the  bottom,  or  they 
may  lay  quite  still.  When  they  do  enter  the  water  column  they  are  capable  of  swimming 
quite  well.  Paranaspides  was  observed  by  me  to  swim  for  hours  near  the  surface  in  a 
small  thermos-container.  In  the  laboratory,  they  were  generally  less  active  animals  than 
Anaspides.  Although  they  have  a  flexure  point  in  the  abdomen  to  facilitate  the  caridoid 
reaction,  when  they  rest  in  their  habitat  on  the  bottom  ooze  or  on  the  water  plants 
they  prefer,  they  reflex  and  hold  the  tailfan  dorsad  off  the  substrate. 

Allanaspides  is  the  smallest  of  the  Anaspididae.  A.  helionomus  is  collected  from 
yabbie  (crayfish)  burrows.  Unlike  the  2  species  above,  A.  helionomus  does  not  beat  its 
exopods  in  a  simple  to-and-fro  pattern,  but  seems  to  rotate  them.  The  exact  manner 
is  not  clear,  but  the  resultant  current  sucks  water  under  the  head  of  the  animal  and 
back  towards  the  tail.  Small,  young  individuals  do  not  beat  their  exopods  when  at  rest; 
only  the  adult  animals  do.  The  anterior  thoracopods  are  oriented  anteriorly  under  the 
head  while  at  rest;  in  combination  with  the  described  current  they  may  function  in 
filter  feeding.  Also,  unlike  Anaspides  and  Paranaspides,  Allanaspides  helionomus  uses 
its  pleopods  in  a  somewhat  different  manner.  The  uniramous  annulate  abdominal 
appendages  are  held  rigid  and  each  pair  is  oriented  in  a  different  direction.  When  they 
assist  in  walking  they  push  off  the  bottom  like  oars  using  just  their  tips.  Walking 
in  A.  helionomus  is  best  described  as  a  "scurrying  crawl,"  occurring  in  intermittent 
bursts  of  activity.  When  at  rest,  the  first  2  pleopods  vibrate  vigorously  to  aerate  the 
fleshy  thoracic  epipodites.  Allanaspides  swims  very  well,  and  was  observed  to  do  so 
even  upside  down.  The  animals  may  be  detritovores  since  in  the  laboratory  they  were 
observed  to  fondle  fecal  pellets  with  their  mouthparts.  The  burrows  these  animals  live 
in  occur  in  grass  marshes  on  surfaces  of  gentle  slope,  aflTording  a  modicum  of  drainage 
and  no  long-standing  water. 


192 


Micraspides  calmani  possess  a  very  flexible  body,  easily  achieving  flexion  dorsally 
and  ventrally  as  well  as  considerable  lateral  bend,  and  is  the  most  infaunal  of  any  of 
the  species  observed  by  me  in  Tasmania.  It  also  lives  in  pools  and  yabbie  burrows  in 
grass  swamps,  but  seems  to  better  tolerate  conditions  with  poorer  drainage  than  do 
species  of  Allanaspides.  Micraspides  moves  with  bursts  of  intermittent  scurrVing,  rem- 
iniscent of  that  seen  in  some  centipedes.  The  annulate  pleopods  are  held  somewhat 
stiffly,  and  operate  only  within  the  metachronal  sequence  of  all  the  limbs,  thus  differing 
only  in  form  and  not  function  from  the  thoracopods.  When  not  moving  they  do  not 
move  any  of  the  appendage  parts.  No  caridoid  escape  reaction  could  be  elicited  from 
Micraspides;  when  startled  or  prodded  the  animals  would  take  evasive  action  by  turning 
laterally  or  flexing  ventrally,  eventually  to  change  their  direction  of  movement  1 80°. 
They  are  thus  ideally  adapted  to  climbing  in,  around,  over,  under,  and  through  obstacles 
in  the  vegetation-choked,  muddy  habitats  they  prefer. 

Several  aspects  of  the  above  have  direct  bearing  on  interpretation  of  the  fossils. 
The  annulate  pleopods  seen  on  the  living  forms  are  noted  as  one  of  the  most  versatile 
and  important  aspects  of  the  anatomy  of  these  creatures;  serving  to  achieve  walking, 
swimming,  and  ventilation  of  epipodites.  They  also  form,  for  the  most  part,  a  functional 
continuum  with  the  thoracopods  (MacMillan  et  al.  1981).  This  has  great  bearing  on 
the  Paleozoic  fossils,  which  were  once  mistakenly  thought  (Brooks  1962/?)  to  all  have 
flap-like  pleopods  and  thoracic  exopods.  The  functional  system  for  pleopods  seen  in 
the  living  forms  is  quite  distinct  from  what  might  be  postulated  for  those  few  paleo- 
caridaceans  with  biramous  flap-like  pleopods,  which  would  appear  to  have  been  capable 
of  only  one  action,  a  to-and-fro  vibration  on  the  ventral  side  of  the  abdomen.  Such 
limbs  would  serve  in  swimming,  but  have  little  or  no  effect  on  walking  on  the  bottom 
or  producing  ventilatory  currents  over  the  thoracic  epipodites.  The  latter  would  have 
to  be  achieved  by  the  vibration  of  the  flap-like  thoracic  exopods.  In  turn,  the  exopods, 
because  of  their  form  and  consequent  limitation  of  movement,  could  not  serve  to 
generate  potential  filtering  currents  around  the  body  as  do  the  rotatory  exopod  move- 
ments of  a  form  like  Allanaspides. 

Palaeocaridaceans  such  as  the  acanthotelsonids,  or  in  part  the  palaeocaridids, 
probably  exhibited  a  functional  system  not  unlike  that  seen  in  other  eumalacostracan 
groups,  such  as  mysidaceans,  euphausiaceans,  and  natant  decapods,  wherein  the  pleo- 
pods are  the  sole  or  primary  organs  of  swimming  in  the  adult  stage.  The  system  seen 
in  the  anaspidaceans,  the  squillitids,  and  in  part  the  palaeocaridids  would  then  appear 
to  possibly  represent  a  functional  advance  in  which  the  entire  trunk  appendage  series 
is  capable  of  acting  as  a  coordinated  unit.  In  this  respect,  it  seems  to  have  been  a 
successful  enough  arrangement  of  parts  to  have  perhaps  evolved  at  least  twice  within 
the  syncarids:  once  in  the  palaeocaridid/squillitid  line  and  again  in  the  anaspidaceans. 

Some  interesting  questions  arise  for  which,  at  the  present,  there  are  no  obvious 
answers.  In  those  living  eumalacostracans  for  which  the  use  of  pleopods  for  swimming 
is  well  developed  there  has  evolved  an  excellent  caridoid  escape  reaction.  Does  the 
existence  of  a  similar  anatomical  system  in  some  of  the  Palaeocaridacea  mean  that 
they  too  may  have  had  a  well-developed  caridoid  escape  reaction,  in  contrast  to  the 
rather  inefficient  single-flexure  behavior  seen  in  living  anaspidaceans?  Dahl  (1983) 
suggests  that  the  caridoid  escape  reaction  of  eumalacostracans  was  independently  evolved 
in  mysidaceans  and  natant  eucarids.  Does  its  possible  existence  in  some  palaeocari- 
daceans mean  there  was  a  third  independent  evolution  of  this  behavior,  or  is  its  possible 
presence  in  the  syncarids  an  argument  for  the  caridoid  escape  reaction  being  considered 
as  a  derived  character  applicable  to  all  eumalacostracans  (Hessler  1983)  and  which  has 
merely  been  repeatedly  lost?  Do  these  functional  considerations  tell  us  anything  about 
character  polarities  (see  next  section)  within  syncarids?  If  the  integrated  system  with 
annulate  appendage  parts  represents  a  functional  advance  within  the  syncarid  line,  this 
might  indicate  that  the  purely  flap-like  structures  are  primitive  and  that  animals  which 
possess  them  are  closer  to  the  stem-group.  Thus  bathynellaceans  and  many  of  the 
palaeocaridaceans  might  be  considered  more  primitive  than  anaspidaceans.  On  the 
other  hand,  if  annulate  limb  parts  and  a  functionally  integrated  trunk  might  represent 


193 


.^^ 


# 

^ 


i 


/ 


^^ 


,^ 


4^ 


Figure  1 .  Classical  arrangement  of  the  superorders  of  Eumalacostraca  sensu  stricto.  Derived  characters  are: 
1)  caridoid  escape  reaction  (and  its  associated  features  of  abdominal  specializations,  see  Hessler  1983),  2) 
antennal  scale  of  a  single  joint,  3)  loss  of  a  carapace,  4)  carapace  fused  to  thoracomeres,  and  5)  oostegite 
brood  pouch. 


an  advance  for  syncarids,  another  scenario  is  possible.  There  is  a  tendency  for  pae- 
domorphosis  in  syncarids  (reduced  or  absent  posterior  limbs,  small  body  size,  free  first 
thoracomere).  Flap-like  limb  parts  and  a  restriction  of  the  pleopods  to  a  swimming 
behavior  may  represent  a  retention  of  "larval"  features,  and  thus  provide  further 
evidence  for  structural  and  behavioral  paedomorphosis  with  palaeocaridaceans  and 
bathynellaceans  being  the  more  derived  groups. 

Higher  Taxonomy  And  Phylogeny 

Problems  arise  in  attempting  to  assess  relationships  of  taxa  within  the  syncarids. 
However,  these  are  no  more  difficult  than  the  problems  associated  with  attempting  to 
assess  the  position  of  syncarids  in  relation  to  other  eumalacostracans.  What  are  the 
unique  characters  which  define  a  taxon  Syncarida?  In  the  classic  scheme  of  Caiman 
(Fig.  1)  Eumalacostraca  sensu  stricto  are  principally  characterized  by  their  caridoid 
escape  reaction  (1)  and  1 -jointed  antennal  scale  (2).  The  syncarids  are  a  sister  group  of 
peracarids  and  eucarids,  defined  by  a  derived  feature  (3),  loss  of  the  carapace  (a  condition 
paralleled  by  a  similar  loss  in  the  line  leading  to  amphipods  and  isopods).  No  opposing 
shared  derived  characters,  however,  join  eucarids.  with  their  carapace  fused  to  the 
thoracomeres  (4),  and  the  peracarids,  with  their  oostegite  brood  pouch  (5). 

Schram  (1981)  and  Watling  (1981,  1983)  have  taken  up  the  problems  engendered 
by  the  Caiman  system  and  have  offered  differing  solutions  to  those  difficulties.  The 
Watling  model  has  difficulties  in  providing  shared  derived  characters  at  the  higher 
taxonomic  levels,  and  won't  be  dealt  with  further  here.  The  system  proposed  by  Schram 
(1981)  had  the  syncarids  as  a  sister  group  to  the  isopods  and  amphipods.  A  difficulty 
with  that  scheme  is  that  it  left  no  derived  features  to  define  the  syncarids.  A  subsequent 
cladistic  analysis  (Schram,  in  press)  utilizing  31  characters  and  a  Wagner  78  program. 


194 


4artices  in 
thoracic  endopods 

5   articles  in 
thoracic  endopods 

first  thoracomere 
free 

Bathynellacea 

Pa  aeocaridacea 

first  thoracomere 
fused  to  cephalon 

Anaspidacea 

Figure  2.     Baupldne  one  can  recognize  with  the  Syncarida.  The  combination  of  the  first  thoracomere  fused 
to  the  cephalon  with  4-segmented  thoracopodal  endopods  was  apparently  never  realized. 


while  confirming  taxa  based  on  Baupldne  derived  from  consideration  of  only  3  char- 
acters (Schram  1981),  does  not  second  the  linking  of  syncarids  with  amphipods  and 
isopods.  This  more  recent  analysis,  however,  does  generally  indicate  that  Syncarida  is 
a  Gilmour-natural  taxon,  and  also  reveals  that  syncarids  probably  are  very  primitive 
animals. 

Indeed,  several  aspects  of  the  biology  of  living  forms  would  reinforce  this  conclu- 
sion. Although  the  living  anaspidaceans  have  a  caridoid  escape  reaction,  it  is  imperfectly 
developed.  It  typically  consists  of  a  single  flick  of  the  abdomen  that  projects  the  animal 
up  into  the  water  column,  after  which  the  animal  either  passively  floats  or  slowly  swims 
back  to  the  bottom.  This  is  in  contrast  to  the  strong  caridoid  escape  reaction  seen  in 
the  mysidaceans  and  eucarids.  In  addition,  syncarids  lay  their  eggs  free  and  gastrulate 
by  involution  into  a  blastocoel— both  primitive  features.  However,  the  development 
within  the  egg  proceeds  to  hatching  at  a  rather  advanced,  free-living  stage  (early  zoeal 
larval  type  in  bathynellaceans,  or  miniature  version  of  the  adult  in  anaspids),  which 
are  generally  considered  derived  features. 

Syncarids  are  thus  generally  considered  to  be  a  monophyletic  taxon.  However, 
discerning  possible  phylogenetic  relationships  within  the  group  poses  problems.  One 
could  recognize  3  basic  morphotypes  or  Baupldne  (Fig.  2)  within  the  syncarids:  a  free 
first  thoracomere  and  less  than  5  segments  in  the  thoracopodal  endopods  (Bathynel- 
lacea), a  free  first  thoracomere  and  5-segment  thoracic  endopods  (Palaeocaridacea), 
and  a  first  thoracomere  fused  to  the  cephalon  and  5  segments  in  the  thoracic  endopods 
(Anaspidacea).  The  characters  used  here  are  the  "traditional"  ones  used  for  decades  to 
sort  major  syncarid  groups. 

Problems  arise  when  one  is  forced  to  choose  between  delineating  syncarids  pri- 
marily on  the  basis  of  endopodal  segment  numbers  or  on  the  basis  of  degree  of  fusion 
of  the  first  thoracomere  into  the  cephalon.  Different  cladograms  and  classifications  of 
the  syncarids  also  result  based  on  whether  palaeocaridaceans  are  to  be  perceived  as  a 
monophyletic  or  paraphyletic  group. 

The  traditional  approach  (Fig.  3)  (Brooks  1969,  Schminke  1975)  essentially  dis- 
criminates between  the  bathynellaceans  with  4-segmented  thoracic  endopods  (1)  and 
palaeocaridaceans  and  anaspidaceans  which  have  5-segmented  thoracic  endopods.  In 
addition,  bathynellaceans  have  the  incisor  process  of  the  mandible  fused  to  the  tooth 
row  (2),  and  the  eighth  thoracopod  of  males  modified  for  copulation  (3).  Character  (2) 
is  difficult  to  assess  in  the  palaeocaridaceans,  since  the  mandibles  generally  are  not 
preserved  well  enough  to  be  able  to  evaluate  whether  they  are  truly  primitive  in  form 
(the  well-preserved  massive  mandibles  seen  in  Palaeocaris,  with  its  distinct  incisor 
process,  may  or  may  not  be  indicative  of  all  palaeocaridaceans).  The  palaeocaridacean- 


195 


Figure  3.  A  "traditional"  presentation  of  relationships  within  Syncarida.  Derived  characters  are:  1 )  4-segment 
thoracic  endopods,  2)  incisor  process  fused  to  tooth  row,  3)  eighth  thoracopod  copulatory,  4)  furcae  lacking, 
5)  first  thoracopod  typically  modified,  6)  "precoxae"  lacking,  7)  paragnaths  lacking,  8)  diagonal  spine  row 
on  uropodal  protopod,  9)  first  thoracopod  reduced,  10)  first  thoracomere  fused  to  cephalon,  11)  eighth 
thoracopod  not  parallel  to  anterior  thoracopods,  1 2)  greatly  shortened  maxillipedal  palp,  1 3)  first  thoracomere 
generally  reduced  in  size,  14)  reduction  and/or  lack  of  pleopods,  15)  pleotelson,  16)  anterior  thoracopods 
rapacious,  17)  annulate  pleopods,  18)  massive  mandibles,  19)  annulate  thoracic  exopods,  20)  large,  setose 
uropod  protopod,  21)  rostrum  separated,  22)  maxillule  palp  lacking,  23)  maxillulary  palp  hook-like.  24) 
mandibular  incisor  process  lacking,  25)  endite  lobe  on  first  thoracopod,  26)  maxilla  proximal  endite  lacking 
spine,  27)  anus  terminal,  28)  mandibular  palp  lacking. 


anaspidacean  line  is  delineated  by  several  synapomorphies:  lack  of  caudal  furcae  (4), 
a  first  thoracopod  typically  modified  in  some  manner  (5),  and  lack  of  thoracopodal 
precoxae  (6).  This  last  character  is  difficult  to  assess,  since  whether  the  presence  of  this 
feature  in  some  bathynellaceans  is  really  a  true  precoxal  leg  joint  or  just  an  articulating 
ring  on  the  body  is  not  clear. 

Within  the  bathynellaceans  the  Parabathynellidae  are  characterized  by  the  lack  of 
paragnaths  (7),  while  the  Bathynellidae  are  marked  by  a  uropodal  protopod  with  a 
diagonal  row  of  spines  (8). 


196 

The  palaeocaridaceans  are  characterized  by  a  derived  feature  that  is  difficuU  to 
evaluate.  Schminke  (1975)  was  the  first  to  point  out  that  the  first  thoracopod  is  ap- 
parently reduced  in  size  as  well  as  number  of  joints  in  the  endopod  (9).  While  this  is 
clearly  true  for  palaeocarids,  acanthotelsonids  and  some  squillitids,  the  form  in  mini- 
carids  is  incompletely  known.  In  the  latter  family  the  first  thoracopod  is  large,  but  it 
is  unclear  if  there  is  a  complete  array  of  5  segments  in  the  endopod.  However,  in  the 
squillitid  genus  Nectotelson  there  is  some  evidence  that  indicates  the  carpus  of  the  first 
thoracopod  is  apparently  not  as  large  as  on  thoracopods  2  through  8.  Generally,  in  the 
former  3  families,  besides  the  greatly  shortened  maxillipedal  palp  (12)  the  first  thorac- 
omere  is  reduced  to  some  degree  (13). 

Minicarididae  are  very  small  animals  which  appear  to  have  a  reduced  number  of 
pleopods  (14).  Admittedly  this  observation  could  be  due  to  vagaries  of  preservation; 
however,  the  rest  of  the  body  and  appendages  of  these  creatures  are  preserved  well 
enough  to  discern  all  pertinent  features  of  their  structure.  The  possible  presence  of  a 
pleotelson  among  these  genera  ( 1 5)  might  eventually  serve  to  further  define  this  group. 

The  Acanthotelsonidae  are  clearly  delineated  by  the  specialized,  rapacious  form 
of  their  anterior  thoracopods  (16).  Within  that  family,  Uronectes  has  only  the  second 
thoracic  appendages  so  modified,  while  Acanthotelson  and  Palaeosyncaris  have  the 
second  and  third  so  specialized.  The  styloid  telson  and  uropods  delineate  the  species 
of  Acanthotelson,  whereas  a  robustly  spinescent  telson  and  lateral  margin  of  the  uro- 
podal  exopod  characterize  Palaeosyncaris. 

The  palaeocarids  and  squillitids  are  characterized  by  annulate  pleopods  (17).  The 
Palaeocarididae,  though  for  the  most  part  rather  generalized,  appear  to  be  distinguished 
by  rather  massive  mandibles  ( 1 8).  The  Squillitidae  are  unified  by  their  acquisition  of 
annulate  thoracic  exopods  (19).  Squillites  has  uniramous  pleopods,  Nectotelson  and 
Praeanaspides  have  biramous  pleopods,  and  Praenaspides  has  a  distinctive  rectangular 
and  laterally  spinose  telson. 

Schminke  (1975)  has  clearly  analyzed  the  distribution  of  characters  within  the 
Anaspidacea,  and  these  are  repeated  here  only  for  completeness.  Anaspidaceans  are 
united  in  possessing  a  first  thoracomere  completely  fused  to  the  cephalon  (10)  and  the 
eighth  thoracopod  offset  at  an  angle  and  not  parallel  to  the  other  thoracopods  (11).  The 
anaspids  and  koonungids  have  a  large  and  setose  uropodal  protopod  (20),  while  psam- 
maspids  and  stygocarids  have  a  separated  rostrum  (21)  and  lack  a  palp  on  the  maxillules 
(22).  Anaspididae  are  distinguished  by  having  the  maxillulary  palp  reduced  to  a  hook- 
like spine  (23),  and  the  Koonungidae  lack  an  incisor  process  on  the  mandible  (24)  and 
have  an  endite  lobe  on  the  first  thoracopod  (25).  The  Psammaspididae  have  no  spine 
on  the  proximal  endite  of  the  maxillae  (26),  while  the  Stygocarididae  have  a  terminal 
anus  (27)  and  lack  a  mandibular  palp  (28). 

A  classification  of  Syncarida  produced  from  the  above  analysis  is  similar  to  that 
usually  encountered  for  the  group,  except  that  a  more  natural  array  of  palaeocaridacean 
families  is  established  than  that  used  by  Brooks  (1962a,  b,  1969). 

Order  Syncarida  Packard,  1885 

Suborder  Bathynellacea  Chappuis,  1915 

Family  Bathynellidae  Chappuis,  1915 

Family  Parabathynellidae  Noodt,  1965 
Suborder  Palaeocaridacea  Brooks,  1962 

Family  Minicarididae,  nov. 

Family  Palaeocarididae  Meek  &  Worthen,  1865 

Family  Squillitidae  Schram  &  Schram,  1974 

Family  Acanthotelsonidae  Meek  &  Worthen,  1865 
Suborder  Anaspidacea  Caiman,  1904 

Family  Anaspididae  Thompson,  1894 

Family  Koonungidae  Sayce,  1908 

Family  Psammaspididae  Schminke,  1974 

Family  Stygocarididae  Noodt,  1963 


197 


Figure  4.  An  alternative  presentation  of  relationships  within  Syncarida.  Derived  characters  are:  1)  first 
thoracomere  reduced  or  fused  to  cephalon.  2)  incisor  process  fused  to  tooth  row,  3)  furcae  lacking,  4)  first 
thoracopod  modified,  5)  reduced  and/or  absent  pleopods,  6)  annulate  thoracic  exopods  and  pleopods,  7) 
4-segmented  thoracic  endopods,  8)  reduced  and/or  absent  pleopods,  9)  eighth  thoracopod  copulatory,  10) 
diagonal  spine  row  on  uropodal  protopod,  1 1)  paragnaths  lacking,  12)  first  thoracomere  fused  to  cephalon, 
13)  eighth  thoracopod  not  parallel  to  anterior  thoracopods,  14)  greatly  reduced  maxillipedal  palp,  15)  large, 
setose  uropodal  protopod,  16)  rostrum  separate,  17)  maxillulary  palp  lacking,  18)  maxillulary  palp  hook- 
like, 19)  mandibular  incisor  process  lacking,  20)  endite  lobe  on  first  thoracopod,  21)  maxilla  proximal  endite 
lacking  spines,  22)  anus  terminal,  23)  mandibular  palp  lacking,  24)  anterior  thoracopods  rapacious,  25) 
annulate  pleopods,  26)  massive  mandibles,  27)  annulate  thoracic  exopods. 


An  alternative  analysis  of  cladistic  relationships  (Fig.  4)  can  be  performed  for 
syncarids  with  initial  assumptions  somewhat  different  from  those  of  the  traditional 
system  presented  above.  Rather  than  make  the  first  dichotomy  one  based  essentially 
on  numbers  of  segments  in  thoracic  endopods,  one  could  distinguish  between  syncarids 
with  no  modification  of  the  first  thoracomere  and  those  with  a  first  thoracomere  mod- 
ified in  some  way  (1).  The  former  line  includes  the  bathynellaceans  and  might  be  further 
characterized  by  mandibles  (2)  with  an  incisor  process  fused  to  the  tooth  row  (as  noted 
above,  a  character  impossible  to  verify  as  yet  on  all  the  fossils).  This  line  divides  into 
a  branch  leading  to  some  fossil  families  which  lack  furcae  (3)  and  which  may  have  a 
first  thoracopod  modified  from  the  form  seen  in  the  second  through  eighth  thoracopods 
(4).  Within  this  branch,  the  Minicaridadae  apparently  lack  or  have  a  reduced  number 
of  pleopods  (5)  while  the  Squillitidae  (in  part,  including  Nectotelson  and  Squillites) 
have  annulate  thoracic  exopods  and  pleopods  (6). 

The  branch  leading  to  the  bathynellaceans  of  course  delineates  the  2  families  on 
the  basis  of  the  bathynellid's  possession  of  a  diagonal  row  of  spines  on  the  uropodal 
protopod  (10)  and  by  the  parabathynellid's  lack  of  a  paragnath  (11). 

The  Anaspidacea  fuse  the  first  thoracomere  into  the  cephalon  (12)  and  have  the 
eighth  thoracopod  offset  from  the  seventh  thoracopod  (13).  The  opposing  branch  with 
its  reduced  but  free  thoracomere  and  parallel  seventh  and  eighth  thoracopods  possess 


198 


a  derived  reduction  in  the  first  thoracopodal  endopod  being  very  short  (14).  The  an- 
aspid/koonungid  line,  as  above,  have  large  and  setose  uropodal  protopods  (15);  and 
within  that  the  Anaspididae  have  a  hook-like  maxillulary  palp  (18),  while  Koonungidae 
lack  a  mandibular  incisor  process  (19)  and  have  gnathobasic  lobes  on  the  first  thora- 
copod  (20).  The  psammaspid/stygocarid  line  has  a  separate  rostrum  (16)  and  lacks  a 
maxillulary  palp  (17).  The  Psammaspididae  lack  spines  on  the  proximal  endites  of  the 
maxillae  (21),  while  the  Stygocarididae  have  a  terminal  anus  (22)  and  lack  a  mandibular 
palp  (23). 

The  Acanthotelsonidae  have  rapacious  anterior  thoracopods  (24),  while  the  pa- 
laeocarid/squillitid  group  has  annulate  pleopods  (25).  The  Palaeocarididae  have  mas- 
sive mandibles  (26),  and  Praeanaspides  (a  squillitid)  has  annulate  thoracic  exopods 
(27). 

This  analysis  could  yield  a  classification  somewhat  different  than  the  traditional, 
in  that  essentially  4  groups  can  be  recognized.  Brooks'  order  Palaeocaridacea  emerges 
as  a  polyphyletic  taxon,  thus  the  major  groups  might  best  be  recognized  as  superfamilies. 

Order  Syncarida 

Superfamily  Bathynelloidea 

Family  Bathynellidae 

Family  Parabathynellidae 
Superfamily  Minicaridoidea 

Family  Minicarididae 

Family  Squillitidae  (in  part) 
Superfamily  Palaeocaridoidea 

Family  Palaeocarididae 

Family  Acanthotelsonidae 

?  Praeanaspides 
Superfamily  Anaspidoidea 

Family  Anaspididae 

Family  Koonungidae 

Family  Psammaspidae 

Family  Stygocarididae 

In  many  respects,  this  second  arrangement  is  an  unsatisfactory  system.  The  dif- 
ferences encapsulated  in  these  2  classifications  of  the  syncarids  arises  from  a  dichotomy 
involved  in  outgroup  comparison  of  the  "palaeocaridaceans,"  and  in  both  schemes  it 
involves  establishing  derived  characters  in  the  Paleozoic  families.  The  problem  could 
be  expressed  as  a  simple  dilemma  derived  from  initial  consideration  of  the  syncarid 
morphotypes:  which  is  more  important,  the  fusion  of  first  thoracomere  into  the  ceph- 
alon,  or  the  loss  of  a  joint  in  the  thoracopodal  endopods?  The  initial  "weighting" 
determines  the  course  of  the  subsequent  analysis. 

In  the  traditional  scheme  (Fig.  3)  28  apomorphies  are  used  to  define  the  10  families 
of  Syncarida;  in  the  alternative  scheme  (Fig.  4)  there  are  27  apomorphies  to  separate 
1 1  "family"  level  taxa.  The  traditional  scheme  thus  seems  to  possess  slightly  more 
information  value.  It  also  more  clearly  justifies  its  initial  dichotomy  (based  on  joint 
number  in  thoracic  endopods)  with  the  greatest  number  of  congruent  features.  For  these 
last  reasons,  as  well  as  the  fact  that  the  alternative  scheme  requires  too  many  uncertain 
judgments  at  this  time  involving  the  poorly  known  minicaridoids,  I  have  opted  in  the 
systematic  section  of  this  monograph  to  retain  the  traditional  classification  of  syncarids 
into  suborders.  However,  more  detailed  and  exacting  knowledge  of  the  minicarids  and 
Nectotelson  someday  may  allow  a  more  reasoned  selection  to  be  made  between  these 
two  systems. 

Nevertheless,  the  problem  of  analyzing  syncarid  phylogeny  is  not  so  easily  disposed 
of  (as  if  the  above  taxonomic  dilemma  were  easy).  Both  of  the  schemes  above  take  for 
granted  essentially  the  same  position  in  regard  to  polarity  of  a  basic  character  in  the 
group,  viz,  that  the  primitive  condition  is  one  in  which  the  first  thoracomere  is  free 


199 


and  large,  and  that  increasing  specialization  is  achieved  as  this  segment  is  reduced  and 
eventually  fused  into  the  cephalon.  Is  this  necessarily  the  case? 

Schminke  (1981)  presents  a  well-documented  series  of  arguments  for  the  progenetic 
paedomorphic  derivation  of  bathynellaceans  from  some  ancestral  syncarid  condition 
in  which  an  adult  animal,  presumably  of  a  palaeocarid  or  anaspid  form,  had  a  long 
larval  sequence.  If  we  extend  Schminke's  arguments,  might  we  not  question  whether 
the  anatomical  stages  seen  in  bathynellaceans  (large  and  free  first  thoracomere,  flap- 
like thoracic  exopods,  reduced  number  of  thoracic  endopodal  segments,  reduced  or 
missing  pleopods,  and  caudal  furcae)  are  really  primitive?  Rather,  might  we  not  consider 
these  features  to  be  actually  derived  by  the  agency  of  progenesis  from  some  ancestral 
adult  in  which  none  of  these  "larval"  features  were  expressed.  In  such  an  interpretation, 
the  most  "primitive"  adult  state  would  be  one  in  which  the  first  thoracomere  is  fused 
to  the  cephalon,  and  that  the  manifestations  of  successive  degrees  of  freedom  of  the 
first  thoracomere  are  increasingly  derived. 

This  assumption  involving  a  reversed  polarity  would  lead  in  turn  to  a  rather 
controversial  cladistic  analysis  (Fig.  5).  A  bathynellacean/palaeocaridacean  line  would 
be  characterized  by  the  presence  of  a  free  first  thoracomere  (1),  the  anaspidacean  line 
by  the  eighth  thoracopod  being  offset  from  and  not  parallel  to  the  seventh  (2). 

The  characterizations  within  the  Anaspidacea  follow  those  already  given  above: 
large,  setose  uropodal  protopods  in  anaspids  and  koonungids  (3),  a  separate  rostrum 
(4)  and  no  maxillulary  palp  (5)  in  psammaspids  and  stygocarids,  a  hook-like  palp  on 
the  maxillule  (6)  in  anaspids,  lack  of  a  mandibular  incisor  process  (7)  and  endite  lobes 
on  the  first  thoracomere  (8)  in  koonungids,  lack  of  spines  on  the  proximal  endite  of 
the  maxilla  (9)  in  psammaspids,  and  a  terminal  anus  (10)  and  lack  of  a  mandibular 
palp  (11)  in  the  stygocarids. 

The  Bathynellacea  share  several  advanced  characters:  a  4-segment  thoracic  en- 
dopod  (12),  fusion  of  the  incisor  process  to  the  tooth  row  in  the  mandible  (13),  a 
copulatory  eighth  thoracopod  in  the  male  (14),  and  a  first  thoracomere  as  large  as  any 
succeeding  thoracomere  (15).  The  Palaeocaridacea  lack  furcae  (16). 

The  bathynellids  have  a  diagonal  row  of  spines  on  the  uropodal  protopod  (17), 
while  the  parabathynellids  lack  paragnaths  (18). 

Within  the  palaeocaridaceans,  the  minicarid/squillitid  (in  part)  line  convergently 
develops  the  enlarged  first  thoracomere  (19),  while  the  other  families  have  a  reduced 
endopod  on  the  first  thoracopod  (20).  The  minicarids  apparently  have  reduced  or  absent 
pleopods  (21),  while  the  squillitids  have  annulate  thoracic  exopods  and  annulate  pleo- 
pods. The  acanthotelsonids  have  rapacious  anterior  thoracopods  (23),  while  the  pa- 
laeocand/ Praeanaspides  line  has  annulate  pleopods  (24).  The  palaeocarids  have  mas- 
sive mandibles  (25)  and  the  Praeanaspides  also  possess  annulate  thoracic  exopods  (26). 

The  above  scheme  in  comparision  with  the  traditional  and  alternative  schemes 
discussed  earlier  unfortunately  uses  only  26  apomorphies  to  define  its  end  points  and 
has  a  rather  high  number  of  convergent  characters.  Note,  however,  that  the  taxonomy 
which  results  from  this  cladogram  is  similar  to  that  of  the  traditional  classification, 
except  that  the  Anaspidacea  in  this  latter  scheme  are  felt  to  be  closest  to  the  primitive 
condition,  and  the  Squillitidae  sensu  stricto  are  separated  from  the  genus  Praeanaspides. 

Still  another  analysis  alternative  to  the  above  is  possible  (Fig.  6)  also  involving 
the  reversed  polarity,  but  utilizing  only  25  apomorphies.  The  anaspidoid  line  is  as 
above.  The  bathynelloid/minicaridoid  line  has  an  enlarged  first  thoracomere  ( 1 2),  while 
the  palaeocaridoid  line  has  a  reduced  endopod  on  the  first  thoracopod.  The  bathynelloid 
hne  has  the  4-segment  endopod  (14),  incisor  process  fusion  (15),  and  copulatory  eighth 
thoracopod  (16)  noted  before;  and  the  minicaridoid  line  has  a  problematic  apomorphy 
difficult  to  assess  because  of  preservation,  i.e.,  the  first  thoracopod  large  but  possibly 
not  structurally  identical  to  the  second  and  following  thoracopods  (17).  The  family 
apomorphies  [Bathynellidae  (18),  Parabathynellidae  (19),  Minicaridiae  (20),  Squilliti- 
dae (in  part)  (21),  Acanthotelsonidae  (22),  Palaeocarididae  (23.  24),  and  Praeanaspides 
(23,  25)]  are  all  those  noted  in  the  schemes  already  discussed,  especially  that  in  Figure  5. 


200 


■c'^ 

"O 


^     /     ■/      ^     /     v^ 

/   /    #   /    /    -^ 


Figure  5.  Relationships  within  Syncarida  involving  reversed  polarity  with  loss  of  fusion  of  the  first  tho- 
racomere  with  the  cephalon  due  to  paedomorphosis.  Derived  characters  are:  1)  first  thoracomere  free,  2) 
eighth  thoracomere  not  parallel  to  anterior  thoracopods,  3)  large,  setose  uropodal  protopod,  4)  rostrum 
separate,  5)  maxillulary  palp  lacking,  6)  maxillulary  palp  hook-like,  7)  mandibular  incisor  process  lacking, 
8)  endite  lobe  on  first  thoracomere,  9)  maxillary  proximal  endite  lacking  spines,  10)  anus  terminal,  11) 
mandibular  palp  lacking,  12)  4-segment  thoracic  endopod,  13)  incisor  process  fused  to  tooth  row,  14)  eighth 
thoracopod  copulatory,  15)  first  thoracomere  subequal  to  any  succeeding  thoracomeres,  16)  furcae  lacking, 
1 7)  diagonal  spine  row  on  uropod  protopod,  1 8)  paragnaths  lacking,  1 9)  first  thoracomere  subequal  to  any 
succeeding  thoracomeres,  20)  greatly  reduced  maxillipedal  palp,  21)  reduced  and/or  absent  pleopods,  22) 
annulate  thoracic  exopods  and  pleopods,  23)  anterior  thoracopods  rapacious,  24)  annulate  pleopods,  25) 
massive  mandibles,  26)  annulate  thoracic  exopods. 


This  last  cladogram  coiresponds  to  the  alternative  classification  given  above,  except 
that  the  superfamily  Anaspidoidea  is  now  presented  as  the  closest  to  a  primitive  con- 
dition for  Syncarida  as  a  whole. 

What  can  we  conclude  about  syncarid  evolution?  We  should  be  cognizant  of  some 
level  of  uncertainty  as  to  just  how  these  taxa  are  related  to  each  other.  However,  a 
more  definitive  resolution  of  the  problem  must  await  better  and  more  detailed  infor- 
mation about  the  Paleozoic  syncarids.  Characters  which  delineate  the  living  and  fossil 
families  are  not  equivalent  in  the  sense  that  the  living  families  are  separated  on  the 
basis  of  details  of  mouthparts,  whereas  the  fossil  families  are  largely  resolved  on  the 
basis  of  gross  form  of  trunk  appendages.  Ideally,  more  mouthpart  data  for  Palaeocar- 
ideacea  could  have  allowed  a  more  complete  data  matrix  than  that  used  here  to  be 
analyzed  with  a  Wagner  78  program.  The  resultant  rigor  could  have  mathematically 
determined  parsimony  and  homoplasy.  However,  phylogenetic  trees  and  taxonomies 
are  pragmatic  instruments  (Charig  1982,  Schram  1983),  and  the  lack  of  any  data  that 
we  would  like  to  have  should  not  be  an  excuse  for  not  attempting  to  organize  that 


201 


Figure  6.  An  alternative  presentation  of  relationships  within  Syncarida  involving  reversed  polarity  with 
loss  of  fusion  of  the  first  thoracomere  with  the  cephalon  due  to  paedomorphosis.  Derived  characters  are:  1 ) 
first  thoracomere  free,  2)  eighth  thoracomere  not  parallel  to  anterior  thoracopods,  3)  large,  setose  uropodal 
protopod,  4)  rostrum  separate,  5)  maxillulary  palp  lacking,  6)  maxillulary  palp  hook-like,  7)  mandibular 
incisor  process  lacking,  8)  endite  lobe  on  first  thoracomere,  9)  maxillary  proximal  endite  lacking  spines,  10) 
anus  terminal,  1 1)  mandibular  palp  lacking,  12)  first  thoracomere,  13)  reduced  endopod  on  first  thoracopod, 
14)  4-segment  thoracic  endopod,  15)  incisor  process  fused  to  tooth  row,  16)  eighth  thoracopod  copulatory, 
17)  ?  form  of  first  thoracopod,  18)  diagonal  spine  row  on  uropod  protopod,  19)  paragnaths  lacking,  20) 
reduced  and/or  absent  pleopods,  21)  annulate  thoracic  exopods  and  pleopods,  22)  anterior  thoracopods 
rapacious,  23)  annulate  pleopods,  24)  massive  mandibles,  25)  annulate  thoracic  exopods. 


information  which  we  do  have.  We  should  simply  recognize  the  limits  of  the  information 
at  hand,  and  be  aware  of  its  effect  on  the  level  of  uncertainty  engendered  in  our  present 
understanding  of  syncarid  evolution.  Nevertheless,  I  would  hope  that  the  organization 
of  the  fossil  Syncarida  used  here  is  more  adequate  than  anything  that  we  have  had 
heretofore. 

Systematics 

The  system  of  annotated  synonymy,  summarized  by  Matthews  (1973),  is  used  in 
this  section  of  the  monograph.  This  should  facilitate  use  and  evaluation  of  my  systematic 
decisions  by  any  future  workers. 


Order  SYNCARIDA  Packard,  1885 

Suborder  PALAEOCARIDACEA  Brooks,  1962 

Family  MINICARIDIDAE  nov. 

Diagnosis.  — Thoracic  exopods  unisegmental.  Pleopods  unisegmental,  if  present. 
First  thoracomere  large,  not  reduced  nor  fused  to  cephalon. 

Type  genus.— Minicaris  Schram,  1979. 

Remarks.  — The  distinctive  nature  of  the  first  thoracomere  and  the  unisegmental 
or  flap-like  form  of  the  pleopods  when  present  clearly  warrants  separate  family  status. 


202 


Figure  7.     A)  Reconstruction  of  Minicahs  brandi.  scale  2  mm  (redrawn  from  Schram  1979a);  B)  tailfan 
to  same  scale  as  body;  C)  dorsal  view  of  right  antenna  slightly  enlarged. 


In  addition,  the  small  size,  possible  absence  of  posterior  pleopods,  the  possible  presence 
of  a  pleotelson,  and  the  early  age  (Lower  Carboniferous)  is  of  interest  with  regard  to  a 
parallelism  to,  or  a  possible  origin  of,  the  Bathynellacea  {see  Higher  Taxonomy  and 
Phylogeny  section). 

Genus  MINICARIS  Schram,  1979a 

Diagnosis.  — VtdunclQS  of  antennules  and  antennae  subequal.  At  least  first  pleopod 
present  and  well  developed.  Uropods  narrow  and  blade-like.  (?)Pleotelson. 
Type  species.— Minicaris  brandi  Schram,  \919a. 

Minicaris  brandi  Schram,  \919a 
Fig.  7 

v.*  1979a  Minicaris  brandi  Schram,  p.  109,  figs.  52  &  53. 
1979b  Minicaris  brandi  Schram.  Schram,  p.  170,  table  2. 

1981  Minicaris  brandi  Schram.  Schram,  p.  131,  table  2,  fig.  6D. 
7952  Minicaris  brandi  Schram.  Wood,  p.  577. 

1982  Minicaris  brandi  Schram.  Schram,  p.  122,  fig.  8A. 

Diagnosis. —Since  there  is  but  one  species,  the  diagnosis  is  the  same  as  that  of  the 
genus. 

Holotype. —GSE  13056.  Long  Livingston  Borehole  no.  25,  West  Lothian,  Scotland. 
1071-1151  foot  section,  below  Pumpherstone  Shell  Bed,  Lower  Oil  Shale  Group,  Di- 
nantian,  Lower  Carboniferous. 

Other  locality.— Questionably  reported  from  along  Manse  Bum,  Bearsden,  near 
Glasgow,  Scotland,  in  shales  equivalent  to  the  Top  Hosie  Limestone,  lowermost  Na- 
murian  (Wood  1982). 

i)€'5'cn/)//oA2.— Antennular  peduncle  3  joints,  proximal-most  joint  one-half  total 
length  of  peduncle,  distal  2  joints  progressively  shorter.  Antennal  protopod  distal  joint 
twice  the  proximal,  scaphocerite  oval  with  distal  tip  pointed  and  setose,  proximal  2 
flagellar  joints  peduncular.  Thoracomeres  with  rounded  pleura,  posterior  comers  acute. 
Thoracopodal  exopods  narrow.  All  thoracopods  appear  equal,  ischium  long,  merus  and 
carpus  short,  propodus  moderate,  dactylus  short.  Abdominal  pleura  rounded.  If  not  a 
tme  pleotelson,  telson  not  sharply  sutured  from  sixth  pleomere.  Telson  spade-like, 
setose.  Uropods  blade-like,  setose,  possibly  with  diaeresis. 

/^emar/:^.  — Reexamination  in  1980  of  the  holotype,  and  still  only  good  specimen 
of  this  species,  confirmed  all  the  pertinent  points  of  the  anatomy  above.  The  lack  of 


203 


Figure  8.     A)  Reconstruction  of  Erythrogaulos  carrizoensis,  scale  2  mm;  B)  tailfan  to  same  scale  as  body. 


all  but  the  first  pleopod  in  the  abdominal  series  is  still  not  completely  understood.  It 
is  possible  the  more  posterior  pleopods  were  not  preserved,  but  it  is  also  possible  that 
they  were  never  there  to  begin  with.  The  small  size  (8  mm)  and  general  form  of  the 
animal  might  indicate  a  possibility  of  paedomorphosis  in  the  evolution  of  this  taxon, 
since  so  many  of  the  living  small  syncarids  do  not  develop  complete  series  of  pleopods 
nor  completely  separate  the  telson  from  the  last  pleomere.  Only  more  and  better  material 
can  allow  us  to  choose  between  these  alternatives. 

Genus  ERYTHROGAULOS  new  genus 

Diagnosis.  — 'PosXtnov  comers  of  pleomere  pleura  serrate.  Telson  distally  spinose. 
Uropodal  exopod  distally  spinose. 

Type  species.— Erythrogaulos  carrizoensis  new  species. 

Etymology.— A.  reference  to  the  stratigraphic  horizon,  Red  Tanks  Member,  Madera 
Formation,  Lower  Permian. 


Erythrogaulos  carrizoensis  new  species 
Fig.  8;  Plate  1,  figs.  A  &  B 

Diagnosis.  SincQ  there  is  but  one  species,  the  diagnosis  is  the  same  as  that  of  the 
genus. 

//o/o/yp^.-SDSNH  25141  (Plate  1,  figs.  A  «fe  B).  Carrizo  Arroyo,  Lucero  Mts., 
southeastern  Valencia  County,  New  Mexico.  Upper  Red  Tanks  Member,  Madera  For- 
mation, Wolfcampian,  Lower  Permian.  (Collected  by  Dr.  Jarmilla  Kukalova-Peck, 
Carlton  University,  Ottawa,  Ontario,  Canada.) 

Etymology.  —  MXqt  the  type  locality  in  Carrizo  Arroyo. 

Description.  — Body  small.  Thoracomeres  subequal,  pleura  apparently  rounded, 
except  eighth  which  appears  posteriorly  serrate.  Pleomeres  subequal,  posterior  comers 
of  at  least  first  3  pleura  serrate.  Telson  rectangular,  developed  distally  with  2  sets  of 
tooth-like  spines,  medial  distal  set  larger  than  lateral  proximal  pair.  Uropodal  rami, 
blade-like  and  subequal,  slightly  longer  than  telson,  exopod  with  distal  tooth-like  spines 
on  lateral  margin  just  anterior  of  where  diaeresis  might  be,  endopod  margins  finely 
setose. 

Remarks.  — Ox\\y  one  specimen  allows  any  inference  to  be  made  concerning  the 
anatomy  of  the  animal.  Two  other  specimens  (SDSNH  29140)  appear  to  preserve  only 
part  of  the  trunk  segment  series.  The  observed  thoracopods  are  of  such  a  diaphanous 
preservation  as  to  preclude  any  more  concrete  conclusions  about  them  other  than  that 
they  seem  to  be  equally  developed  back  to  and  including  the  eighth  pair.  None  of  the 


204 


Plate  1 

Figures  A  &  B.  Erythrogaulos  carhzoensis  new  species,  holotype,  SDSNH  25 1 4 1 ;  A)  whole  specimen,  x  7; 
B)  closeup  of  posterior  abdomen  and  tailfan,  note  spines  on  posterior  of  pleura  (p),  spines  on  distal  telson 
(t)  and  lateral  margin  of  uropodal  exopod  (e),  and  setose  margins  of  uropodal  rami  (arrows),  x  19.6. 

Figures  C-E.  Acanthotelson  stimpsoni  Meek  and  Worthen,  1865;  C  &  D)  latex  peels  of  holotype,  X  346, 
X  1.7;  E)  syntype  oi  A.  event  Meek  and  Worthen,  1868,  ISGS  3066  (made  a  junior  synonym  of  A.  stimpsoni, 
by  Packard  1886),  x  1.2. 


205 


anterior  limbs  seemed  specialized  in  any  way,  though  these  had  to  be  partly  destroyed 
in  preparation  in  order  to  fully  reveal  the  cephalon.  No  traces  of  pleopods  were  noted, 
and  this,  combined  with  the  fact  that  thoracopods  are  clearly  detectable  and  with  the 
general  small  size  of  the  body,  might  suggest  the  possibility,  as  with  Minicaris  brandi, 
that  pleopods  were  either  greatly  reduced  or  not  present  on  this  species. 

The  distinctively  serrate  pleura  and  spinose  telson  warrant  separate  generic  status 
for  this  species  from  its  nearest  relative,  Minicaris  brandi  of  the  Lower  Carboniferous. 

The  associated  biota  in  the  Red  Tanks  Member  includes:  numerous  plants  dom- 
inated by  the  gymnosperm  genera  Walchia  and  Cordaites,  but  also  including  Callipteris, 
sphenopsids,  and  lycopsids;  a  most  diverse  array  of  uniramians  including  insects  and 
myriapods;  the  eurypterid  Adelophthalmus  luceroensis;  ostracodes;  brachiopods;  and 
spirorbid  worms.  Kues  and  Kietzke  (1981)  interpret  the  paleoecology  of  the  Carrizo 
Arroyo  fauna  as  representing  a  fresh  to  brackish  water  habitat  on  a  delta  plain.  The 
extreme  delicacy  of  the  preservation  from  this  locality  also  indicates  quick  burial  under 
anoxic  conditions  with  little  postdepositional  disturbance. 

Family  ACANTHOTELSONIDAE  Meek  and  Worthen,  1865 

Diagnosis.— Thoracic  exopods  unisegmental  and  flap-like.  Anterior  thoracopods 
raptorial.  Pleopods  biramous  and  flap-like. 

Type  genus.— Acanthotelson  Meek  &  Worthen,  1865. 

Remarks.  — brooks  (1962a,  b)  chose  to  place  the  genera  Acanthotelson  and  Uro- 
nectes  in  separate  families  based  on  the  degree  of  raptorial  development  expressed  in 
the  anterior  thoracopods.  Although  this  is  an  important  character,  it  is  best  utilized 
for  distinction  at  the  generic  level.  The  unisegmental,  flap-like  nature  of  the  thoracic 
exopods  and  rami  of  the  pleopods  in  comparison  to  other  palaeocaridacean  families 
herein  recognized  warrants  uniting  all  species  with  raptorial  thoracopods  into  a  single 
family. 

Genus  ACANTHOTELSON  Meek  and  Worthen,  1865 

Diagnosis.  — First  thoracopod  markedly  reduced.  Second  and  third  thoracopods 
raptorial.  Telson  and  uropods  styliform. 

Type  species.— Acanthotelson  stimpsoni  Meek  and  Worthen,  1865. 

Acanthotelson  stimpsoni  Meek  and  Worthen,  1865 
Fig.  9;  Plate  1,  figs.  C-E,  Plate  2,  fig.  A 

v.*  1865  Acanthotelson  stimpsoni  Meek  and  Worthen,  p.  47. 

V.  1866  Acanthotelson  stimpsoni  Meek  and  Worthen.  Meek  and  Worthen,  p.  401,  pi.  32,  figs.  6,  6a-f. 

1868a  Acanthotelson  eveni  Meek  and  Worthen.  Meek  and  Worthen,  p.  27. 

V.  18686  Acanthotelson  stimpsoni  Meek  and  Worthen.  Meek  and  Worthen,  p.  549,  2  figs. 

V.  1868Z?  Acanthotelson  eveni  Meek  and  Worthen.  Meek  and  Worthen,  p.  551.  4  figs. 

1880  Acanthotelson  stimpsoni  Meek  and  Worthen.  Brocchi,  p.  10,  pi.  1,  fig.  11. 

1884  Acanthotelson  stimpsoni  Meek  and  Worthen.  White,  p.  176,  pi.  37,  fig.  4-5. 

1 184  Acanthotelson  eveni  Meek  and  Worthen.  White,  p.  177,  pi.  38,  figs.  4-7. 

V.  1886<2  Acanthotelson  stimpsoni  Meek  and  Worthen.  Packard,  p.  123,  pi.  1,  figs.  1-3,  pi.  2,  figs.  1-3. 

1886a  Acanthotelson  eveni  Meek  and  Worthen.  Packard,  p.  125. 

1890  Eileticus  anthracinus  Scudder,  p.  420,  pi.  38,  fig.  5. 

1890  Eilecticus  aequalis  Scudder,  p.  421,  pi.  38,  figs.  6-9. 

1896  Acanthotelson  stimpsoni  Meek  and  Worthen.  Caiman,  p.  799,  pi.  2,  fig.  16. 

1901  Acanthotelson  species  Fritsch,  p.  74,  fig.  398. 

1909  Acanthotelson  stimpsoni  Meek  and  Worthen.  Smith,  p.  575,  fig.  62. 

1911a  Acanthotelson  stimpsoni  Meek  and  Worthen.  Caiman,  p.  159. 

191  la  Acanthotelson  eveni  Meek  and  Worthen,  Caiman,  p.  159. 

1916  Acanthotelson  stimpsoni  Meek  and  Worthen.  Cockerell,  p.  234. 

1916  Acanthotelson  stimpsoni  Meek  and  Worthen.  Vanhoffen,  p.  146,  fig.  12. 

1916  Acanthotelson  eveni  Meek  and  Worthen.  Vanhoffen,  p.  148. 

1916  Acanthotelson  species.  Vanhoffen,  p.  148,  fig.  14. 

1919  Acanthotelson  species.  Pruvost,  p.  85. 

7927  Acanthotelson  stimpsoni  Meek  and  Worthen.  Chappuis,  p.  605. 


206 


Figure  9.  A)  Reconstruction  of  Acanthotelson  stimpsoni,  scale  5  mm  (modified  from  Brooks  19626);  B) 
lailfan  to  same  scale  as  body;  dorsal  views  of  C)  right  antennule  and  D)  antenna;  E)  posterior  thoracopod. 
Appendages  slightly  enlarged. 


1931  Acanthotelson  event  Meek  and  Worthen.  Van  Straelen,  p.  1 1. 

1931  Acanthotelson  stimpsoni  Meek  and  Worthen.  Van  Straelen,  p.  12. 

1959  Acanthotelson  event  Meek  and  Worthen.  Slewing,  p.  2. 

1959  Acanthotelson  stimpsoni  Meek  and  Worthen.  Siewing,  p.  3. 

1962a  Acanthotelson  stimpsoni  Meek  and  Worthen.  Brooks,  p.  236. 

V.  19626  Acanthotelson  stimpsoni  Meek  and  Worthen.  Brooks,  p.  230,  pis.  55-59;  Text-pl.  10,  11a. 

7965  Acanthotelson  species  Noodt,  p.  83. 

1969  Acanthotelson  stimpsoni  Meek  and  Worthen.  Brooks,  p.  R355,  figs.  165-2,  171. 


1969a  Acanthotelson  stimpsoni 
1969b  Acanthotelson  stimpsoni 
1976a  Acanthotelson  stimpsoni 
1976b  Acanthotelson  stimpsoni 
1979a  Acanthotelson  stimpsoni 
1979b  Acanthotelson  stimpsoni 
1981a  Acanthotelson  stimpsoni 
1981b  Acanthotelson  stimpsoni 


Meek  and  Worthen.  Schram,  p.  219,  Table  1. 

Meek  and  Worthen.  Schram,  p.  201. 

Meek  and  Worthen.  Schram,  p.  21. 

Meek  and  Worthen.  Schram,  p.  411. 

Meek  &  Worthen.  Schram,  p.  28,  Table  1. 

Meek  and  Worthen.  Schram,  p.  167,  fig.  1,  Table  2. 

Meek  and  Worthen.  Schram,  p.  131,  text-fig.  5b,  Table  2. 

Meek  and  Worthen.  Schram,  p.  9,  fig.  in  text. 


7952  Acanthotelson  event  Meek  and  Worthen.  Kent,  p.  15. 

Diagnosis. —  Second  joinX  of  antennal  peduncle  shorter  than  first  or  third.  Telson 
styHform,  equal  to  or  slightly  longer  than  uropods. 

Lectotype.  —  X  346  (Plate  1 ,  figs.  C  and  D).  Mazon  Creek  area,  Will  County,  Illinois. 
Francis  Creek  Shale,  Carbondale  Formation  (Westphalian  C),  Pennsylvanian.  (Im- 
properly designated  a  holotype  by  Brooks  \962b.) 

Other  localities.  — {Sqq  Schram  1976a)  Illinois  State  Geol.  Surv.  core  T-4  (816  feet) 
NW  V4,  SW  '/4,  SE  »/4,  sec  25,  T2S,  R14W,  Wabash  County  Ilhnois;  Dykersburg  Shale, 
Carbondale  Formation,  Pennsylvanian.  Sec.  4,  T9S,  RIE,  '/z  mile  west  of  Carterville, 
Williamson  County;  gray  shale  above  #6  (Herrin)  Coal,  Brereton  Cyclothem,  Penn- 
sylvanian. Abandoned  Chieftan  Mine,  7  miles  south  of  Terre  Haute,  Indiana,  east  of 
Highway  41;  Lower  Shelbum  Formation,  Pennsylvanian. 

Description.— CQXi^3\on  with  short  rostrum,  cervical  and  precervical  grooves.  Eyes 
small  and  stalked.  Antennular  peduncle  3-segmented,  proximal  and  distal  joints  large, 
medial  segment  short,  flagella  well  developed  with  inner  branch  shorter  than  outer 
branch.  Antennal  protopod  with  short  proximal  segment  bearing  nephropore  and  long 
distal  segment,  no  scaphocerite,  very  long  flagellum  with  proximal  2  joints  peduncular. 
Antennules  and  antennae  with  setose  inner  peduncular  margins.  Mandible  massive, 
palp  well  developed.  Maxillule  with  3-segment  palp.  Maxilla  with  at  least  proximal 
segment  of  palp  large. 

First  thoracomere  reduced  in  length.  Second  through  fourth  thoracomeres  pro- 
gressively longer  than  first.  Last  4  thoracomeres  subequal,  last  3  thoracomeres  have 


207 

anterior  margins  with  raised  ridge.  Thoracic  pleura  simple.  First  thoracopod  reduced, 
possibly  as  short  maxillipede.  Second  and  third  thoracopods  biramous,  endopods  large, 
spinose,  and  raptorial  in  form.  Five  posterior  thoracopods  of  ambulatory  form,  with 
epipodites,  exopods  of  single  segment  (flap-like),  endopods  with  short  ischium  and 
dactylus,  and  long  merus,  carpus,  and  propodus  joints. 

Pleomeres  similar  in  size  to  posterior  thoracomeres,  first  through  fifth  pleura  with 
posterio-ventral  comers  serrate,  fifth  and  sixth  pleomere  posterior  margins  serrate. 
Sixth  pleomere  not  elongate.  Pleopods  as  biramous  setose  paddles.  Telson  as  long  spike, 
margins  with  alternating  spines  and  setae.  Uropodal  rami  as  blades,  margins  with 
alternating  spines  and  setae.  Spikes  or  uropods  and  telson  reinforced  with  median 
ridges. 

Remarks.— Though  recognized  as  a  distinct  taxon  since  the  time  of  Meek  and 
Worthen  (1865);  a  complete  and  reliable  description  and  accurate  reconstruction  of  .4. 
stimpsoni  was  not  available  until  Brooks  {\962b).  However,  actual  photo  illustrations 
of  the  type  series  of  A.  stimpsoni  have  not  been  prepared  until  now,  except  for  the 
single  exception  of  Brooks  {\962b,  pi.  54,  fig.  4).  Latex  peels  illustrated  here  of  the 
lectotype,  X346  (Plate  1,  figs.  C  and  D),  are  taken  from  the  specimen  which  was  used 
as  the  basis  for  one  of  the  drawings  in  Meek  and  Worthen  (1868/^:549,  fig.  B).  A 
paralectotype,  X3442,  (Plate  2,  fig.  A)  was  the  basis  for  another  drawing  in  Meek  and 
Worthen  (1868^7:549,  fig.  A).  The  tail  on  X3442  was  also  apparently  used  by  Meek 
and  Worthen  as  an  addition  to  augment  their  drawing  (p.  551,  fig.  A)  of  ISGS  3066 
(Plate  1,  fig.  E).  ISGS  3066  is  also  a  syntype  of  another  species,  A.  eveni,  since  syn- 
onymized  (Packard  1886)  with  A.  stimpsoni. 

Heretofore,  understanding  of  what  constitutes  the  genus  Acanthotelson  has  been 
clouded  incredibly  by  the  incorrect  use  of  the  name  Eileticus  Scudder,  1882,  by  Eu- 
ropean workers.  Eileticus  {sensu  stricto)  is  now  generally  conceded  to  be  a  myriapod. 
However,  as  pointed  out  by  Brooks  (1962Z):258),  Scudder  designated  as  a  separate 
taxon,  E.  aequalis,  what  turned  out  to  be  a  poorly  preserved  specimen  of  A.  stimpsoni. 
Some  European  workers  built  upon  this  confusion  and  have  applied  the  name  Eileticus 
to  other  taxa  that  have  turned  out  not  to  be  Acanthotelson.  For  example,  E.  cf.  aequalis 
of  Pruvost  ( 1 9 1 9)  is  probably  Pleurocaris,  and  E.  pruvosti  Vandenberghe  ( 1 960)  is  likely 
better  placed  in  Nectotelson.  Each  of  these  cases  is  discussed  in  detail  elsewhere,  under 
the  appropriate  taxon  designation. 

Acanthotelson  kentuckiensis  new  species 

by 

Frederick  R.  Schram  and  Donald  Chesnut* 

*Kentucky  Geological  Survey,  Lexington 

Fig.  10;  Plate  2,  figs.  B-F 

Diagnosis.  — ^QgmQnXs  of  antennular  peduncles  subequal.  Telson  subtriangular  but 
long  and  narrow,  shorter  than  uropods. 

//o/o/y/7£'.-SDSNH  23722  (Plate  2,  fig.  B).  Black  Oak  Coal,  Inc.  strip  mine,  near 
Silverville,  McCreary  County,  Kentucky,  2  miles  north  of  Tennessee  state  line;  84°26'30" 
N,  36°38'42"  W.  Black  fissile  shale  above  River  Gem  Rider  Coal,  Lower  Breathitt 
Formation,  Middle  Pennsylvanian. 

Other  material. -SDSNH  23723  (Plate  2,  fig.  E),  23724,  23725. 

Descriptions.  — Aniennules  well  developed,  with  3  subequal  segments  (Plate  2,  fig. 
F)  composing  the  peduncle,  about  equal  in  size  to  peduncle  of  antennae.  Antennal 
peduncular  segments  apparently  short,  no  scaphocerite  noted. 

All  pleomeres  about  equal  in  length,  last  3  with  paired  longitudinal  dorsal  ridges, 
dorsal  posterior  margin  of  sixth  pleomere  concave.  Telson  (Plate  2,  figs.  C  &  D)  tri- 
angular in  outline,  narrow,  with  dorsal  median  ridge,  margins  furrowed  and  setose, 
distal  setae  more  strongly  developed.  Uropodal  rami  styliform,  each  with  reinforcing 
rib  flanked  by  slight  furrows,  exopodal  margins  with  strong  setae  (especially  laterally), 
endopodal  margins  finely  setose. 


208 


,—■ -^ 

"l    1  ^' ~T~T~Ttt 

-' 

'  /  /   ;  '  '  '  '  > 

B 


Figure  10.     A)  Diagrammatic  rendition  of  what  is  currently  known  about  the  form  of  Acanthotelson  ken- 
tuckiensis.  scale  5  mm;  B)  tailfan  to  same  scale  as  body. 


Remarks.— T\iQ  most  complete  specimen  (SDSNH  23722)  lacks  a  thorax  and  all 
other  specimens  are  of  tailfans  only.  However,  though  anatomical  information  about 
this  species  is  minimal,  the  distinctive  nature  of  the  tailfan,  especially  the  styliform 
uropods,  is  so  different  from  that  seen  in  A.  stimpsoni  as  to  require,  pending  some 
future  evidence  to  the  contrary  concerning  thoracopods,  a  separate  species  for  this 
material  within  the  genus  Acanthotelson. 

The  shorter  and  broader  telson  of  ^.  kentuckiensis  is  more  primitive  than  the  long 
styliform  tail  oi  A.  stimpsoni.  Apparently,  the  styliform  expression  is  allometric  since 
the  smallest  specimen  of  ^.  kentuckiensis,  SDSNH  23723,  has  the  broadest  and  shortest 
telson  (Table  1)  while  the  larger  specimens  are  narrower  and  longer.  A.  stimpsoni,  by 
contrast,  is  a  generally  larger  and  more  robust  species  than  A.  kentuckiensis,  and  has 
a  very  long,  styliform  telson. 

The  biota  associated  with  A.  kentuckiensis  indicates  a  fresh  to  brackish  water  facies. 
In  addition  to  occasional  fish  scales  and  teeth,  abundant  remains  were  collected  of  the 
pelecypod  Anthraconaia,  and  fossils  of  the  plant  Calamites  were  common.  These  ob- 
servations on  the  biota  are  reinforced  by  the  nature  of  the  black,  fissile,  canneloid  shale 
in  which  the  animals  are  found.  The  fossils  occur  at  the  base  of  a  generally  coarsening 
sequence  of  shales  and  sandstones,  beginning  with  the  carbonaceous  shales  with  abun- 
dant fossils,  and  grading  into  an  increasing  arenaceous  sequence  with  interbedded  gray 
and  black  shales.  These  beds  overlay  another  coarsening  sequence  with  the  River  Gem 


Table  1.  Measurements  in  mm  on  material  of  Acanthotelson  kentuckiensis.  Comparative  data  on  telson 
measurements  included  for  two  representative  specimens  of  A.  stimpsoni.  see  text  for  discussion.  *  Ho- 
lotype. 


Specimen 


Length  Max. 

Length  A,  Length        Length       width       Ration 

cephalon    peduncle  a^  telson       telson        tl:tw 


Length      Length 

uropodal  uropodal 

exopod    endopod 


A.  kentuckiensis 

♦SDSNH  23722 

SDSNH  23723 

SDSNH  23724 

SDSNH  23725 

A.  stimpsoni 

SDSNH  17454 
SDSNH  5210 


-3.0 


1.8 


0.9 

3.2 

1.0 

3.2 

4.0 

3.3 

0.5 

1.5 

0.8 

1.9 

2.4 

1.3 

3.0 

1.1 

2.7 

0.8 

1.1 

0.9 

2.9 

3.5 

7.5 

1.7 

4.4 

7.1 

1.6 

4.4 

209 


. ■ttj '.-,•"..  ."j<  >•■ 


© 


®      ^^ 


_  J- 


Plate  2 

Figure  A.    Acanlhotelson  stimpsoni  Meek  and  Woilhen,  1865,  paralectotype  X  344-Z,  x  1.3. 

Figures  B-F.  Acanthotelson  kentuckiensisnev/  species;  B-D,  F)  holotype  SDSNH  23722;  B)  whole  specimen 
displaying  cephalon  and  abdomen,  thorax  missing,  x3.9;  C,  D)  closeup  of  tailfan  counterparts,  x7.8;  F) 
closeup  of  cephalon,  note  antennular  peduncle  with  3  subequal  segments  (arrow).  E)  SDSNH  23723,  note 
relatively  wider  telson  (t)  in  relation  to  length  than  that  seen  in  C  or  D,  x6. 


Rider  Coal  at  the  base,  on  top  of  which  are  a  siltstone  and  shale  grading  into  a  heavy 
burrowed  argillaceous  sandstone. 

Genus  URONECTES  Bronn,  1850  (=GAMPSONYCHUS  Burmeister,  1855) 

Diagnosis.  — No  rostrum.  First  thoracomere  moderately  reduced.  Second  thora- 
copod  raptorial.  Telson  and  uropods  broad  and  rounded,  uropods  with  straight  diaere- 
sis, broad  tailfan  formed  from  overlapping  elements. 


210 


Figure  11.     A)  Reconstruction  of  Uronectes  fimbriatus,  scale  5  mm  (modified  and  corrected  from  Brooks, 
1962Z?);  B)  tailfan  to  same  scale  as  body;  C)  dorsal  view  of  right  antenna  slightly  enlarged. 


Type  species.  —  Gampsonyx  fimbriatus  Jordan,  1 847. 


Uronectes  fimbriatus  (Jordan),  1847 
Fig.  1 1 

*1847  Gampsonyx  fimbriatus  Jordan,  p.  89,  pi.  2. 
1848  Gampsonyx  fimbriatus  Jordan.  Bronn,  p.  575. 
7550  Gampsonyx  fimbriatus  Jordan.  Bronn,  p.  575. 
1850  Uronectes  fimbriatus  (Jordan).  Bronn,  p.  575. 

1854  Gampsonyx  fimbriatus  Jordan.  Jordan  &  von  Meyer,  p.  1,  pi.  2. 

1855  Gampsonychus  fimbriatus  (JoTdan).  Burmeister,  p.  191,  pi.  10,  figs.  12-14. 

1856  Uronectes  fimbriatus  (Jordan).  Roemer,  p.  202. 

1856  Gamsonychus  fimbriatus  (Jordan).  Roemer,  p.  202,  p.  672. 

1873  Gampsonyx  fimbriatus  Jordan.  Feistmantel,  p.  593,  pi.  18,  figs.  9-11. 

1877  Carcinurus fimbriatus  (Jordan).  Goldenburg,  p,  35,  pi.  2,  figs,  lb,  2-7. 

1880  Gampsonyx  fimbriatus  Jordan.  Brocchi,  p.  10,  pi.  10,  fig.  7. 

1885  Gampsonychus  fimbriatus  (Jordan).  Zittel,  p.  672,  fig.  857  (in  part). 

1886b  Gampsonychus  fimbriatus  (Jordan).  Packard,  p.  130,  fig.  1. 

1896  Gampsonyx  fimbriatus  Jordan.  Caiman,  p.  798,  pi.  2,  fig.  17. 

1900  Gampsonyx  fimbriatus  Jordan.  Eastman  in  Zittel,  p.  659,  fig.  1382. 

1901  Gampsonychus  fimbriatus  (Jordan).  Fritsch,  p.  72,  pi.  159,  text-fig.  377. 

1902  Uronectes  fimbriatus  (Jordan).  Caiman,  p.  66. 

1909  Gampsonyx  fimbriatus  Jordan.  Smith,  p.  568,  fig.  53-55. 
1916  Gampsonychus  fimbriatus  (Jordan).  Vanhoffen,  p.  143,  fig.  7-8. 
1927  Uronectes  fimbriatus  (Jordan).  Haack,  p.  733,  3  figs. 
1927  Uronectes  fimbriatus  (Jordan).  Chappuis,  p.  605. 
1931  Uronectes  fimbriatus  (Jordan).  Van  Straelen,  p.  18. 

1958  Uronectes  fimbriatus  (Jordan).  Malzahn,  p.  355. 

1959  Gampsonychus  fimbriatus  (Jordan).  Slewing,  p.  1. 
1962(2  Uronectes  fimbriatus  (Jordan).  Brooks,  p.  236. 

1962^  Uronectes  fimbriatus  (Jordan).  Brooks,  p.  230,  text-pl.  1  lb. 

1963  Uronectes  species.  Noodt,  p.  82. 

1969  Uronectes  fimbriatus  Jordan.  Brooks,  p.  R355,  figs.  165-3,  173. 

1969a  Uronectes  fimbriatus  Jordan.  Schram,  p.  221,  table  1. 

7972  Uronectes  species  Jordan.  Boy,  p.  47,  fig.  2. 

1974  Uronectes  fimbriatus  Jordan.  Schram  &  Schram,  p.  101. 

7979  Uronectes  fimbriatus  Jordan.  Schram  &  Schram,  p.  170. 

7952  Uronectes  species  Schneider,  et  al.,  p.  75,  fig.  5. 

Diagnosis. —  Sixih  thoracomere  somewhat  enlarged  over  adjacent  segments.  Sixth 
pleomere  long. 

Lectotype.—]k  4a,  b,  from  the  Kramer  Ironworks  of  Lebach,  near  Saarbriicken, 
Saarland,  West  Germany.  Rotliegende,  Lower  Permian. 


211 


Paralectotype.—Jk  5. 

Other  localities.  — Pfeffelbach,  near  Kusel,  Rheinlandpfalz,  West  Germany;  Rot- 
liegende.  Oberhof  (Schweitzerhatte),  near  Zella-Mehlis,  Thiiringia,  East  Germany;  Ob- 
erhofer  Beds,  Lower  Permian. 

Diagnosis.  —Cephalon  with  faint  cervical  groove,  no  rostrum.  Antennule  peduncles 
3-segmented,  proximal  segment  very  long,  distal  2  joints  short,  flagella  moderately 
developed.  Antennal  protopod  with  2  subequal  segments,  scaphocerite  oval,  flagellum 
moderately  long  with  proximal  2  segments  peduncular. 

First  thoracomere  moderately  reduced.  Thoracic  pleura  simple,  with  slight  furrow 
along  margins.  Sixth  thoracomere  somewhat  longer  dorsally  than  others.  Second  thor- 
acopod  large,  spinose,  and  raptorial.  Second  through  eighth  thoracopods  ambulatory, 
more  or  less  subequal,  ischium  very  short,  merus  through  propodus  moderate,  dactylus 
very  small. 

First  5  pleomeres  with  finely  serrate  posterior  margins,  pleura  acuminate  anteriorly 
with  slight  furrow  on  margins.  Sixth  pleomere  elongate.  Telson  rounded,  margins  setose. 
Uropods  flap-like,  margins  setose,  exopod  with  straight  diaeresis  and  reinforced  with 
lateral  thickened  rib. 

Remarks.— The  reconstructions  of  U.  fimbriatus prepared  by  Brooks  (1962^?,  1969) 
generally  reflect  an  accurate  view  of  the  creature,  except  for  the  fact  that  he  mistakenly 
drew  7  abdominal  segments  instead  of  6  (corrected  here  in  Fig.  1 1).  The  description 
in  his  text  indicates  the  proper  number. 

The  classic  Lebach  locality  has  been  the  source  of  U.  fimbriatus  specimens  in 
museums  around  the  world.  The  freshly  collected  material  was  a  black  shale.  The 
characteristic  red  rock  with  white  fossils  developed  only  after  the  specimens  were 
"roasted"  at  the  Kramer  Ironworks,  driving  off"  the  volatile  organics  in  the  shale  and 
fossils,  and  leaving  a  calcitic  residue  behind  on  a  rock  residue  high  in  siderite.  Specimens 
from  other  localities  were  found  by  me  while  searching  various  European  collections. 
The  Staatssamlungen  flir  Palaontologie  in  Munich  has  an  "unroasted"  specimen  from 
the  Rothegende  (BS  1975  I  164)  from  Pfefffelbach,  near  Kusel,  not  too  far  from  Lebach. 
The  Munich  collection  also  has  a  specimen  from  the  Oberhofer  Beds  from  near  Oberhof, 
in  Thiiringia  (BS  1953  XXVIII  21)  in  a  strange  3-dimensional  preservation.  The  Pa- 
laontologisches  Museum  of  the  Museum  fiir  Naturkunde  in  Berlin  also  has  specimens 
(PMB  A.  62-67)  identified  as  U.  fimbriatus  from  Thiiringia  (H.-E.  Gruner,  pers.  comm.), 
as  does  the  San  Diego  Natural  History  Museum  (these  a  gift  of  Dr.  J.  Schneider  of  the 
Bergakademie,  Freiberg). 

Uronectes  kinniensis  Schram  and  Schram,  1979 

Fig.  12 

v.*  1979  Uronectes  kinniensis  Schram  and  Schram,  p.  169,  pi.  1,  text-fig.  1. 
1981a  Uronectes  kinniensis  Scharm  and  Schram.  Schram,  p.  133,  text-fig.  4g. 

Diagnosis.  — YourXh.  thoracomere  moderately  reduced;  eighth  thoracomere  with 
lateral  semicircular  ridges.  Fifth  pleomere  elongate. 

Holotype.-\]^NM  235625.  Kinney  Clay  Pit,  SE  'A,  Sec.  18,  T9N,  R6E,  Bernalillo 
County,  New  Mexico.  Madera  Formation,  Virgilian,  Pennsylvanian. 

Description.— CephaXon  apparently  undecorated,  no  rostrum.  Antennal  protopod 
of  2  subequal  segments,  scaphocerite  subtriangular  and  setose,  at  least  proximal-most 
joint  of  flagellum  peduncular. 

First  and  fourth  thoracomeres  moderately  reduced.  All  thoracomeres  except  eighth 
undecorated,  pleura  simple.  Eighth  thoracomere  with  small,  lateral,  paired,  semicircular 
ridges.  Thoracomeres  3  through  8  subequal  (details  obscure). 

Pleopods  with  finely  serrate  posterior  margins.  Fifth  pleomere  elongate.  Telson 
rectangular,  rounded  distally,  and  apparently  distally  serrate.  Uropods  as  broad  flaps, 
exopod  with  straight  diaeresis,  at  least  endopods  setose. 

Remarks.— The  lack  of  knowledge  about  the  first  and  second  thoracopods  makes 
it  difficult  to  place  this  species  in  Uronectes  without  any  hesitation.  However,  as  orig- 


212 


Figure  12.     A)  Reconstruction  of  Uronectes  kinniensis,  scale  5  mm  (modified  from  Schram  and  Schram 
1979);  B)  tailfan  to  same  scale  as  body;  C)  dorsal  view  of  right  antenna  slightly  enlarged. 


inally  reported  (Schram  and  Schram  1979:170)  the  overall  aspects  of  the  anatomy 
(especially  the  serrate  pleomere  margins,  lack  of  rostrum,  moderately  reduced  first 
thoracomere,  and  straight  diaeresis)  come  closest  to  Uronectes,  and  U.  kinniensis  is 
best  left  within  that  genus  for  the  time  being. 

Genus  PALAEOSYNCARIS  Brooks,  \962b 

Diagnosis.  — First  thoracomere  very  reduced,  second  thoracomere  moderately  re- 
duced. Second  and  third  thoracopods  raptorial.  Telson  oval,  with  spinose  margins. 
Uropodal  exopods  laterally  spinose. 

Type  species.—  Palaeosyncaris  dakotensis  Brooks,  \962b. 

Palaeosyncaris  dakotensis  Brooks,  \962b 
Fig.  13;  Plate  3,  figs.  B-E 

v.*  1962^  Palaeosyncaris  dakotensis  Brooks,  p.  251;  pi.  65,  figs.  3,  4,  pi.  66;  text-pl.  14,  fig.  a. 
1969  Palaeosyncaris  dakotensis  Brooks.  Brooks,  p.  R355,  figs.  169-2,  170-2. 


Figure  13.     A)  Reconstruction  of  Palaeosyncaris  dakotensis,  scale  5  mm  (corrected  from  Brooks  1962^); 
B)  tailfan  to  same  scale  as  body;  C)  dorsal  view  of  right  antenna  slightly  enlarged. 


213 


Plate  3 

Figure  A.     Palaeosyncaris  micra  new  species,  holotype,  PE  2496,  x4.1. 

Figures  B-E.  Palaeosyncaris  dakotensis  Brooks,  1 962*;  B-D)  holotype,  USNM  1 43409;  B)  whole  specimen. 
x2.4;  C)  closeup  of  lailfan,  x7;  D)  closeup  of  anterior  thoracopods,  note  the  inflated  meri  on  thoracopods 
(2)  and  (3)  and  antennal  scale  (arrow),  x  8;  E)  counterpart  of  holotype,  Univ.  of  North  Dakota  collection, 
note  inflated  meri  on  thoracopods  (2)  and  (3)  and  antennal  scale  (arrow),  x5.5. 


1969a  Palaeosyncaris  dakotensis  Brooks.  Schram.  p.  216.  table  1. 
1974  Palaeosyncaris  dakotensis  Brooks.  Schram  and  Schram,  p.  95. 

Diagnosis.— A\\  segments  with  transversely  striate  decoration.  Abdominal  pleura 
with  posterior  margins  serrate. 

Holotype.  — \]SNM  14309  (Plate  3,  figs.  B-D)  (counterpart,  unnumbered,  in  col- 
lection of  University  of  North  Dakota,  Plate  3,  fig.  E).  Borehole  Casimer  Duletski  No. 


214 


1,81 70-8 1 80  feet  NW  'A,  NW  'A,  Sec.  1 6,  T 1 39N,  R99W,  Stark  County,  North  Dakota. 
Heath  Shale,  Upper  Mississippian. 

Description.  — Eyes  small,  eye  stalk  long.  Antennules  with  3-segment  peduncle, 
proximal  joint  equal  to  distal  2  joints,  flagella  well  developed.  Antennae  with  small, 
oval,  finely  setose  scaphocerite,  proximal  2  joints  of  flagellum  peduncular  witfi  median 
margins  setose,  flagellum  well  developed  (but  of  undetermined  length). 

First  thoracomere  greatly  reduced,  second  thoracomere  moderately  reduced;  sec- 
ond and  all  other  somites  (Plate  3,  fig.  B)  with  transverse  striae,  pleura  rounded,  and 
ventral  margins  with  furrow.  First  thoracopod  apparently  reduced;  second  through 
eighth  thoracopods  robust,  merus  somewhat  longer  than  other  subequal  joints,  merus 
on  second  and  third  thoracopods  (Plate  3,  figs.  D-E)  inflated  (possibly  spinescent), 
dactylus  long  and  pointed,  second  and  third  thoracopods  raptorial.  Eighth  thoracic 
pleuron  posteriorly  extended  and  margin  serrate.  All  tergites  with  marginal  furrows, 
especially  prominent  on  pleura. 

Abdominal  pleura  anteriorly  somewhat  rounded,  posteriorly  pointed  with  mar- 
gins serrate.  Last  pleomere  somewhat  elongate.  Telson  long,  oval,  and  marginally  spi- 
nose,  terminal  median  spines  reduced  in  comparison  to  adjacent  members  of  series. 
Uropodal  exopod  laterally  spinose,  endopod  margins  finely  setose  (Plate  3,  fig.  C). 

Remarks.  — Though  obviously  well-preserved  thoracic  epipodites  and  exopods  ap- 
pear not  to  have  been  present  on  the  type  specimen,  there  is  some  indication  on  the 
coxa  of  the  third  thoracopod  of  USNM  143409  of  a  foramen  for  an  epipodite.  This 
same  appendage  may  also  preserve  part  of  an  exopod  arising  from  the  basis. 

The  thoracopodal  endopods  of  this  species  are  all  strongly  developed.  The  inflated 
meri  on  the  second  and  third  thoracopods  may  well  have  been  capable  of  acting  like 
subchelae  in  opposition  to  the  carpi  on  these  appendages,  which  appear  to  be  proximally 
narrow  and  with  a  rather  disto-posterior  spiniform  crest.  In  this  regard  USNM  143409 
appears  to  have  partially  preserved  the  sockets  of  articulating  spines  on  the  merus  of 
the  second  thoracopod  disto-posteriorly. 

Brooks  (1962)  compared  P.  dakotensis  to  Praeanaspides  praecursor,  mainly  on  the 
basis  of  similarities  of  tergal  ornament.  The  tailfans,  however,  are  now  known  to  be 
quite  different  (Schram  1 979a).  Furthermore,  the  identification  of  raptorial  thoracopods 
on  P.  dakotensis  would  appear  to  ally  this  species  with  members  of  the  Acanthotel- 
sonidae.  Placement  in  this  family  should  not  be  without  query,  however,  since  complete 
knowledge  of  the  thoracic  exopods  and  pleopods  would  be  necessary  before  unques- 
tioned affiliation  could  be  sanctioned.  Pleopods  are  not  preserved  on  the  type  coun- 
terparts. 

The  other  syncarid  of  the  Heath  Shale  is  Squillites  spinosus.  This  latter  species  is 
collected  from  "paper  shale"  outcrops  of  the  Heath  in  central  Montana,  while  the  rock 
of  the  North  Dakota  core  which  contains  P.  dakotensis  is  a  well-indurated,  blocky, 
black  shale.  The  associated  fauna  on  the  core  section  with  P.  dakotensis  is  largely 
composed  of  partially  pyritized  cyzicoid  branchiopods  and  casts  of  indeterminate  os- 
tracodes. 

Palaeosyncaris  micra.  new  species 
Fig.  14;  Plate  3,  fig.  A;  Plate  4,  figs.  A-E 

Diagnosis.  — 'Qody  small.  Segments  mooth,  undecorated.  Abdominal  pleura  not 
serrate.  Telson  with  spinose  setae  increasing  in  size  distally. 

Holotype.  —  PE  2496  (Plate  3,  fig.  A).  Mazon  Creek  area;  Will,  Crundy,  and  Kan- 
kakee Counties,  Illinois.  Francis  Creek  Shale,  Carbondale  Formation  (Westphalian  C- 
D),  Pennsylvanian. 

Description.  — Body  small,  tergites  smooth.  Cephalon  with  short  rostrum.  Anten- 
nules with  3  subequal  joints  in  peduncle.  Antenna  with  small  setose  scaphocerite. 

First  thoracomere  greatly  reduced,  second  moderately  reduced,  third  through  eighth 
subequal  with  anterior  comers  rounded  (Plate  4,  figs.  A,  C).  First  thoracopod  reduced, 
about  one-half  the  length  of  ambulatory  thoracopods.  Second  and  third  thoracopods 


215 


Figure  14.     A)  Reconstruction  of  Palaeosyncaris  micra,  scale  5  mm;  B)  tailfan  to  same  scale  as  body;  C) 
dorsal  view  of  right  antenna  slightly  enlarged. 


robust,  raptorial,  dactyli  with  well-developed  terminal  spines  (Plate  4,  fig.  A).  Third 
through  eighth  thoracopods  ambulatory;  meri  and  carpi  long,  ischia,  propodi,  and 
dactyli  short;  exopods  flap-like  (Plate  4,  fig.  C). 

Abdominal  pleura  anteriorly  and  posteriorly  reduced.  Sixth  pleomere  somewhat 
longer  than  anterior  pleomeres.  Telson  oval  (Plate  4,  fig.  D),  medial  margins  with 
spinose  setae,  with  setae  increasing  in  size  distally.  Uropods  (Plate  4,  figs.  D,  E)  setose, 
setae  of  lateral  margin  of  exopod  spinose,  exopod  reinforced  with  medial  rib  and 
apparently  possessing  an  oval  diaeresis. 

Remarks.  — ThQ  small  size  and  generally  incomplete  preservation  of  these  fossils 
has  resulted  in  their  being  mistakenly  sorted  by  previous  workers  (including  myself) 
as  "small  and  poorly  preserved"  examples  of  the  other  two  Mazon  Creek  syncarids, 
Acanthotelson  stimpsoni  and  Palaeocaris  typus.  We  are  indebted  to  Mr.  Stephen  L. 
May  for  recognizing  these  specimens  as  a  separate  species,  and  bringing  it  to  our 
attention.  Representative  measurements  are  given  in  Table  2. 

The  establishment  of  a  third  Mazon  Creek  syncarid  now  brings  the  crustacean 
assemblage  of  the  brackish  water  biotope  in  the  American  Pennsylvanian  into  accord 
with  that  of  the  European  Carboniferous  (Schram  1981a).  Both  faunas  now  have  a 
pygocephalomorph  associated  with  3  species  of  syncarid  (see  Table  3).  However,  there 
does  not  appear  to  be  a  point-for-point  analogy  between  the  syncarid  species.  The 
American  faunas  have  2  rapacious  acanthotelsonids  and  1  palaeocarid,  whereas  the 
British  fauna  syncarids  are  in  apparently  3  different  families.  Palaeosyncaris  micra 
occurs  in  both  the  Essex  and  Braidwood  faunas  of  Johnson  and  Richardson  (1966). 


Table  2.     Representative  measurements  in  mm  of  species  of  Palaeosyncaris.  *  Holotypes. 


Head 

Thorax 

Abdomen 

Telson 

T, 

T, 

T, 

A. 

P.  dakotensis 

*USNM  143409 

3.5 

10.5 

11.0 

4.0 

0.6 

1.1 

1.3 

2.5 

P.  micra 

*PE  2496 

0.8 

7.8 

7.5 

0.4 

0.5 

0.8 

PE  11670 

-1.8 

8.8 

0.6 

1.0 

1.1 

PE  1268 

7.1 

2.6 

0.8 

1.2 

PE  12174 

1.0 

7.9 

7.0 

0.4 

0.7 

0.9 

1.1 

PE  37912 

1.0 

6.8 

5.5 

0.4 

0.6 

0.8 

PE  37915 

1.3 

8.1 

0.5 

0.7 

1.0 

216 


1 

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V 


Plate  4 

Figures  A-E.  Palaeosyncaris  micra  new  species;  A)  latex  peel  of  holotype,  PE  2496.  closeup  of  anterior 
end,  note  the  raptorial  thoracopods  2  and  3  (arrows),  and  the  reduced  first  thoracomere  (1),  x9.8;  B)  latex 
peel  of  PE  11670,  with  rounded  thoracic  and  abdominal  pleura,  x4.5;  Q  latex  peel  of  PE  37915,  note 
progressively  increasing  lengths  of  anterior  thoracomeres  (1-3)  and  large  flap-like  exopods  (arrows),  x  7;  D) 
latex  peel  of  PE  12168,  showing  spination  on  telson,  x  10.7;  E)  latex  peel  of  PE  12174,  showing  spines  on 
lateral  margin  of  uropodal  exopod  and  possible  diaeresis  (arrow),  setation  of  endopod,  and  all  but  distal- 
most  portions  of  the  telson,  x  7.4. 


217 
Table  3.     Crustaceans  found  in  Late  Carboniferous  brackish  water  habitats  in  North  America  and  Europe. 


Mazon  Creek  Faunas 

Westphahan  British  Coal  Measures 

Pygocephalomorpha 

Acanthotelson  stimpsoni 

Pygocephalus  cooperi 

Syncarida 

Acanthotelson  stimpsoni 
Palaeocaris  typus 
Palaeosyncaris  micra 

Praeanaspides  praecursor 
Palaeocaris  retractata 
Pleurocaris  annulatus 

Family  PALAEOCARIDIDAE  Meek  and  Worthen,  1865 

Diagnosis.— Thovdic'xc  exopods  flap-like,  pleopods  annulate. 

Type  genus.— Palaeocaris  Meek  and  Worthen,  1865. 

Remarks.  — The  genus  Paleocaris  has  long  been  treated  as  a  catchall  taxon  for  every 
incompletely  known  Paleozoic  eumalacostracan  which  has  not  had  any  evident  cara- 
pace. This  has  been  complicated  by  the  fact  that  Palaeocaris  itself  was  incorrectly 
understood,  in  the  sense  of  Brooks  (1962).  Unfortunately,  much  remains  to  be  discov- 
ered about  this  most  important  genus  of  Paleozoic  syncarids;  however,  the  diagnostic 
combination  of  annulate  pleopods  with  flap-like  thoracic  exopods  provides  a  focus 
upon  which  further  work  in  the  group  can  be  based.  It  is  possible  the  genera  Brook- 
syncaris,  Palaeorchestia,  and  Williamocalmania  may  belong  to  this  family,  but  these 
fossils  yet  lack  relevant  information  about  the  appendages  to  allow  definitive  placement 
of  them  into  the  Palaeocarididae. 

Genus  PALAEOCARIS  Meek  and  Worthen,  1865 

Diagnosis.  — M.3.n6\h\QS  massive,  first  thoracomere  greatly  reduced.  Sixth  pleomere 
posterior  margin  deeply  concave.  Uropodal  rami  margins  very  setose,  exopod  with 
pronouced  diaeresis,  exopod  distinctly  longer  than  endopod,  endopod  distinctively 
longer  than  telson.  Telson  oval,  margins  bearing  stout  setae. 

Palaeocaris  typus  Meek  and  Worthen,  1865 

[=Acanthotelson  inaequalis  Meek  and  Worthen,  1865] 

Fig.  15;  Plate  5,  figs.  A-C 

1865  Acanthotelson  inaequalis  Meek  and  Worthen,  p.  48. 
v.*  1865  Palaeocaris  typus  Meek  and  Worthen,  p.  49. 

1866  Acanthotelson  inaequalis  Meek  and  Worthen.  Meek  and  Worthen,  p.  403,  pi.  32,  fig.  7. 
V.  1866  Palaeocaris  typus  Meek  and  Worthen.  Meek  and  Worthen,  p.  405,  pi.  32,  figs.  5,  5a-d. 
1868(3  Palaeocaris  typus  Meek  and  Worthen.  Meek  and  Worthen,  p.  28. 

18686  Palaeocaris  typus  Meek  and  Worthen.  Meek  and  Worthen,  p.  552,  figs.  1,  2. 

1880  Palaeocaris  typus  Meek  and  Worthen.  Brocchi,  p.  9,  pi.  1,  figs.  8-10. 

1884  Palaeocaris  typus  Meek  and  Worthen.  White,  p.  179,  pi.  38,  figs.  1-3. 

V.  18866  Palaeocaris  typus  Meek  and  Worthen.  Packard,  p.  129,  pi.  7,  figs.  1-2. 

1889  Palaeocaris  typus  Meek  and  Worthen.  Packard,  p.  213. 

1896  Palaeocaris  typus  Meek  and  Worthen.  Caiman,  p.  796,  pi.  2,  fig.  15. 

7909  Palaeocaris  typus  Meek  and  Worthen.  Smith,  p.  570.  text-figs.  56-58. 

79/6  Palaeocaris  typus  Meek  and  Worthen.  Vanhoffen,  p.  141,  fig.  5. 

7976  Acanthotelson  inaequalis  Meek  and  Worthen.  Vanhoffen,  p.  147. 

7927  Palaeocaris  typus  Meek  and  Worthen.  Chappuis,  p.  605. 

V?  1957a  Palaeocaris  species.  Copeland,  p.  595;  pi.  6,  fig.  5. 

V?  1957b  Palaeocaris  of.  typus  Meek  and  Worthen.  Copeland,  p.  47;  pi.  15,  fig.  1. 

7959  Palaeocaris  typus  Meek  and  Worthen.  Siewing,  p.  3. 

1959  Acanthotelson  inequalis  Meek  and  Worthen.  Siewing,  p.  102. 

7967  Palaeocaris  typus  Meek  and  Worthen.  Rolfe,  p.  548. 

V.  19626  Palaeocaris  typus  Meek  and  Worthen.  Brooks,  p.  240,  pis.  60-64,  text-pis.  12  (fig.  a),  13. 

7965  Palaeocaris  species  Noodt,  p.  82. 

1969  Palaeocaris  typus  Meek  and  Worthen.  Brooks,  p.  R348,  figs.  165-1,  167,  170- la,  171. 

1969a  Palaeocaris  typus  Meek  and  Worthen.  Schram,  p.  219,  table  1. 

1969b  Palaeocaris  typus  Meek  and  Worthen.  Schram,  p.  201. 

7972  Palaeocaris  typus  Meek  and  Worthen.  Secretan,  p.  3. 


218 


^ 


Figure  15.  A)  Reconstruction  oi Palaeocaris  typus,  (corrected  from  Brooks  1962^?),  scale  5  mm;  B)  tailfan 
to  same  scale  as  body;  dorsal  views  of  right  C)  antennule  and  D)  antenna;  E)  posterior  thoracopod.  Appendages 
slightly  enlarged. 


797^  Palaeocaris  typus  Meek  and  Worthen.  Schram  and  Schram,  p.  101. 

1976a  Palaeocaris  typus  Meek  and  Worthen.  Schram,  p.  21. 

1976b  Palaeocaris  typus  Meek  and  Worthen.  Schram,  p.  411. 

7975  Palaeocaris  typus  Meek  and  Worthen.  Schminke,  p.  235,  fig.  17. 

1979a  Palaeocaris  typus  Meek  and  Worthen.  Schram,  p.  28,  table  1. 

1979b  Palaeocaris  typus  Meek  and  Worthen.  Schram,  p.  167,  table  2. 

1981a  Palaeocaris  typus  Meek  and  Worthen.  Schram,  p.  131,  text  fig.  5c,  table  2. 

1981^  Palaeocaris  typus  Meek  and  Worthen.  Schram,  p.  9,  fig.  in  text. 

Diagnosis.  — HQdid  to  thorax  ratio  1:4.  Scaphocerite  longer  than  the  two  peduncular 
segments  of  antennal  flagellum.  Sixth  thoracomere  larger  dorsally  than  any  other  tho- 
racomere.  Uropodal  diaeresis  slightly  curved  to  straight,  outer  margin  of  exopod  with 
widely  spaced  spinose  setae  along  its  length  terminating  in  3  spines  just  anterior  to 
diaeresis.  Telson  ovoid  but  wider  proximally  than  distally. 

Holotype.  —  X33S  (Plate  5,  fig.  A),  Mazon  Creek  area,  Will  County,  Illinois.  Francis 
Creek  Shale,  Carbondale  Formation  (Westphalian  C),  Pennsylvanian.  [Brooks  (\962b: 
248)  states  that  the  types  of  this  species  "are  misplaced  or  lost."  However,  close 
comparison  of  X338  with  the  description  of  Meek  and  Worthen  (1868,  fig.  A:552), 
and  the  more  detailed  treatment  in  their  original  description  (Meek  and  Worthen  1 865), 
reveal  that  this  specimen  was  undoubtedly  the  basis  for  these  texts  and  the  1868 
illustration,  and  is  thus  almost  certainly  the  holotype.] 

Other  localities.— Abandoned  Chieftan  Mine,  7  miles  south  of  Terra  Haute,  In- 
diana, east  of  Highway  US-4 1 ;  Lower  Shelbum  Formation,  Pennsylvanian.  Abandoned 
strip  mine  talus  1.8  miles  west  of  Windsor,  Missouri,  on  Highway  MO-2. 

Description.— Cephalic  shield  smooth,  except  for  slight  lateral  groove  at  level  of 
mandible.  Rostrum  small.  Optic  notch  prominent.  Ventral  margins  of  cephalic  shield 
rounded  and  whole.  Eyes  moderate  in  size,  stalk  with  prominent  (peracarid-like)  papilla. 
Antennular  peduncle  3-segmented,  proximal-most  joint  as  long  as  distal  2  joints,  medial 
margins  setose,  inner  flagellum  shorter  than  outer  flagellum,  outer  flagellum  about  one- 
third  body  length.  Antennal  protopod  with  short  proximal  joint  and  large  distal  joint, 
scaphocerite  oval  and  setose,  2  basal  flagellar  joints  peduncular,  medial  margins  of 
peduncular  joints  setose,  flagellum  equal  to  body  length.  Mandible  massive,  promi- 
nently projecting  below  cephalic  shield  margin,  palp  of  at  least  2  segments.  Maxillules 
and  maxillae  with  palps  (details  uncertain). 

First  thoracomere  markedly  reduced,  sixth  thoracomere  larger  than  any  other. 
Thoracic  pleura  broadly  rounded  anteriorly,  posterior  margin  straight.  First  thoracopod 


219 


Plate  5 

Figures  A-C.  Palaeocaris  typus  Meek  and  Worthen,  1 865;  A)  latex  peel  of  holotype,  X  388,  x  1 .9;  B)  YPM 
19765,  showing  the  flap-like  thoracopodal  exopods,  x  4.5;  C)  PE  23237.  showing  annulate  pleopods  (arrows), 
x8.9. 

Figure  D.  Palaeocaris  retracta  Caiman,  1932,  I  13971,  with  short  section  of  an  annulate  pleopod  (arrow), 
x6.7. 


about  one-half  the  size  of  succeeding  appendages.  Thoracopods  2  through  8  subequal, 
epipods  present  (details  not  known),  exopods  broadly  flap-like  (Plate  5,  fig.  B)  and 
setose,  endopods  with  short  ischia  and  dactyli,  meri  and  carpi  long  and  subequal. 
propodi  one-half  the  length  of  carpi. 

Pleomeres  with  posterior  margins  finely  setose,  pleura  of  first  5  abdominal  segments 
as  in  thoracomeres.  Pleopods  annulate  (Plate  5.  fig.  C)  [not  flap-like  as  reported  by 
Brooks  1962].  Telson  oval,  somewhat  wider  anteriorly  than  medially  or  posteriorly, 
margins  with  strong  setae.  Uropods  flap-like,  faint  median  reinforcing  rib  on  setose 


220 


Figure  16.     A)  Reconstruction  of  Palaeocaris  retractata,  (modified  from  Schram  1979a),  scale  5  mm;  B) 
tailfan  to  same  scale  as  body;  C)  dorsal  view  of  right  antenna;  D)  thoracopod.  Appendages  slightly  enlarged. 


rami,  exopods  with  lateral  margin  spinose,  setae  distally  developed  as  3  small  spines 
just  anterior  of  straight  to  slightly  curved  diaeresis. 

Remarks.  — brooks  (1962Z?)  made  some  errors  in  anatomical  interpretation  of  the 
P.  typus  specimens  available  to  him.  He  felt  the  antennal  peduncle  had  5  segments,  2 
protopodal  and  3  flagellar.  The  mistake  arose  in  interpreting  a  preservation  anomaly 
on  the  distal  protopodal  joint,  mistaking  a  longitudinal  crack  in  that  joint  for  a  lon- 
gitudinal suture. 

More  importantly.  Brooks  reconstructed  the  pleopods  of  P.  typus  as  flap-like,  and 
compared  them  to  those  of  Acanthotelson  stimpsoni.  The  pleopods  are  rarely  well- 
preserved  on  P.  typus.  The  thoracic  exopods  are  clearly  flap-like  (e.g.,  YPM  19765, 
Plate  5,  fig.  B).  The  pleopods,  however,  are  annulate  [YPM  19731,  YPM  19765,  PE 
23237  (Plate  5,  fig.  C),  PE  37893,  PE  37957,  PE  37976].  It  is  the  correction  of  our 
understanding  of  this  feature  and  its  detection  on  other  species  of  Palaeocaris  that 
delineates  the  family  Palaeocarididae  from  other  palaeocaridacean  syncarids. 


Palaeocaris  retractata  Caiman  1932 
Fig.  16;  Plate  5,  fig.  D;  Plate  6,  fig.  A 

V.  191  Id  Palaeocaris  praecursor  (Woodward).  Caiman,  p.  488,  figs.  1,  2a,  3. 

V.  1914  Palaeocaris  species.  Peach,  p.  146,  pi.  4,  fig.  9. 

*1932  Palaeocaris  retractata  Caiman,  p.  541. 

1959  Palaeocaris  retractata  Caiman.  Siewing,  p.  101. 

V.  1961  Palaeocaris  retractata  Caiman.  Rolfe,  p.  546,  pi.  68,  fig.  8,  text-fig.  1. 

1962b  Palaeocaris  retractata  Caiman.  Brooks,  p.  248. 

1979(2  Palaeocaris  retractata  Caiman.  Schram,  p.  106,  figs.  50,  51. 

1979b  Palaeocaris  retractata  Caiman.  Schram,  p.  170,  table  2. 

1981a  Palaeocaris  retractata  Caiman.  Schram,  p.  131,  text  fig.  5e,  table  2. 

1982  Palaeocaris  retractata  Caiman.  Schram,  p.  123,  fig.  8. 

Diagnosis.  — Head  to  thorax  ratio  1:2.8.  All  thoracic  segments  subequal.  Uropodal 
diaeresis  a  rounded  to  sigmoid  curve,  outer  margin  of  exopod  armed  with  spines.  Telson 
ovoid,  margin  with  spinose  setae. 

Holotype.  — In  29012.  Clay  Craft  open  works,  Cosely  near  Dudley,  Worcestershire. 
Ten  foot  Ironstone  Measures,  Lower  Similis-Pulchra  Zone,  Middle  Coal  Measures. 

Other  locality.  — WqsI  flank  Bilberry  Hill,  in  Lickey  Hills  southwest  of  Birmingham, 
Warwickshire;  Keele  Beds,  Weslphalian  D. 


Plate  6 

Figure  A.     Palaeocaris  retractata  Caiman,  1932,  I  13973,  showing  setose  margins  of  telson,  x7. 

Figures  B-G.  Palaeocaris  secretanae  new  species:  B)  holotype,  AM  7423,  x  3.6;  C)  AM  7243,  closeup  of 
cephalon,  showing  stalked  eyes,  bases  of  antennules  and  antennae,  and  mouthparts,  x  10.7;  D)  AM  4293, 
closeup  of  annulate  pleopod,  x  22.3;  E)  AM  7861,  showing  optic  notch,  cephalic  groove,  large  mandible,  and 
thoracopodal  exopods,  x  10.7;  F^  AM  5019,  showing  cephalic  groove  and  the  markedly  reduced  first  tho- 
racomere,  xio.7;  G)  AM  7810,  lateral  preservation  of  posterior  abdomen,  with  telson  (t)  and  pleomeres 
(numbered),  x  10.7. 

Figure  H.     Nectotelson  /cre/"c/7  (Fritsch)  1875,  B  77621,  tailfan,  x6.6. 


221 


222 

Descriptions.  — Cephalic  shield  smooth,  slight  lateral  groove  at  level  of  mandible, 
premandibular  portion  of  cephalon  as  long  or  longer  than  the  posterior  region.  Rostrum 
small.  Optic  notch  slight.  Eyes  small  to  medium  in  size.  Antennular  peduncle  with  3 
subequal  joints,  the  most  proximal  with  an  optic  fossa.  Antennal  protopod  with  distal 
joint  long,  scaphocerite  small  and  oval,  2  basal-most  joints  of  flagellum  petiuncular. 
Mandible  large  with  prominent  incisor  process. 

All  thoracomeres,  except  the  markedly  reduced  first,  subequal  in  length.  Second 
through  fourth  thoracic  pleura  subquadrangular,  posterior  pleura  broadly  rounded  an- 
teriorly, all  pleura  with  slightly  marginal  furrows.  Second  through  eighth  thoracopods 
with  epipods  possibly  flap-like,  moderate  flap-like  exopods,  endopodal  joints  subequal 
with  a  tendency  to  shorten  as  one  proceeds  distally. 

Second  through  sixth  pleomeres  with  setose  posterior  margins.  Pleopods  annulate 
(Plate  5,  fig.  D).  Telson  oval,  margins  setose  (Plate  6,  fig.  A).  Uropodal  exopod  with 
rounded  to  sigmoid  diaeresis,  lateral  margins  spinose,  other  margins  of  rami  setose  (at 
least  distally). 

Remarks.— Reexamination  and  preparation  of  available  material.  In  29013,  In 
29014,  and  especially  I  13971  (Plate  5,  fig.  D),  indicates  that  the  pleopods  of/',  retractata 
are  annulate.  This  was  not  noticed  in  the  redescription  of  Schram  {\919a). 

Now  that  other  species  of  Palaeocaris  are  better  understood,  the  large  cephalon 
(small  head  to  thorax  ratio)  is  seen  as  quite  diagnostic  for  this  species.  Other  species, 
P.  typus  and  P.  secretanae,  have  relatively  smaller  heads. 

Restudy  in  July  of  1980  of  all  P.  retractata  material  mentioned  in  Schram  (1979<3) 
aflowed  me  an  opportunity  to  reconsider  the  identity  of  doubtful  specimens  in  light  of 
these  collateral  studies  of  all  fossil  syncarids.  I  now  feel  that  an  incomplete  specimen, 
GSL  RAE  1 29 1 ,  is  not  an  example  of  P.  retractata.  This  correction  does  not  affect  our 
understanding  of  the  anatomy  of  this  species.  However,  it  does  shorten  the  biostratig- 
raphic  range  of  P.  retractata  (Schram  1979(3:7,  fig.  1),  now  understood  to  extend  only 
from  Westphalian  B  to  D,  i.e.,  from  the  Lower  Similis-Pulchra  Zone  up  into  the  Tenuis 
Zone. 

Palaeocaris  secretanae  new  species 
Fig.  17;  Plate  6,  figs.  B-F;  Plate  7,  figs.  A  &  B 

V.  1980a  palaeocarid  syncarid.  Secretan,  p.  24,  pi.  1. 

V.  1980a  ?ceratiocarid  phyllocarid.  Secretan,  p.  28,  pi.  2,  figs.  1,  2. 

V.  1980a  ?eocaridacean.  Secretan,  p.  28,  pi.  3,  figs.  2,  3. 

V.  1980a  pygocephalomorph.  Secretan,  p.  30,  pi.  3,  figs.  5,  6. 

V.  1980a  ?palaeostomatopod.  Secretan,  p.  30. 

V.  1980a  "specimens  enigmatiques."  Secretan,  p.  32,  pi.  4,  figs.  2  &  6. 

1980b  Palaeocaris.  species.  Secretan,  p.  414,  pis.  1-4,  fig.  1. 

1981  Palaeocaris  cf.  P.  retractata  Caiman.  Pacaud  et  al.,  p.  40. 

1982  Palaeocaris  cf.  P.  retractata  Caiman.  Rolfe  et  al.,  p.  426. 

Diagnosis.  — Head  to  thorax  ratio  1:4.6.  All  thoracic  segments  subequal.  Uropodal 
diaeresis  markedly  circular,  outer  margin  of  exopod  armed  with  small  spines  distally 
near  diaeresis,  margins  of  rami  with  long  dense  setae,  rami  reinforced  with  heavy 
median  ribs.  Telson  ovoid  with  distal  end  blunt,  margin  with  long  spines. 

Holotype.— AM  7423-24  (Plate  6,  figs.  B  &  C).  From  shales  above  First  Blanzy- 
Montceau  Coal  (=-Puits  St.  Louis),  Stephanian  B,  Upper  Carboniferous.  St.  Louis  open 
cast  mine,  Montceau-les-Mines,  France. 

Etymology.  — Named  in  honor  of  Dr.  Sylvie  Secretan,  who  first  recognized  the 
nature  and  significance  of  this  material,  and  who  has  been  a  major  figure  in  organizing 
and  coordinating  the  scientific  study  of  the  important  biota  of  the  Montceau-les-Mines 
locality. 

Description.  — Cephalic  shield  smooth,  slight  lateral  groove  at  level  of  mandible 
(AM  5019;  7861,  Plate  6,  figs.  E  &  F).  Rostrum  small  to  moderate  in  size.  Optic  notch 
prominent  (AM  7861).  Eyes  moderate  to  large  (AM  7423-24).  Antennular  peduncle 
with  3  subequal  segments,  median  margins  setose  (AM  6 1 37-38),  flagella  well  developed 


223 


Figure  17.     A)  Reconstruction  of  Palaeocaris  secretanae,  scale  5  mm;  B)  tailfan  to  same  scale  as  body; 
dorsal  views  of  C)  antennule  and  D)  antenna;  E)  thoracopod.  Appendages  slightly  enlarged. 


(but  of  undetermined  length).  Antennal  protopod  with  median  margins  setose  (AM 
7794),  scaphocerite  oval,  small  in  size  and  setose,  2  proximal-most  joints  of  flagellum 
peduncular  and  very  large  (AM  7524).  Mandible  (AM  5744)  with  large  incisor  process 
and  well-developed  palp. 

Thoracic  and  abdominal  pleura  with  gently  rounded  anterior  comers  and  well- 
developed  posterior  comers  (AM  7810,  Plate  6,  fig.  G),  all  body  segments  (except  first) 
subequal.  First  thoracopod  reduced  in  size  (AM  5019).  Second  through  eighth  thora- 
copods  with  long  cylindrical  epipodites  (AM  7861),  exopods  large  and  flap-like  with 
setose  margins,  endopodal  ischium  and  dactylus  small,  merus  and  propodus  long,  and 
carpus  moderate  in  length  (AM  7423-24). 

Annulate  pleopods  setiferous  (AM  4293,  Plate  6,  fig.  D;  4377,  7424).  Telson  with 
margin  spinose  (AM  7436,  Plate  7,  fig.  B).  Uropodal  rami  spatulate  (AM  5080,  Plate 
7,  fig.  A),  reinforced  with  strong  median  ribs,  exopodal  lateral  margin  with  small  spines 
distally  (AM  5080),  diaeresis  strongly  circular  (Plate  7,  fig.  B),  ramal  margins  with 
dense  array  of  long  setae. 

Remarks.— Though  closely  resembling  P.  retractata,  P.  secretanae  is  easily  distin- 
guished by  its  short  cephalon  (shorter  than  that  of  any  species  of  Palaeocaris)  and  the 
spatulate  nature  of  the  uropodal  rami,  i.e.,  more  narrow  proximally  than  distally. 

Although  the  general  preservation  of  most  of  the  material  of  P.  secretanae  is 
exceptionally  fine,  those  specimens  that  were  not  so  well-preserved  can  be  rather  con- 
fusing to  interpret.  This  accounts  for  the  variety  of  tentative  assignments  made  by 
Secretan  (1980a,  b).  Her  non-syncarid  identifications  are  all  based  on  poorly  preserved 
specimens.  In  point  of  fact,  though  the  biota  at  Montceau-les-Mines  is  among  the  most 
diverse  in  the  Carboniferous,  P.  secretanae  remains  the  only  malacostracan  presently 
known  from  that  fauna,  save  for  one  specimen  that  is  possibly  a  phreatoicid  isopod. 
Measurements  comparable  to  those  made  by  Brooks  (1962b)  on  P.  typus  are  provided 
in  Table  4. 


Family  SQUILLITIDAE  Schram  and  Schram,  1974 

Diagnosis.  — Thoracic  exopods  annulate,  pleopods  annulate  and  either  uni-  or  bi- 
ramous. 

Type  genus.  —  Squillites  Scott,  1938. 


224 


\ 


V'*^  '♦> 


^^•^•*^ 


Plate  7 
Figures  A  &  B.     Palaeocaris  secretanae  new  species;  A)  AM  5080,  tailfan,  x  10.7;  B)  AM  7436,  tailfan, 
xlO.7. 

Figures  C-G.  Nectotelson  krejcii  (Fritsch)  1875;  C)  B  7762b,  showing  prominent  cephalic  groove  (arrow) 
and  8  subequal  thoracomeres  (numbered),  x  5.6;  D)  M  1 042,  typical  form  and  preservation  of  Czech  localities, 
X  7;  E)  B  7762i,  showing  subequal  trunk  segments  (numbered)  and  fragments  of  annulated  pleopods  (arrows), 
X  7;  F)  B  7762d,  latex  peel  showing  antennae  and  thoracopodal  endopods,  x  6.7;  G)  M  1033,  showing  setose 
margins  of  telson,  x5. 


225 


Table  4.     Representative  length  measurements  in  mm  of  species  of  Palaeocaris*  Holotype. 


Cephalon 

Thorax 

Abdomen 

T. 

A. 

A. 

Telson 

Body 

P.  secretaneae 

AM  3424 

1.8 

8.3 

1.2 

0.9 

4293 

1.5 

7.7 

1.3 

1.1 

4301-02 

9.4 

1.4 

2.4 

2.7 

4385-86 

1.6 

6.8 

1.1 

4800 

2.5 

5019 

1.5 

6.3 

5032-33 

1.6 

10.0 

8.1 

1.4 

1.2 

1.8 

5080 

6.8 

5610-11 

1.8 

2.3 

5616 

2.0 

5617 

2.5 

5744 

1.4 

6.4 

4.4 

1.0 

7103-04 

3.9 

0.7 

1.0 

7352 

1.5 

7.8 

1.0 

7372-73 

5.4 

0.8 

0.8 

1.5 

7420 

2.0 

8.0 

0.9 

*7423-24 

1.5 

7427-28 

2.0 

7429-30 

1.3 

11.5 

7436 

1.4 

2.3 

7440 

7.8 

1.4 

1.9 

2.3 

7454-55 

2.8 

7528 

5.1 

0.8 

1.3 

7534 

1.4 

5.4 

0.8 

0.5 

7556 

1.1 

5.0 

4.2 

0.7 

0.6 

1.2 

10.0 

7766-67 

4.5 

0.7 

1.2 

7794 

9.3 

7.5 

7810 

3.2 

7845 

1.8 

2.2 

7861 

2.3 

10.2 

7.8 

1.5 

1.4 

1.9 

20.0 

8236 

5.9 

1.5 

11.8 

8338-39 

1.9 

X 

1.67 

7.6 

5.9 

1.1 

1.0 

1.6 

2.5 

13.3 

P.  typus  X 

2.3 

9.3 

2.2 

1.2 

1.9 

3.6 

20.4 

P.  retractata  x 

2.0 

5.8 

5.3 

3.4 

Remarks.— TogQXhtx  the  annulate  thoracic  exopods  and  pleopods  of  these  species 
most  resemble  those  seen  in  the  living  Anaspididae  within  the  order  Anaspidacea. 
However,  the  first  thoracomere  is  not  fused  to  the  cephalon  in  these  fossils:  equal  in 
size  to  all  other  thoracomeres  in  Nectotelson,  slightly  reduced  in  Squillites,  and  greatly 
reduced  in  Praenaspides. 

Genus  SQUILLITES  Scott,  1938 

Diagnosis.  — ¥'\rs,\  thoracomere  only  slightly  reduced.  Uropods  narrow,  spatulate, 
and  setose.  Telson  subtriangular,  armed  with  moveable  spines. 
Type  species.— Squillites  spinosus  Scott,  1938. 


Squill  it  es  spinosus  Scott,  1938 
Fig.  18 

v.*  1938  Squillites  spinosus  Scott,  p.  508,  figs.  1,  2. 

1939  Squillites  spinosus  Scott.  Berry,  p.  467. 

1962^  Squilites  spinosus  Scott.  Brooks,  p.  254,  pi.  53,  figs.  1,  3;  text-pl.  14,  fig.  d. 

1965  Squillites  species.  Noodt,  p.  82. 

7967  Squillites  spinosus  Scott.  Secretan,  p.  173,  fig.  8. 

1969  Squillites  spinosus  Scotx.  Brooks,  p.  R355,  figs.  169-1  and  170-3. 

1969a  Squillites  spinosus  Scott.  Schram,  p.  216,  table  1. 

V.  1974  Squillites  spinosus  Scott.  Schram  and  Schram,  p.  96,  pis.  1-2,  text-figs.  1,  2. 


226 


Figure  18.     A)  Reconstruction  of  Squillites  spinosus,  (modified  from  Schram  and  Schram  1974),  scale  5 
mm;  B)  tailfan  to  same  scale  as  body;  C)  dorsal  view  of  right  antenna  slightly  enlarged. 


J 97 9a  Squillites  spinosus  Scott.  Schram,  p.  114. 

1981  Squillites  spinosus  Scott.  Schram,  p.  133,  text-fig.  4. 

7952  Squillites  spinosus  Scott.  Schram,  p.  122. 

Diagnosis.  —  Since  there  is  but  one  species,  the  diagnosis  of  the  species  is  the  same 
as  that  of  the  genus. 

Holotype.  —  Xl  2 19.  One-half  mile  south  of  Heath,  Fergus  County,  Montana;  Heath 
Shale,  Big  Snowy  Group,  Upper  Mississippian. 

Description.  — Stalked  compound  eyes  small  and  spherical.  Antennule  with  3-seg- 
ment  peduncle,  flagella  long.  Antennal  protopod  with  only  1  segment  observed,  scaph- 
ocerite  oval  and  setose,  very  long  flagellum  with  proximal  2  segments  large  and  pe- 
duncular. Cephalon  with  marked  broad  rostrum,  lacking  any  cervical  grooves,  pair  of 
semicircular  mid-dorsal  ridges. 

Thoracomeres  with  4  anterior  pleura  medially  pointed  and  4  posterior  pleura 
rounded,  pair  of  semicircular  ridges  mid-dorsal  on  each  thoracomere,  first  thoracomere 
slightly  shorter  than  others.  Thoracopods  subequal,  ischia  short  and  equal  to  bases, 
meri  long,  carpi  through  dactyli  short. 

Pleomeres  variously  decorated,  first  through  fifth  with  setose  posterior  margins, 
first  and  second  with  mid-dorsal  paired  semicircular  ridges,  and  third  through  fifth  with 
large  immobile  posteriorly  directed  spines.  First  pleuron  rounded,  second  through  fifth 
pleura  with  posterior  comers  denticulate.  Setose  pleopods  robust  and  uniramous.  Sixth 
pleomere  elongate.  Uropods  as  oval  flaps,  margins  finely  setose.  Telson  subtriangular, 
with  median  keel  and  1 7  pairs  of  moveable  marginal  spines. 

Remarks.— There  is  a  slight  reduction  in  the  size  of  the  first  thoracomere,  but 
reconsideration  of  the  original  material  of  Schram  and  Schram  (1974)  leaves  some 
question  as  to  whether  their  first  thoracopod  is  as  well  developed  as  those  on  the  other 
thoracic  segments. 

The  apparent  single  joint  on  the  antennal  protopods  may  be  an  artifact  of  pres- 
ervation, but  oddly  coincides  with  an  apparent  similar  phenomenon  on  Praeanaspides 
praecursor,  which,  if  it  is  confirmed,  may  provide  another  derived  feature  to  characterize 
the  family. 


Genus  PRAEANASPIDES  Woodward,  1908 

Diagnosis.  — ¥\rs\  thoracomere  very  reduced.  Pleopods  biramous.  Uropodal  exo- 
pod  with  distinct  circular  diaeresis.  Telson  rectangular,  laterally  spinose. 


227 


Figure  19.     A)  Reconstruction  of  Praeanaspides  praecursor,  scale  5  mm  (redrawn  from  Schram  1979(2);  B) 
tailfan  to  same  scale  as  body;  C)  dorsal  view  of  right  antenna  slightly  enlarged. 


Type  species.— Praeanaspides  praecursor  Woodward,  1 908. 

Praeanaspides  praecursor  Woodward,  1 908 
Fig.  19 

v.*  1908  Praeanaspides  praecursor  Woodv/ard,  p.  385,  figs.  1-5. 

V.  1908  Paleocaris  lansboroughi  Peach,  p.  55,  pi.  8,  figs.  8-10. 

1911  Paleocaris  lansboroughi  Peach.  Woodward,  p.  363. 

7927  Paleocaris  praecursor  {V^ood\^ard).  Chappuis,  p.  605. 

1959  Praeanaspides  praecursor  Woodward.  Siewing,  p.  10. 

1959  Paleocaris  praecursor  V^oodvjavd.  Siewing,  p.  101. 

1932  Paleocaris  praecursor  (V^oodward).  Caiman,  p.  537,  figs.  1,  2. 

1962b  Paleocaris  praecursor  (Woodward).  Brooks,  p.  249. 

1969a  Paleocaris  praecursor  (Woodward).  Schram,  p.  220,  table  1. 

7976  Praeanaspides  praecursor  Woodward.  Schram,  p.  411. 

1979a  Praeanaspides  praecursor  Woodward.  Schram,  p.  112,  figs.  54,  55. 

1979b  Praeanaspides  praecursor  Woodward.  Schram,  table  2. 

7957  Praeanaspides  praecursor  Woodward.  Schram,  p.  131,  table  2. 

Diagnosis.  — S>mcQ  there  is  but  one  species,  the  diagnosis  of  the  species  is  the  same 
as  that  of  the  genus. 

Lectotype.—GSh  30213-14.  Shipley  Hall,  I'A  miles  NW  of  Ilkestone,  Derbyshire, 
England.  Clay  Ironstone,  top  Modiolaris  Zone,  Middle  Coal  Measures. 

Other  locality.— GvQQnhxW,  or  Woodhill  Quarry,  near  Kilmaurs,  Ayrshire;  roof  of 
Pinnies  Main  Coal,  Middle  Coal  Measures,  Westphalian  B. 

Description.— A.r\\Qr\n\x\Qs  with  3-segment  peduncle,  middle  joint  shorter  than  either 
proximal  or  distal  unit,  flagella  relatively  short.  Antennae  with  only  single  segment 
visible  in  protopod,  oval  non-setose  scaphocerite,  flagellum  very  long  with  proximal  2 
joints  peduncular.  Cephalon  unomamented,  slight  rostral  projection. 

First  thoracomere  smooth  with  no  ornament,  all  other  segments  with  3  or  4  tergal 
ridges  and  rounded  pleura.  First  thoracopod  apparently  reduced  (never  seen).  Second 
through  eighth  subequal,  ischium  and  merus  longer  than  carpus  and  propodus  (dactyls 
not  seen).  Pleopods  with  exopod  slightly  longer  than  endopods,  rami  thin.  Uropodal 
exopod  lateral  margin  spinose,  endopod  subtrapezoidal  with  its  longer  margin  medial. 
Telson  rectangular  but  somewhat  bilobed  terminally,  lateral  margin  with  12-13  pair 
of  moveable  spines. 


228 


Figure  20.     A)  Reconstruction  of  Nectotelson  krejcii,  scale  5  mm;  B)  tailfan  to  same  scale  as  body;  C)  dorsal 
view  of  right  antenna;  D)  posterior  thoracopod.  Appendages  slightly  enlarged. 


Remarks.  — ThQ  apparently  single  segment  antennal  protopod  is  possibly  an  artifact 
of  preservation,  a  short  proximal  segment  just  may  not  be  observable  on  known  material 
(however,  see  remarks  on  5".  spinosus).  The  same  problem  applies  to  the  thoracopodal 
dactyli;  they  were  probably  short  and  simple,  but  were  simply  not  preserved  on  any 
specimens  now  available. 

Of  all  the  palaeocaridaceans,  P.  praecursor  comes  closest  to  resembling  the  anas- 
pidid  anaspidaceans  in  regard  to  its  very  reduced  first  thoracic  segment  and  in  the 
character  of  its  thoracic  and  abdominal  appendages. 


Genus  NECTOTELSON  Brocchi,  1880 

Diagnosis.  — A\\  thoracopods  (?)  and  thoracomeres  subequal.  Pleopods  biramous. 
Uropods  spatulate,  diaeresis  circular.  Telson  oval  and  spinose. 
Type  species.  — Gampsonychus  krejcii  Fritsch,  1875. 

Nectotelson  krejcii  (Fritsch)  1875 
Fig.  20;  Plate  6,  fig.  H;  Plate  7,  figs.  C-G;  Plate  8,  fig.  A-D 

1870  Gampsonychus  species,  Fritsch,  p.  34. 

1873  Gampsonychus  fimbriatus  Jordan.  Feistmantel,  p.  593,  pi.  18,  figs.  9-12. 

v.*  1875  Gampsonychus  krejcii  Fritsch,  p.  104,  fig.  265. 

V.  1880  Nectotelson  rochei  Brocchi,  p.  10,  pi.  1. 

1885  Gampsonychus  fimhriatus  (Jordan).  Zittel,  p.  672  (in  part). 

1885  Nectotelson  rochei  Brocchi.  Zittel,  p.  673. 

V.  1901  Gasocaris  krejcii  (Fritsch).  Fritsch,  p.  66,  figs.  371-376,  pis.  156-158. 

790/  Nectotelson  rochei  Brocchi.  Fritsch,  p.  74. 

1909  Gasocaris  krejcii  (Fritsch).  Smith,  p.  572,  figs.  59-61. 

1919  Gasocaris  species  Fritsch.  Pruvost,  p.  85. 

1919  Nectotelson  species  Brocchi.  Pruvost,  p.  85. 

1931  Gasocaris  species  Fritsch.  Van  Straelen,  p.  5. 

1959  Gasocaris  krejcii  (Fritsch).  Siewing,  p.  5. 

1959  Nectotelson  rochei  Brocchi.  Siewing,  p.  103. 

?  1960  Eileticus  pruvosti  Vandenberghe,  p.  690,  fig.  2,  pi.  17. 

1965  Nectotelson  species  Noodt,  p.  82. 

1965  Gasocaris  species  Noodt,  p.  82. 


229 


• '''^- ^-JR*^- 15^V      ^^    **    '    >  ■•-V:■'■■■ 


Plate  8 

Figures  A-D.  Nectotelson  krejcii  (Fritsch)  1875:  A)  lectotype,  M  1050,  x4.5;  B)  Me  40,  showing  diaeresis 
on  uropodal  exopod  (arrow),  x5;  C)  B  7762k,  latex  peel  showing  cephalon  (c)  and  thoracopodal  epipodites 
(arrows),  x7;  D)  B7762J,  showing  antennules  and  antennae,  and  thoracopods,  x7. 

Figure  E.  Pleurocaris  annulatus  Caiman,  1911;  I  13814,  latex  peel  showing  thoracic  pleura,  tergite  deco- 
ration, and  proximal  portions  of  thoracopods,  x6.2. 


1969  Palaeocaris  krejcii  {¥hX's,c\\).  Brooks,  p.  R355,  figs.  170- lb. 
1969  Palaeocaris  rochei  (QtoccYlx).  Brooks,  p.  R355,  figs.  170-lc. 
1969a  Palaeocaris  krejcii  (Fritsch).  Schram,  p.  221,  table  1. 
1969a  Palaeocaris  rochei  (Brocchi).  Schram,  p.  221,  table  1. 
1972  Nectotelson  rochei  Brocchi.  Secretan,  p.  1,  1  fig. 


230 


Diagnosis.— Since  but  a  single  species  is  known,  the  diagnosis  of  the  species  is  the 
same  as  that  of  the  genus. 

Lectotype.  —  M  1050  (Plate  8,  fig.  A).  Humboldt  Mine,  Nyfan,  near  Pilsen,  Bo- 
hemia. Gaskohle,  Lower  Permian. 

Other  localities.  — FriXsch  (1901)  also  recorded  this  species  from  the  Gaskohle  of 
the  Krimitz  Mine  and  in  Tremosna.  Brocchi  (1880)  described  the  same  species  under 
another  name  from  the  Lower  Permian  shales  near  Autun,  central  France. 

Description.— Cephalon  lacks  prominent  rostral  extensions  (M  1035,  B  7762g,  1), 
marked  by  rather  deep  and  prominent  mid-dorsal  (B  7762b,  Plate  7,  fig.  C)  groove 
parallel  to  posterior  margin  which  shallows  as  it  extends  toward  ventral  margin  (B 
7762h).  Eye  oval,  moderate  in  size  (B  7762j,  Plate  8,  fig.  D).  Antennular  peduncles 
subequal,  3-segmented  middle  joint  slightly  shorter  than  other  two  (M  1035,  B  7762b, 
e  &  j,  Plate  8,  fig.  D.  Antennal  protopod  with  short  proximal  segment  and  longer  distal 
joint  (B  7762e,  B  7763b),  these  with  slight  longitudinal  ridges  (Me  40).  Scaphocerite 
short,  oval,  and  setose  (Me  40,  B  7762e),  overlapping  2  proximal-most  peduncular 
joints  of  flagellum.  Antennular  and  antennal  flagella  well  developed,  [of  undetermined 
length  because  of  lack  of  preservation  of  distal  joints  (M  1042,  Plate  7,  fig.  D;  B  11626. 
&  j,  Plate  7,  fig.  F)].  Median  margin  of  antennal  peduncle  marked  by  row  of  short 
denticulae  (B  7762e,  B  7763k). 

Mandible  large  but  not  apparently  heavily  sclerotized  or  mineralized  (B  7763a), 
with  an  incisor  process  (B  7762i,  B  7763c).  Maxillules  and  maxillae  with  small  palps, 
maxillary  palp  seems  to  have  several  segments  (B  7763a). 

First  thoracomere  large  (B  7762i,  Plate  7,  fig.  E),  with  fine  serrations  on  posterior 
margin  (B  7763k).  Thoracic  pleura  rounded  posteriorly  (M  1050,  B  77621,  and  the  TV. 
rochei  holotype).  First  thoracopod  large  [but  not  completely  preserved  on  either  the 
Prague  or  Paris  specimens  (e.g.,  B  7763g)],  with  moderately  long  proximal  unit  (?is- 
chium)  followed  by  short  merus,  long  carpus,  and  distal  to  the  knee  may  be  at  least 
another  short  unit.  Second  through  eighth  thoracopods  (Plate  7,  fig.  F)  subequal;  with 
small  epipods  (B  7762k,  Plate  8,  fig.  C);  stout  annulate  exopods  (M  1042,  B  7762d  & 
k);  short  coxae,  bases,  and  ischia;  long  meri;  and  distal  to  knee  progressively  shorter 
carpi,  propodi,  and  dactyli  (M  1042  in  part,  B  7762e,  B  7763f  &  g);  posterior  margins 
of  endopods  with  apparently  fine  setae  (B  7762a). 

Pleomeres  with  acute  postero-ventral  comers  (M  77631);  second  through  sixth 
pleomeres  with  finely  serrate  posterior  margins  (M  42,  M  1033,  M  1054,  B  7762b). 
Sixth  pleomere  almost  twice  as  long  as  any  other  segment  (M  1054,  B  7762f,  B  7763b). 
Pleopods  with  robust,  annulate  rami  subequal  (M  1054,  B  7762b  &  m,  B  7763),  and 
setose  (M  42). 

Uropods  longer  than  telson;  protopod  short,  with  2  faint  longitudinal  ridges,  spat- 
ulate  rami  (B  77621,  Plate  6,  fig.  H),  densely  setose,  and  reinforced  with  sclerotized 
ribs  along  most  of  their  lengths  (M  1044,  M  1054,  B  7763h,  A^.  rochei  type).  Exopods 
laterally  spinose  (M  1054,  B  7762a,  B  7763e)  diaeresis  circular  (Me  40,  Plate  8,  fig.  B), 
segment  beyond  diaeresis  as  a  narrow  oval  (M  1033,  B  7763d).  Telson  elongate  and 
oval;  ornamented  with  stout,  short,  moveable  spines  set  in  sockets  (M  1033,  Plate  7, 
fig.  G;  M  1044,  M  1054,  B  7762b  &  m,  B  7763e). 

Remarks.  — Both  the  Bohemian  and  French  specimens  are  preserved  as  carbonized 
films,  with  varying  degrees  of  pyrite  replacement,  on  a  fine-grained  thinly-bedded  black 
shale.  The  pyrite,  especially  on  the  Bohemian  material,  is  highly  reflective,  and  makes 
studying  and  photographing  specimens  extremely  difficult.  In  only  a  few  instances  were 
fossils  preserved  as  external  molds,  and  these  tend  to  occur  on  shale  fragments  from 
Autun  with  a  higher  content  of  clay  mineral.  The  matrix  in  these  latter  cases  was 
cemented  enough  to  allow  one  latex  peel  to  be  made,  but  otherwise  the  shale  and  fossils 
are  too  friable  to  allow  such  treatment.  In  general,  the  French  material  is  somewhat 
better  preserved  in  contrast  to  the  Czech  specimens. 

There  are  subtle  differences  between  the  two  suites  of  specimens,  especially  as 
regards  the  antennal  peduncles.  That  described  above  generally  represents  that  seen  on 
the  better  preserved  French  specimens.  The  Bohemian  material  seems  to  have  a  some- 
what shorter  distal  segment  of  the  protopod,  a  somewhat  longer  scale,  and  no  detectable 


231 


Table  5.     Representative  measurements  in  mm  of  Nectotelson  krejcii. 


Cephalon 

Thorax 

Abdomen 

Telson 

Blieia 

-3.0 

-0.8 

B 7762b 

1.1 

3.8 

2.8 

0.5 

B  7762c 

0.8 

B7762d 

2.5 

0.6 

B  7762g 

1.2 

B  7762h 

1.4 

B  77621 

4.6 

3.8 

B  77621 

0.8 

B  7763a 

3.2 

2.6 

B  7763b 

1.0 

B  7763c 

4.1 

B7763e 

3.3 

0.8 

B7763g 

-1.3 

B7763h 

1.0 

B  77631 

3.4 

B  7763J 

1.0 

B  7763k 

1.4 

B  77631 

0.9 

M  1033 

0.7 

M  1035 

-1.3 

-5.0 

X 

1.2 

4.1 

3.2 

0.8 

medial  denticulae.  The  Czech  specimens  also  have  slightly  more  distinctive  spines 
along  the  pleomere  margins.  However,  in  light  of  all  the  other  fine  and  detailed  points 
of  agreement  between  the  two  series  of  specimens,  and  caveats  due  to  vagaries  of 
preservation,  I  feel  that  A^.  rochei  Brocchi  must  be  synonymized  with  A^.  krejcii  (Fritsch). 
Nor,  in  light  of  the  distinctive  form  of  the  thoracic  exopods  and  the  first  thoracomere, 
can  this  species  be  maintained  in  the  genus  Palaeocaris  in  the  sense  of  Brooks  (1969). 
Rather,  A^.  krejcii  is  more  closely  aligned  with  the  Squillitidae. 

I  found  no  evidence  of  any  of  the  sexual  dimorphism  reported  by  Fritsch  (1901: 
70-71).  Representative  measurements  of  specimens  are  given  in  Table  5. 

A  vexing  problem  in  this  study  has  been  what  to  do  with  Eilecticus  pruvosti 
Vandenberghe,  1960.  The  published  paper  contains  no  reference  as  to  where  the  types 
were  deposited;  the  photographs,  while  good  enough  to  be  intriguing,  are  not  of  sufficient 
quality  to  facilitate  detailed  study;  and  the  description  of  the  specimens  is  too  vague 
to  be  of  any  real  help.  Attempts  to  find  the  specimens  or  locate  Dr.  Vandenberghe  have 
proved  futile.  However,  certain  items  in  the  published  description  seem  to  hint  that 
this  material  may  belong  within  the  genus  Nectotelson.  Vandenberghe  related  that  E. 
pruvosti  had  an  abdomen  of  7  segments,  a  thorax  of  7  segments,  and  the  first  thoracic 
segment  fused  with  the  cephalon.  Counts  of  body  segments  on  the  published  photo- 
graphs indicate  14  subequal  segments  between  the  head  and  tailfan.  Thus  segment 
count  for  the  abdomen  must  be  wrong,  and  it  seems  logical  to  infer  that  the  abdomen 
had  6  segments  and  the  thorax  8.  The  description  also  makes  reference  to  a  deep  groove 
on  the  cephalon,  and  spatulate  uropodal  rami.  All  these  features  would  seem  to  cor- 
respond to  identical  characters  noted  above  in  Nectotelson.  Consequently,  I  choose  to 
assign  this  species,  with  a  query,  to  A^.  krejcii.  The  question  mark  can  be  removed  only 
by  rediscovery  and  study  of  the  type,  or  recollection  at  the  original  locality. 

E.  pruvosti  occurs  at  the  top  of  the  Griiner  Group,  near  the  Middle  and  Upper 
Stephanian  boundary.  The  material  came  from  a  borehole  in  the  Saint-Etienne  basin, 
and  is  associated  with  the  limuline  Pringlia  demaistrei,  insects,  fish  scales,  and  the 
plant  Odontopteris  pseudoschotheimi. 


Family  uncertain 

Remarks.— The  familial  taxonomy  of  the  Palaeocaridacea  being  adopted  here  is 
an  attempt  to  establish  a  more  natural  system  than  any  used  heretofore,  and  is  based 
on  comparative  morphology  of  thoracic  and  abdominal  appendages.  Such  a  system, 


232 


however,  requires  that  certain  features  of  the  anatomy  of  these  fossils  be  known  before 
famiHal  assignments  can  be  made.  Unfortunately,  fossils  do  not  always  preserve  all  the 
features  that  one  would  like  to  have  information  about.  The  fossil  syncarids  are  no 
exception  to  this,  and  as  a  result  there  are  some  Paleozoic  taxa  that  cannot  be  placed 
within  a  family  with  any  degree  of  certitude,  though  we  can  recognize  them  as  distinct 
and  valid  genera  and  species.  It  was  felt  here  that  the  issue  should  not  be  forced,  and 
that  it  was  preferable  to  simply  recognize  the  uncertainty  and  treat  these  taxa  as  presently 
"unassignable."  Thoughts  as  to  their  affinities  can  sometimes  be  offered,  but  it  seems 
better  to  patiently  await  future  data  which  will  allow  someone  to  definitively  place 
these  problematica. 

Genus  PLEUROCARIS  Caiman,  1911a 

Diagnosis.— Cephalon  small,  2  cephalic  grooves  not  joined  laterally.  Tergites  dec- 
orated with  lateral  ridges.  Thoracic  pleura  very  large.  Telson  and  uropodal  rami  styli- 
form. 

Type  species.— Pleurocaris  annulatus  Caiman,  1911a. 

Pleurocaris  annulatus  Caiman,  1911a 
Fig.  21;  Plate  8,  fig.  E 

?  1881  Palaeocaris  burnetii  Woodward,  p.  534,  pi.  14,  figs.  3a,  3b. 

v.*  19 11a  Pleurocaris  annulatus  Caiman,  p.  156,  fig.  1. 

V.  191  \b  Pleurocaris  annulatus  Caiman.  Caiman,  p.  494,  fig.  5. 

.1912  Eileticus  cf.  aequalis  Scudder.  Pruvost,  p.  66,  pi.  2,  figs.  6,  7. 

79/5  Pleurocaris  annulatus  Caiman.  Chappuis,  p.  173. 

1919  Pleurocaris  annulatus  Caiman.  Pruvost,  p.  86,  fig.  21,  22;  pi.  25,  fig.  11. 

.1919  Eileticus  cf.  aequalis  Scudder.  Pruvost,  p.  89,  fig.  23;  pi.  25,  fig.  12. 

1922  Eileticus  aequalis  Scudder.  Pruvost,  p.  149. 

1923  Pleurocaris  annulatus.  Caiman.  Pruvost,  p.  149. 
7927  Pleurocaris  annularis  Caiman.  Chappuis,  p.  605. 
7959  Palaeocaris  burnetii  Woodward.  Slewing,  p.  101. 
7959  Pleurocaris  annulatus  Caiman.  Slewing,  p.  103. 
1962a  Pleurocaris  annulatus  Caiman.  Brooks,  p.  236. 
7965  Pleurocaris  species.  Noodt,  p.  83. 

1969  Pleurocaris  annulatus  Caiman.  Brooks,  p.  355,  fig.  169-5,  172. 

1969a  Pleurocaris  annulatus  Caiman.  Schram,  p.  220,  table  1. 

?  1969a  Palaeocaris  burnetii  Woodward.  Schram,  p.  220,  table  1. 

7976  Pleurocaris  annulatus  Caiman.  Schram,  p.  411. 

1979a  Pleurocaris  annulatus  Caiman.  Schram,  p.  103,  figs.  48,  49. 

1979b  Pleurocaris  annulatus  Caiman.  Schram,  p.  167,  table  2. 

1981(2  Pleurocaris  annulatus  Caiman.  Schram,  p.  131,  text-fig.  5f;  table  2. 

Diagnosis.  — ^mcQ  there  is  but  one  species,  the  diagnosis  of  the  species  is  the  same 
as  that  of  the  genus. 

Holotype.  — In  29008.  Clay  Croft  mine,  Coseley,  near  Dudley,  Worcestershire, 
England.  Ten  foot  Ironstone  Measures,  Lower  Similis-Pulchra  Zone,  Middle  Coal 
Measures,  Upper  Carboniferous. 

Other  localities.  — Pit  no.  9,  near  Lens,  Belgium;  Black  shale  of  the  Insect  beds, 
beneath  the  "veine  Girard,"  Edouard  Group,  Westphalian  C.  Pit  no.  4,  Vicoigne  Mines, 
France;  Black  shale,  top  of  the  "veine  du  Nord,"  Olympe  Group,  Westphalian  A. 

Descriptions.— Cepha\on  short,  rostrum  small.  Stalked  compound,  eyes  small  and 
rounded.  Details  of  antennules  and  antennae  uncertain. 

First  thoracomere  short,  about  half  the  length  of  any  other.  All  thoracomeres  with 
2  laterally  directed  ridges  on  tergites;  pleura  very  large,  rounded,  set  off  from  tergites 
as  lappets.  Second  through  eighth  thoracopods  with  well-developed  endopods,  meri 
long,  carpi  short  (other  joints  indeterminable). 

Pleomeres  decorated  dorsally  with  lateral  ridges  as  thoracomeres,  pleura  decrease 
in  development  posteriorly.  Telson  styliform,  margins  with  5  pair  moveable  spines. 
Pleopods  possibly  flap-like  (I  14449).  Uropods  as  blades;  exopod  straight,  serrated 
laterally  and  less  conspicuously  so  medially;  endopod  curved  mediad,  lateral  margin 
faintly  spined,  medial  margin  distinctly  so. 


233 


Figure21.     A)  Reconstruction  of  P/eMrocamawwMtoiw,  (modified  from  Caiman  1 9 1 1  a;  and  Schram  1979a), 
scale  5  mm;  B)  tailfan  to  same  scale  as  body. 


Remarks.  One  specimen,  In  14449,  appears  to  have  some  poorly  preserved  prox- 
imal portions  of  the  pleopods.  These  appear  to  be  flap-like.  However,  they  are  of  such 
quality  as  to  be  almost  impossible  to  photograph.  This  was  not  noticed  in  the  original 
study  of  Schram  (1979a).  Though  the  few  British  specimens  known  of  this  species 
(Schram  1979<3:121)  are  for  the  most  part  moderately  well  preserved,  our  knowledge 
of  this  taxon  suffers  because  there  are  so  few  examples  of  it.  All  continental  European 
material  attributable  to  this  species  has  been  lost.  However,  examples  of  this  species 
on  the  continent  have  apparently  never  been  common,  just  as  in  Britain. 

Confusion  in  the  identification  of  specimens  of  this  species  with  Acanthotelson 
stimpsoni  arises  from  the  somewhat  similar  syliform  telson  and  uropods.  However,  the 
short  cephalon,  subequal  second  through  eighth  thoracomeres,  large  thoracic  pleura, 
and  tergal  decoration  clearly  justify  a  separate  generic  status  for  this  species  (Plate  8, 
fig.  E).  However,  exact  familial  affinities  must  remain  uncertain  until  such  time  as  the 
structure  of  the  thoracic  exopods,  distal  joints  of  the  endopods,  and  pleopods  can  be 
ascertained.  The  general  form  of  the  tailfan  noted  here,  as  well  as  a  supposed  analogue 
correspondence  to  A.  stimpsoni  in  the  Carboniferous  brackish  water  habitat  community 
(Schram  1981a)  may  suggest  that  P.  annulatus  could  be  eventually  assigned  to  the 
acanthotelsonids. 

Pruvost  (1912,  1919)  described  2  syncarid  specimens  which  he  variously  referred 
to  Eileticus  cf.  aequalis  and/or  Pleurocaris  annulatus.  The  specimens  were  deposited 
in  the  museum  at  the  University  of  Lille,  but  are  now  lost.  The  published  descriptions 
and  illustrations  of  these  specimens,  combined  with  Pruvost's  own  stated  reservations 
on  what  he  called  E.  cf.  aequalis,  indicate  these  were  indeed  examples  off.  annulatus. 


Genus  WILLIAMOCALMANIA  new  genus 

Diagnosis.  — ¥'\rs\  thoracomere  markedly  reduced,  second  thoracomere  longer  than 
first  but  less  than  any  other  thoracomeres.  Thorax  shorter  than  abdomen,  ratio  about 
0.9:1.  Telson  elongate,  subtriangular,  distal  end  rounded.  Uropodal  rami  oval,  some- 
what longer  than  telson. 

Type  species.  — Palaeocaris  vandergrachti  Pruvost,  1922. 

jE'/.vwo/o^.  —  Named  in  honor  of  W.  T.  Caiman,  among  whose  many  accomplish- 
ments was  his  expertise  on  fossil  and  recent  syncarids. 

Remarks.— The  distinctive  pattern  of  reduction  of  the  anterior  thoracomeres,  the 
unique  thorax-abdomen  ratio,  and  the  characteristic  tailfan  clearly  separate  this  species 


234 


Figure  22.    A)  Reconstruction  of  Williamocalmania  vandergrachti,  scale  5  mm;  B)  tailfan  to  same  scale  as 
body. 


from  those  herein  included  in  the  newly  redefined  genus  Palaeocaris.  Consequently,  a 
separate  generic  designation  is  necessary  for  this  taxon.  Though  obviously  distinct  from 
Palaeocaris  or  any  other  known  fossil  syncarid,  the  lack  of  sufficient  information  about 
body  appendages  dictates  an  uncertain  family  affinity  for  this  species. 

Williamocalmania  vandergrachti  (Pniyost)  1922 
Fig.  22;  Plate  9,  figs.  A-C 

v*1922  Palaeocaris  vandergrachti  Pruvost,  p.  147,  fig.  1. 

7927  Palaeocaris  vandergrachti  Pruvost.  Chappuis,  p.  605. 

1930  Palaeocaris  vandergrachti  Pruvost.  Pruvost,  p.  181,  fig.  5,  pi.  8. 

1959  Palaeocaris  vandergrachti  Pruvost.  Slewing,  p.  101. 

1969a  Palaeocaris  vandergrachti  Pruvost.  Schram,  p.  220,  table  1. 

Diagnosis.  — Since  but  one  species  is  known,  the  diagnosis  of  the  species  is  the 
same  as  that  of  the  genus. 

Lectotype.—NB  7183  Ech.  no.  1  (see  Pruvost  1930,  plate  8,  fig.  la).  Woensdrecht 
borehole  (1 164-1 167  m).  The  Netherlands;  Chokier  Ampelite,  Lower  Namurian. 

Description.  — Body  moderate  to  large  in  size.  Antennular  peduncle  large,  with  3 
subequal  segments.  Antennal  peduncles  large,  scaphocerite  appears  large  and  ovoid,  2 
(or  3)  proximal-most  segments  of  flagellum  peduncular  and  very  large.  Cephalon  with 
rostrum,  prominent  optic  notch. 

First  thoracomere  reduced  and  closely  associated  with  cephalon,  second  thorac- 
omere  shorter  than  posterior  thoracomeres  but  larger  than  first,  all  other  thoracic  and 
anterior  abdominal  segments  subequal.  Pleura  somewhat  subtriangular,  attenuated  along 
anterior  margin  (Plate  9,  fig.  C),  with  slight  marginal  furrows.  Thoracopods  apparently 
equally  developed,  exopods  possibly  flap-like  (Plate  9,  fig.  A).  Thorax  somewhat  shorter 
in  length  than  abdomen. 

Sixth  pleomere  elongate.  Telson  long,  subtriangular  with  distal  end  rounded,  mar- 
gins with  stout  spinose  setae  (Plate  9,  figs.  B  &  C).  Uropodal  rami  oval;  exopod  rein- 
forced proximally  with  medial  rib,  with  slightly  curved  diaeresis,  somewhat  longer  than 
endopod. 

Remarks.— The  later  treatment  by  Pruvost  (1930)  of  this  species  is  superior  in 
most  respects  to  the  original  description  (Pruvost  1922),  especially  in  regards  to  the 
plate  figures  which  illustrate  all  4  of  the  available  specimens.  Pruvost,  however,  claimed 
in  his  original  description  to  have  studied  1 2  specimens.  The  only  substantially  complete 


235 


\ 


© 


Plate  9 


Figures  A-C.  Williamocalmania  vandergrachti  (Pruvost)  1922;  A)  NB  7183  Ech.  no.  3,  note  reduced  first 
thoracomere  (arrow),  x7.6;  B)  NB  7183  Ech.  no.  4,  showing  long,  subtriangular  telson,  x7;  Q  NB  7183 
Ech.  no.  2,  showing  somewhat  acute  abdominal  pleura,  x7.8. 

Figures  D  &  E.  Palaeorchestia  parallela  (Fritsch)  1876,  part  and  counterpart  of  lectotype,  CGH  593, 
showing  the  characteristic  parallel-sided,  distally  circular  telson;  D)  under  water  x3.6;  E)  x2.9. 


236 


Table  6. 

Measurements  of  lengths  in 

mm 

of  Williamocalmania  vandergrachti. 

*  Lectotype. 

Cephalon 

Thorax 

Abdomen 

A, 

Telson 

NB  7183  Ech 

1 

-1.2 

4.7 

5.6 

1.3 

2.3 

2 

6.4 

1.2 

*  2.3 

3 

0.8 

4.5 

4 

4.0 

0.9 

2.2 

specimen  now  available,  NP  7183  Ech.  no.  1,  clearly  is  the  basis  for  the  1922  and  1930 
figure  drawings,  and  thus  designated  here  as  the  lectotype.  Some  measurements  are 
provided  in  Table  6. 

BROOKSYNCARIS  new  genus 

Diagnosis.  —  First  thoracomere  only  slightly  reduced,  sixth  through  eighth  thorac- 
omeres  slightly  larger  than  second  through  fifth.  Thoracomeres  each  with  2  transverse 
grooves. 

Type  species.— Palaeocaris  canadensis  Brooks,  1962^7. 

Etymology.— '^2iVC\Qd.  in  honor  of  H.  K.  Brooks. 

Brooksyncaris  canadensis  (Brooks),  \962b 
Fig.  23;  Plate  10,  fig.  A 

v*1962^  Palaeocaris  canadensis  Brooks,  p.  248;  pi.  15,  figs.  1,  2. 
1969a  Palaeocaris  canadensis  Brooks.  Schram,  p.  220,  table  1. 

Diagnosis.  —  Since  there  is  but  one  species,  the  diagnosis  of  the  species  is  the  same 
as  that  of  the  genus. 

Holotype.  —  MCZ  5435  (Plate  10,  fig.  A);  Confluence  of  Diligent  and  Ramshead 
Rivers,  south  of  Diligent  River,  Cumberland  County,  Nova  Scotia;  Riversdale  Group, 
Westphalian  A. 

Description.  -Cephsdon  short  (cephalon  to  thorax  ratio  1:5.4);  prominent  cephalic 
groove;  small  postcephalic  groove  extending  in  arc  dorsad  from  posterior  margin. 


//     //      // 
'/      II      ,1 


n] 


Figure  23.     Diagrammatic  rendition  of  what  is  currently  known  about  the  form  oi  Brooksyncaris  canadensis, 
scale  5  mm. 


237 

Antennual  and  antennal  peduncles  well  developed  [too  poorly  preserved  to  discern 
details]. 

First  thoracomere  not  much  reduced.  Second  through  eighth  thoracopods  ambu- 
latory, with  large(?)  epipodites. 

Remarks.— The  specimens  discussed  by  Copeland  (1951  a,  b)  and  referred  by  Brooks 
(1962/))  to  this  species  have  been  reexamined  by  me.  I  concur  with  Copeland's  original 
treatment  of  that  material  and  refer  those  specimens  to  Palaeocaris  cf.  typus.  Thus,  the 
only  material  that  is  referable  to  B.  canadensis  is  the  holotype. 

Brooks  ( 1 962^)  described  the  thoracopods  of  this  species  as  bearing  epipodites.  I 
have  concurred  with  this  for  the  time  being,  but  it  is  difficult  to  clearly  discern  whether 
these  structures  are  epipodites  or  flap-like  exopods.  These  features  are  located  very 
close  to  the  base  of  the  limbs,  and  I  would  also  assume  that  the  exopods  probably 
would  have  been  as  poorly  preserved  as  the  endopods.  However,  the  question  remains 
open. 

The  new  genus  is  required  because  this  species  obviously  does  not  belong  in 
Palaeocaris  as  now  understood,  the  latter  taxon  being  characterized  in  part  by  the 
extreme  reduction  of  the  first  thoracomere.  However,  none  of  the  pertinent  features  of 
the  thoracopods,  abdomen,  pleopods,  or  tailfan  are  preserved  on  the  holotype  that 
would  allow  us  to  place  this  species  in  any  of  the  known  genera  of  palaeocaridaceans, 
let  alone  family.  Thus  the  establishment  of  a  separate  genus  seems  prudent. 

Genus  PALAEORCHESTIA  Zittel,  1885 

Diagnosis.— AnXennuXar  peduncles  smaller  than  those  of  antennae.  Telson  dis- 
tinctly rectangular. 

Type  species.  —  Gampsonychus  parallelus  Fritsch,  1876. 

Remarks.  — DQlerminrng  the  proper  name  for  this  genus  poses  a  classic  problem 
in  untying  the  twisted  strands  of  available  names  among  Paleozoic  syncarids.  Jordan 
(1847)  originally  described  Gampsonyx  fimbriatus,  unaware  of  the  fact  that  the  name 
of  this  genus  was  preoccupied  in  a  bird,  Gampsonyx  swainsoni  Vigors,  1825.  Bronn 
(1850)  did  detect  the  synonymy  and  suggested  the  name  Uronectes  be  applied  to  G. 
fimbriatus;  while  Burmeister  (1855)  independently  caught  the  same  synonymy  and, 
unaware  of  Bronn's  work,  suggested  the  use  of  the  name  Gampsonychus  for  G.  fim- 
briatus. Subsequent  authors,  until  Chappuis  (1927),  ignored  Bronn  and  used  either 
Gampsonyx  (and  thus  also  ignoring  the  synonym)  or  Gampsonychus. 

In  this  context  Fritsch  (1876)  described  a  new  species  Gampsonychus  parallelus 
and  allied  it  to  G.  fimbriatus.  Zittel  (1885)  recognized  the  distinctive  generic  status  of 
this  species  from  fimbriatus  and  erected  a  new  combination  Palaeorchestia  parallela. 
Brooks  (1969)  seemingly  concurred  with  the  Zittel  distinction  of/*,  parallelus  from 
what  by  then  was  known  as  U.  fimbriatus,  but  implied  that  the  proper  generic  assignment 
of  the  species  was  supposedly  with  Palaeocaris  when  he  synonymized  Palaeoorchestia 
{lapsus  calumni),  along  with  other  syncarid  genera,  with  Palaeocaris.  Brooks  was 
mistaken,  since  the  taxa  in  question  are  distinctly  different.  One  might  be  technically 
entitled  to  return  to  the  use  of  Gamsonychus,  since  the  use  of  Gampsonychus  in  Fritsch 
(1876)  is  not  affected  by  any  subjective  synonymy  in  Burmeister  (1855).  However,  to 
do  so  would  be  to:  1)  return  to  the  19th  and  early  20th  century  confusion  over  the  use 
of  Gamsonyx-Gampsonychus,  2)  minimize  the  importance  of  Zittel's  initial  recognition 
of  the  separate  generic  status  of  the  type  species,  and  3)  overlook  Brooks'  implicit 
acceptance  of  Zittel's  work.  For  these  reasons,  I  think  the  decisions  of  Brooks  andZhtel 
should  prevail  and  the  name  Palaeorchestia  be  used. 

Palaeorchestia  parallela  (Fritsch),  1876 
Fig.  24;  Plate  9,  figs.  D  &  E 

1859  cf.  Gampsonychus  fimbriatus.  Krejci,  p.  79. 

v*1876  Gampsonychus  parallelus  Fritsch,  p.  4,  pi.  3  fig.  1,  pi.  4. 

1885  Palaeorchestia  parallela  (Fritsch).  Zittel  p.  673,  fig.  858. 


238 


Figure  24.     Dorsal  reconstruction  of  what  is  currently  known  about  the  form  of  Palaeorchestia  parallela, 
scale  5  mm. 


7907  Palaeorchestia  parallela  (Fritsch).  Fritsch,  p.  73. 
7976  Palaeorchestia  parallela  (Fritsch).  Vanhoffen,  p.  146, 
7959  Palaeorchestia  parallela  (Fritsch).  Slewing,  p.  103. 
7965  Palaeorchestia  species  Noodt,  p.  82. 
7969  Palaeocaris  parallela  (Fritsch).  Brooks,  p.  R355. 


fig.  11. 


Diagnosis. —SmcQ  but  one  species  is  known,  the  diagnosis  of  the  species  is  the 
same  as  that  of  the  genus. 

Lectotype.—CGYi  593  (Plate  9,  figs.  D  &  E),  from  Lisek,  northwest  of  Beraun, 
Bohemia,  Czechoslovakia.  Rodnitz  Horizon,  Coal  Measures,  Lower  Permian.  [The 
stratigraphic  horizon  is  somewhat  vague  in  the  literature.  One  specimen  in  the  British 
Museum  (Natural  History),  In  35327,  which  resembles  the  preservation  seen  in  the 
Czech  types,  is  marked  "Carboniferous,  Nirzan,  near  Pilsen,  Bohemia— Old  Colin." 
seems  to  indicate  the  type  Lower  Permian  area  in  Czechoslovakia.] 

Description.  — ^ody  moderate  in  size.  Antennular  peduncle  with  3  subequal  seg- 
ments, medial  margin  of  second  segment  spinose,  flagella  well  developed.  Antennal 
protopod  with  short  proximal  segment,  scaphocerite  large  and  setose,  basal  joints  of 
flagellum  peduncular  with  distal  segment  twice  as  long  as  proximal.  No  rostrum. 

First  thoracomere  somewhat  shorter  than  any  other  body  segment.  Thorax  length 
more  than  one  and  one-half  times  that  of  abdomen. 

Sixth  pleomere  very  long,  with  faint  lateral  groove  about  mid-length.  Telson  long, 
subrectangular,  marginally  setose.  Uropodal  rami  spatulate,  with  strong  median  ribs, 
margins  finely  setose,  exopod  larger  than  endopod,  endopod  equal  to  or  shorter  than 
telson,  exopod  with  circular  diaeresis. 

Remarks.  — ¥v\Xsc\\  (1876,  1901)  referred  to  only  one  specimen.  CGH  593  is  ob- 
viously the  basis  of  Plate  4  in  his  1876  paper,  and  is  thus  designated  the  lectotype  here. 
However,  one  additional  specimen,  CGH  592,  is  in  the  collections  of  the  National 
Museum  in  Prague.  It  confirms  the  form  of  the  head  and  antennae  noted  on  the  type, 
and  clearly  reveals  the  slight  reduction  in  size  of  the  first  thoracomere. 

Aside  from  the  antennules,  antennae,  and  uropods,  virtually  nothing  is  known 
concerning  any  of  the  other  appendages.  CGH  593  preserves  some  remnants  of  anterior 
thoracopods.  Both  specimens  are  preserved  in  dorsal  view  in  a  brownish-gray  mudstone, 
and  details  on  the  thorax  and  anterior  abdomen  are  almost  totally  lacking.  The  addi- 
tional specimen  in  the  British  Museum,  In  35327,  contributes  nothing  towards  un- 
derstanding this  species. 


239 


^f 


Figure  25.     A)  Partial  reconstruction  of  Clarkecaris  brasilicus,  scale  5  mm;  B)  tailfan  to  same  scale  as  body; 
dorsal  views  of  C)  right  antennule  and  D)  antenna  slightly  enlarged. 


Genus  CLARKECARIS  Mezzalira,  1952 

Diagnosis.— C^phdiXon  with  well-developed  groove.  Eight  free  thoracic  segments 
short.  Abdominal  segments  long,  especially  the  sixth,  pleura  styliform.  Telson  subtrian- 
gular,  with  a  narrow  bifid  terminus. 

Type  species.  — Gampsonyx  brasilicus  Clarke,  1920. 

Remarks.— Tht  higher  taxonomic  placement  of  this  species  has  been  rather  per- 
ipatetic. It  was  originally  placed  among  the  "gamsonychids,"  but  when  Mezzalira  (1952) 
recognized  the  separate  generic  status  of  this  creature  he  assigned  it  to  the  Uronectidae. 
However,  Mezzalira's  understanding  of  "uronectids"  was  not  that  of  Brooks  (1962a). 
The  latter  placed  Clarkecaris  in  its  own  family  within  the  anaspidaceans;  though  later 
(Brooks  1969)  he  reassigned  it  to  the  stygocaridaceans,  still  within  its  own  family. 
Stygocaridines  have  since  been  recognized  as  a  group  within  the  Anaspidacea.  However, 
reexamination  of  the  types  and  an  additional  specimen  studied  and  illustrated  by  Brooks 
(1962a,  1969),  as  well  as  information  from  new  material  from  Brazil  (Dr.  Iraja  Domiani 
Pinto,  pers.  comm.),  reveals  that  there  are  8  free  subequal  thoracomeres,  placing  this 
species  within  the  Palaeocaridacea. 

Clarkecaris  brasilicus  (Clarke)  1920 
Fig.  25;  Plate  10,  figs.  B-D 

\.*1920  Gampsonyx  brasilicus.  Clarke,  p.  137,  pi.  3,  figs.  9,  10. 

1927  Uronectes  brasilicus  (Clarke).  Chappuis,  p.  605. 

1931  Uronectes  braziliensis  (Clarke).  Van  Straelen,  p.  18. 

1946  Gampsonyx  brasilicus  Clarke.  Mezzlira,  p.  118,  figs.  9,  10. 

1946  Uronectes  brasilicus  (Clarke).  Mezzalira,  p.  118. 

1948  Uronectes  brasilicus  (Clarke).  Mezzalira,  p.  250. 

1952  Clarkecaris  brasilicus  (Clarke).  Mezzalira,  p.  46,  pi.  3. 

1954  Clarkecaris  brasilicus  (Clarke).  Mezzalira,  p.  168. 

1959  Gampsonyx  brasilicus  Clarke.  Slewing,  p.  100. 

V.  1962a  Clarkecaris  brasilicus  (Clarke).  Brooks,  p.  231;  Fig.  2b;  pi.  5,  fig.  2. 

1962b  Clarkecaris  brasilicus  (Clarke).  Brooks,  p.  274. 

1969  Clarkecaris  brasilicus  (ClaTke).  Brooks,  p.  R358,  figs.  169-3,  174-2. 

1969  Clarkecaris  brasilicus  (Clarke).  Schram,  p.  221,  table  1. 

1971  Clarkecaris  brasilicus  (Clarke).  Mezzalira,  p.  319,  pi.  1,  fig.  1,  pi.  3,  figs.  1-6. 

1977  Clarkecaris  brasilicus  (Clarke).  Schram,  p.  370. 

1978  Clarkecaris  brasilicus  (Clarke).  Brito  and  de  Quadros,  p.  417,  fig.  3. 

1979  Clarkecaris  brasilicus  (Clarke).  Schram,  p.  170. 
1981  Clarkecaris  brasilicus  (Clarke).  Schram,  p.  130. 

Lectotype. -NYSM  9738  (Plate  10,  fig.  D);  near  Guare'i,  Sao  Paulo,  Brazil;  Irati 
Formation,  Permian. 


240 


Other  /oca//7/>5.  — Innumerable  localities  for  this  species  are  known  in  the  states  of 
Panama  and  Sao  Paulo,  in  Brazil.  These  are  summarized  in  Mezzalira  (1948,  1954) 
and  Brito  and  de  Quadros  (1978). 

Z)6'5<:/'//7//o«.— Cephalon  with  well-marked  groove  (Plate  10,  fig.  C).  Antennules 
large,  with  well-developed  peduncles  medially  serrate  and  setose.  Antennae  with  large, 
oval,  setose  scaphocerite.  Flagella  well  developed. 

Thoracomeres  short,  subequal  in  length,  anterior  margins  marked  with  row  of 
papillae,  pleura  apparently  rounded. 

Abdominal  segments  long  (Plate  10,  fig.  D),  sixth  longer  than  any  others.  Pleura 
various:  first  rounded,  second  through  fifth  with  styliform  posteriorly  directed  processes. 
Uropodal  protopod  with  lateral  styliform  processes;  exopod  long  and  thin  (possibly  a 
straight  diaeresis);  endopod  diaphanous,  long,  and  oval.  Telson  shorter  than  uropod 
rami,  narrow,  subtriangular,  terminus  developed  as  bifed  process  (Plate  10,  fig.  B). 

Remarks.— T\iQ  description  here  is  based  on  observations  derived  from  study  of 
the  types  (NYSM  9738,  9739),  USNM  1 12766,  and  the  published  figure  in  Brito  and 
de  Quadros  (1978).  Considerable  more  information  should  become  available,  however, 
as  the  collections  available  to  Dr.  Damiani  Pinto  of  the  Instituto  de  Geosciencias,  Porto 
Alegre,  Brazil,  are  eventually  studied  and  described.  These  should  allow  a  definitive 
placement  of  C.  brasilicus  within  the  palaeocaridacean  families. 

Suborder  ANASPIDACEA  Caiman,  1904 

Infraorder  ANASPIDINEA  Caiman,  1904 

Family  ANASPIDIDAE  Thompson,  1894 

Genus  ANASPIDITES  Brooks,  1962a 

Diagnosis.  — 'R.osXnxm.  broad.  Thoracomeres  relatively  short  compared  to  anterior 
pleomeres.  Telson  subtriangular,  distally  pointed. 

Type  species.— Anaspides  antiquus  Chilton,  1929. 

Remarks.  — ThQ  initial  observations  on  this  taxon  (Chilton  1929,  Brooks  1962a) 
were  largely  based  on  one  incompletely  preserved  specimen.  A  search  in  1980  of  the 
reserve  collections  of  the  Australian  Museum  uncovered  one  additional  specimen  (F 
25226),  which  preserves  the  abdomen  and  parts  of  the  tailfan.  In  addition,  two  spec- 
imens were  found  in  the  British  Museum  (Natural  History).  One  of  these  (In  46114) 
is  the  finest  example  of  this  species  known,  revealing  considerable  details  about  ap- 
pendage anatomy.  The  other  (In  46056)  is  of  the  uropodal  exopods.  Consequently,  a 
redescription  of  the  species  and  new  reconstruction  are  presented  here. 

Anaspidites  antiquus  (Chilton)  1929 
Fig.  26;  Plate  10,  figs.  E&F 

v.*  1929  Anaspides?  antiquus  Chilton,  p.  366,  pi.  30. 

1962a  Anaspidites  antiquus  (Chilton).  Brooks,  p.  234;  pi.  5,  fig.  1;  figs.  1  &  2c. 

1962b  Anaspidites.  species  Brooks.  Brooks,  pp.  267,  274. 

1969  Anaspidites  antiquus  (Chilton).  Brooks,  p.  R356,  figs.  169-4, 

J  982  Anaspidites  antiquus  (Chilton).  Schram,  p.  122. 

Diagnosis. —  Since  there  is  but  one  species  known,  the  diagnosis  of  the  species  is 
the  same  as  that  of  the  genus. 

Holotype.  —  \JS  7903.  Brookvale  Brick  Quarry,  New  South  Wales.  Hawksbury 
Sandstone,  Triassic. 

Description.— Cephalon  with  broad  rostrum,  prominent  cervical  groove.  Anten- 
nular  peduncles  large,  with  3  subequal  segments,  flagella  well  developed  [but  length 
indeterminate  because  of  lack  of  preservation].  Antennal  peduncles  with  4  (?)  segments 
(US  7903).  Mandibles  large,  massive. 

Thoracomeres  somewhat  shortened,  almost  one-half  the  length  of  anterior  pleo- 
meres (US  7903,  In  461 14).  Pleura  somewhat  rounded  (In  461 14).  Thoracopods  (Plate 
10,  fig.  F)  with  short  coxae,  bases,  and  ischia,  long  meri  (In  461 14);  beyond  knee,  long 


174-1. 


241 


Plate  10 

Figure  A.  Brooksyncaris  canadensis  (Brooks)  1962,  closeup  of  anterior  end  with  cephalon  (c)  and  first  4 
thoracomeres  (numbered),  xg.Q. 

Figures  B-D.  Clarkecaris  brasilicus  (Clark),  1920;  B)  paralectotype,  NYSM  9739,  showing  telson  with 
pointed  tip  (t)  and  styliform  uropods  (u),  x7;  C)  USNM  1 12766,  closeup  of  anterior  end,  note  cephalon, 
size  of  thoracomeres  (numbered)  and  anterior  pleomeres  (numbered),  x  4.5;  D)  NYSM  9738,  lectotype,  with 
posterior  thorax  and  abdomen,  x  2.8. 

Figures  E  &  F.  Anaspidites  antiquus  (Chilton)  1929;  E)  F  25226,  with  abdomen  and  telson,  x  1.8;  F)  In 
461 14,  whole  body,  note  annulate  pleopods  (arrows),  x  1.6. 


242 


^ 


Figure  26.    Reconstruction  of  Anaspidites  antiquus,  scale  5  mm. 


carpi  and  propodi  (US  7903,  In  46114),  dactyli  incompletely  preserved  (In  46114). 
Neither  thoracic  epipodites  nor  exopods  preserved. 

Pleomeres  undecorated.  Sixth  pleomere  length  twice  that  of  any  anterior  to  it. 
Pleopods  long,  uniramous,  annulate  (In  46114,  Plate  10,  fig.  F),  protopods  well  de- 
veloped. Telson  long,  subtriangular,  distally  pointed  (F  25226,  Plate  10,  fig.  E)  (perhaps 
some  faint  indication  that  terminus  possibly  flanked  by  set  of  small  furcae).  Uropodal 
protopod  simple,  well  developed  (In  46056,  In  461 14).  Exopod  blade-like  (F  25226, 
In  461 14),  reinforced  with  thick  struts  along  lateral  and  medial  margins  (In  46056). 

Remarks.  — brooks  (1962^)  interpreted  a  2-segment  protopod  on  the  antennae 
(with  only  the  distal  segment  visible),  a  straight-edged  scaphocerite,  and  the  3  most 
proximal  joints  of  the  flagellum  as  peduncular.  I  found  no  evidence  for  an  antennal 
scale  on  either  US  7903  or  In  461 14.  Brooks  also  felt  that  the  thoracopods  were  widely 
spaced,  on  opposite  ends  of  well-developed  thoracic  stemites.  Close  examination  of 
US  7903  indicates  that  the  supposed  foramina  of  the  thoracopods  are  more  likely 
preservational  anomalies  of  the  cuticular  wrinkles  or  possibly  ridges  on  the  anterior 
thoracomeres.  Finally,  the  pleopod  that  Brooks  noted  is  in  fact  part  of  a  posterior 
thoracopod. 

Although  the  general  mode  of  preservation  of  these  fossils  obscures  much  of  the 
detail,  enough  can  be  discerned  to  be  reasonably  certain  A.  antiquus  is  an  anaspid.  The 
body  is  large  and  well  developed,  but  the  first  thoracomere  is  fused  into  the  cephalon, 
and  the  rami  of  the  pleopods  are  clearly  uniramous  and  annulate.  However,  the  narrow 
thoracic  somites  and  styliform  telson  clearly  separate  this  Triassic  species  from  the 
living  forms  found  today  in  Tasmania.  Unfortunately,  the  diagnostic  features  of  the 
mouthparts  are  not  visible  on  any  of  the  available  material  of  Anaspidites,  and  as  a 
result  exact  assurance  as  to  family  affinities  within  the  Anaspidacea  must  remain  un- 
certain. 


Acknowledgments 

This  research  was  carried  out  under  NSF  grant  GB  79-03602,  which  allowed  me 
to  examine  specimens  in  museums  in  Australia,  Europe,  and  North  America,  and  to 
study  and  collect  living  syncarids  in  the  wilds  of  Tasmania.  The  following  individuals 
were  of  direct  help  on  various  aspects  of  this  study:  Drs.  H.-E.  Griiner,  Museum  fur 
Naturkunde,  Humboldt  Universitat,  Berlin;  S.  Morris,  British  Museum  (Natural  His- 
tory); S.  Secretan,  Institut  de  Paleontologie,  Paris;  G.  Pacaud,  Museum  d'Histoire 
Naturelle,  Autun;  R.  Prokop,  Narodni  Museum,  Prague;  A.  Richardson  and  R.  Swain, 
University  of  Tasmania,  Hobart;  H.  K.  Schminke,  Universitat  Oldenburg;  G.  Ubaughs, 
Laboratoire  de  Paleontologie  Animale,  Universite  de  Liege;  H.  van  Amerom,  Rijks 


243 


Geologische  Dienst,  Heerlen;  and  R.  Wilson,  Institute  of  Geological  Sciences,  Edin- 
burgh. Photographic  and  technical  assistance  was  rendered  by  Messrs.  B.  R.  Burnett, 
R.  M.  Chandler,  and  T.  A.  Demere,  San  Diego  Museum;  and  the  reconstructions  were 
drawn  by  Mr.  M.  J.  Emerson. 

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TRANSACTIONS 
OF  THE  SAN  DIEGO 
SOCIETY  OF 
NATURAL  HISTORY 


Volume  20  Number  14  pp.  247-276        20  November  1984 


The  Late  Wisconsinan  Vertebrate  Fauna  from  Deadman  Cave, 
Southern  Arizona  ,^ 


Jim  I.  Mead 

Center  for  the  Study  of  Early  Man.  Institute  for  Quaternary  Studies, 
University  of  Maine,  Orono,  Maine  04469 


NOVg 


Edward  L.  Roth  J^^J^^fO 

Department  of  Biology,  Howard  Payne  University,  '^'^iMi/Tv, 

Brownwood,  Texas  76801  *Y 

Thomas  R.  Van  Devender 

Arizona- Sonora  Desert  Museum,  Route  9,  Box  900, 
Tucson,  Arizona  85743 

David  W.  Steadman 

Division  of  Birds,  Smithsonian  Institution, 
Washington,  DC.  20560 

Abstract.  We  report  a  particularly  rich  assemblage  of  fossil  vertebrates  from  a  cave  in  southern 
Arizona.  This  fauna  provides  new  data  for  reconstructing  the  inadequately  known  Late  Pleistocene- 
Early  Holocene  biota  of  the  Sonoran  Desert  and  nearby  mountains.  The  vertebrate  fauna  of  Deadman 
Cave  includes  5  amphibians,  25  reptiles  (13  lizards  and  12  snakes),  12  birds,  and  22  mammals  for  a 
total  of  64  species.  Only  one  amphibian  {Bufo  woodhousei),  three  reptiles  (Callisaurus  draconoides, 
Phrynosoma  modestum,  Gyalopium  canum),  and  one  mammal  {Microtus  species)  are  locally  extirpated, 
although  all  still  occur  in  southern  Arizona.  An  unidentified  icterine  bird  may  prove  to  be  an  extinct 
species.  Extinct  mammals  include  Euceratherium  collinum,  Equus  species,  and  Nothrotheriops  shasten- 
sis,  all  large  herbivores.  Other  than  the  extinct  animals,  the  fauna  dating  of  the  Late  Pleistocene  and 
Early  Holocene  is  little  different  from  that  which  is  available  in  southern  Arizona  today.  What  appears 
to  have  changed  is  the  mosaic  of  the  plant  and  animal  community.  Distinctly  boreal  animals  are  lacking 
from  the  fauna.  The  climate  during  the  time  of  deposition  of  the  cave  sediment  appears  to  have  been 
equable;  certain  animals  now  confined  to  deserts  were  able  to  live  in  more  diverse  woodland  com- 
munities. 

Introduction 

Deadman  Cave  is  a  medium-sized,  limestone  cave  at  1400  m  (4600  ft)  elevation 
on  the  northeastern  side  of  the  Santa  Catalina  Mountains,  Pima  County,  Arizona  (Fig. 
1).  The  cave  is  located  in  the  lower  portion  of  the  mountain  range  where  various 
Paleozoic  limestone  formations  are  exposed  and  it  appears  to  be  formed  in  the  Mis- 
sissippian  Escabrosa  Formation  (Wallace  1955). 

The  present  vegetation  of  this  highly  dissected  area  is  desert-grassland  intermixed 
with  the  lower  boundary  of  the  oak  woodland  {Quercus  species);  juniper  {Juniperus 
erythrocarpa  and  J.  deppeana)  is  thinly  scattered  (Whittaker  and  Niering  1968).  Desert- 
grassland  elements  such  as  agave  (Agave parryi'),  ocotillo  {Fouqieria  sp/endens),  variable 
prickly  pear  (Opuntia  phaeacantha),  and  assorted  grasses  occur  on  the  limestone  and 
conglomerate  hillslopes.  Tributaries  between  the  hills  and  the  areas  of  the  lower  hill- 
slopes  are  covered  with  velvet  mesquite  {Prosopis  velutina),  netleaf  hackberry  (Celt is 


248 


CONTOUR      LINES      •     S500     <l      ELEVATION 


Figure  1 .  Map  of  southern  Arizona  with  the  locations  of  Deadman,  Papago  Springs,  and  Ventana  caves. 
Contour  lines  (1650  m,  5500  ft)  denote  the  positions  of  large  mountain  ranges,  illustrating  that  southeastern 
Arizona  is  mountainous  with  great  expanses  of  woodland  and  boreal  habitats.  The  southwestern  portion  of 
the  state  contains  predominately  small  desert  mountains. 


reticulata),  and  catclaw  {Acacia  greggii).  The  area  around  Deadman  Cave  is  presently 
the  ecotone  between  the  creosotebush  desertscrub  communities  of  the  San  Pedro  Valley 
(715m  elevation  at  the  town  of  Mammoth)  and  the  oak  woodland  above.  Between 
Deadman  Cave  and  the  top  of  the  Santa  Catalina  Mountains  (2790  m  elevation)  there 
are  Mexican  pine-oak  woodland,  and  ponderosa  pine  and  mixed  conifer  forests. 

The  approximately  three-by-five  meter  entrance  to  Deadman  Cave  is  a  collapsed 
cavern  ceiling  (Fig.  2).  The  cave  once  contained  an  elaborate  system  of  active  speleo- 
thems.  Travertine  building  is  now  very  rare  to  absent,  possibly  because  of  the  devel- 
opment of  the  present  entrance  and  resultant  dessication;  rimstone  pools  rarely  contain 
any  water  (William  Peachey  1980,  personal  communication). 

During  the  late  1800s  miners  entered  the  cave  to  explore  the  numerous  passages. 
A  cabin  was  constructed  across  the  entrance  to  the  cave.  Possibly  at  the  same  time,  a 
shaft  was  begun  in  a  back  portion  of  the  cave.  This  shaft  penetrated  a  travertine  surface 
layer  and  2.4  m  of  cemented  rubble  containing  bones,  and  provided  access  to  a  lower, 


HAFT 


BONE 
DEPOSIT 


m 


Figure  2.  Generalized  cross  section  of  Deadman  Cave,  southern  Arizona.  The  present  entrance  appears 
to  have  been  the  opening  during  the  Late  Pleistocene.  A  shaft  built  by  miners  during  the  1800s  cuts  through 
two  travertine  layers  and  provides  access  to  a  lower  room  and  the  bone  deposit,  sealed  off  from  the  rest  of 
the  cave  since  approximately  8000  B.P. 


249 


sealed-off,  small  room.  Here  a  second  travertine  layer  covered  a  fine  carbonate  silt.  At 
this  point  a  trench  was  excavated  horizontally  through  this  loose  sedimentary  layer 
and  then  all  mining  operations  ceased,  leaving  the  exposed  shaft  and  trench  walls. 

William  Peachey  informed  us  of  the  exposed  sediments  in  Deadman  Cave.  One 
of  us  (ELR)  and  W.  Peachey,  entered  the  cave  in  1972  and  excavated  an  approximately 
1.0  by  0.5  by  0.5  m  layer  of  loose  sediment  from  below  the  travertine  layer  capping 
the  material  exposed  in  the  trench  wall.  All  excavated  sediments  were  screened  through 
window  mesh  (2  mm)  sieves.  Most  of  the  fossils  were  identified  using  the  comparative 
collections  at  the  University  of  Arizona,  Tucson  and  the  Division  of  Birds,  Smithsonian 
Institution,  Washington,  D.C. 

Chronology 

The  faunal  assemblage  contains  three  extinct  species:  the  Shrub  ox  {Euceratherium 
collinum),  the  Shasta  ground  sloth  {Nothrotheriops  shastensis),  and  the  Horse  {Equus 
species).  The  remainder  of  the  fauna  can  be  found  today  living  in  various  habitats  in 
southern  Arizona.  Euceratherium,  Nothrotheriops,  and  Equus  apparently  were  extinct 
along  with  many  other  large  mammals  by  approximately  1 1  000  to  10  200  B.P.  (years 
before  present;  Martin  1967;  Mosimann  and  Martin  1975;  Haynes  1968;  Meltzer  and 
Mead  1983).  The  youngest  known  radiocarbon  date  on  dung  of  A^.  shastensis  is  about 
10  500  B.P.  (Thompson  et  al.  1980).  These  dates  suggest  that  at  least  some  of  the 
Deadman  Cave  faunal  assemblage  is  of  Late  Wisconsinan  age. 

A  radiocarbon  date  of  6080  ±  250  B.P.  (A-1617)  was  determined  on  14.8  grams 
of  endocarps  of  Cehis  reticulata  found  directly  associated  with  the  fauna.  Unfortunately 
there  was  insufficient  CO2  available  for  a  '^C  correction.  Cehis  endocarps  are  notorious 
for  containing  very  little  carbon  and  for  being  easily  contaminated  by  carbonates  in 
percolating  water.  The  sediments  had  been  leached  of  all  organics;  some  of  the  bones 
were  encrusted  with  carbonates.  For  this  reason  we  believe  that  the  ''^C  age  of  6000 
B.P.  is  probably  too  young  by  at  least  2000  years.  The  Deadman  Cave  faunal  assemblage 
presented  here  most  likely  dates  between  12  000  and  8000  B.P.,  grading  across  the  Late 
Wisconsinan-Early  Holocene  boundary.  Because  of  the  uncertainties  of  the  dating  it  is 
not  possible  to  establish  unequivocally  whether  all  the  reported  taxa  lived  contem- 
poraneously in  the  local  community.  The  thick  travertine  cap  and  the  sealing  off' of  the 
lower  room  suggests  that  the  deposit  has  not  been  contaminated  with  Middle  or  Late 
Holocene  bones. 

The  Fossil  Deposit 

The  fossiliferous  layer  is  a  pebbly  silt  (very  pale  brown,  7/3  10  YR  dry,  Munsell 
color)  and  shows  no  physical  indication  that  the  sediments  were  deposited  by  flowing 
water.  The  bones  may  have  accumulated  by  a  number  of  mechanisms.  Bassariscus 
astutus  (Ringtail)  is  a  small  carnivore  that  inhabits  the  cave  today;  small  pockets  of 
bones  and  seeds  are  presently  developing  as  the  scats  of  the  Ringtail  decay.  Some  of 
the  fossil  bone  deposit  in  Deadman  Cave  may  have  been  developed  by  the  Ringtail  as 
it  was  2000  B.P.  in  Vulture  Cave,  Grand  Canyon,  Arizona  (Mead  and  Van  Devender 
1981).  Spilogale  putorius  (Spotted  skunk)  is  the  most  common  carnivore  in  the  fossil 
deposit  and  it  too  may  have  helped  in  the  accumulation  of  animal  remains. 

Owls  {Otus  species— Screech  owl;  Micrathene  whitneyi—E\{ o-wX;  and  Asio  otus— 
Long-eared  owl)  were  also  recovered  from  the  Deadman  Cave  deposit.  The  Long-eared 
owl  is  known  to  prey  upon  Spotted  skunks  and  is  the  only  owl  that  inhabits  deep 
recesses  in  caves,  that  was  recovered  from  the  deposit. 

The  Ringtail  and  the  Long-eared  owl  could  very  well  account  for  the  entire  fossil 
deposit  except  for  the  larger  mammal  remains.  The  cave  also  may  have  been  a  den 
and  a  food  cache  for  Felis  concolor  (Mountain  lion)  or  other  large  carnivores,  which 
could  account  for  the  occasional  fossil  remains  of  the  Mountain  lion.  Horse,  Ground 
sloth,  and  Shrub  ox. 

The  area  of  the  bone  deposit  (Fig.  2)  is  just  beyond  view  of  the  light  coming  in 


250 


from  the  present  and  the  presumed  Late  Wisconsinan-aged  cave  entrance.  From  here 
to  the  area  of  the  fossil  deposit,  it  is  an  easy  passage  for  an  owl  or  a  mammalian 
carnivore,  across  an  open,  large  cavern  room.  The  predators  may  have  had  easy  access 
to  the  lower  room  where  the  fossil  bone  deposit  occurs,  but  the  passage  has  since  filled 
in  with  travertine,  bones,  and  rock  rubble. 

Thus,  the  Deadman  Cave  fossil  deposit  is  probably  time  transgressive  by  possibly 
a  few  thousand  years,  and  appears  to  be  an  in  situ  deposit  from  the  predator  accu- 
mulation(s)  dating  some  8000  to  12  000  years  ago. 

Results 
Fauna 

The  vertebrate  fauna  recovered  from  Deadman  Cave  (University  of  Arizona,  Lab- 
oratory of  Paleontology,  UALP,  locality  78121),  is  represented  by  5  amphibians,  25 
reptiles  (13  lizards  and  12  snakes),  12  birds,  and  22  mammals,  for  a  total  of  64  species 
(Table  1). 

The  following  is  a  systematic  account  of  the  fauna.  After  the  skeletal  element  is 
the  number  of  specimens,  if  more  than  one,  and  the  UALP  specimen  number.  "R" 
and  "L"  refer  to  right  and  left  respectively.  A  brief  discussion  of  identifying  criteria 
and  present-past  distributions  within  the  southwestern  United  States  and  northern 
Mexico  is  included.  Our  taxonomic  sequences,  distributional  data,  and  descriptive 
osteological  nomenclature  is  as  follows:  amphibians  and  reptiles— Stebbins  (1966); 
birds— Baumel  et  al.  (1979),  Phillips  et  al.  (1964);  and  mammals— Jones  et  al.  (1982) 
for  extant  species  and  Kurten  and  Anderson  (1980)  for  extinct  species.  Amphibians 
and  reptiles  were  identified  by  TRVD  and  JIM,  birds  by  DWS,  and  mammals  by  ELR 
and  JIM. 

Class  AMPHIBIA— Amphibians 

Order  ANURA  —Toads  and  frogs 

Family  Pelobatidae— Spadefoot  toads 

Scaphiopus  species— Spadefoot  toad 

Material  — Kdidio-uXna.  (11411). 

Scaphiopus  cowc/z/- Couch's  spadefoot  toad 

Ma/ma/.- Vertebra  (1 1409);  R  ilium  (15312). 

Remarks.  —  Skeletal  elements  from  adult  individuals  of  Scaphiopus  couchi  and  S. 
hammondi  are  distinctive.  The  shape  of  the  ilium,  including  the  Ala  ossa  ilei  and  the 
Margo  dorsalis,  identify  the  fossil  specimen  as  belonging  to  a  spadefoot  toad.  Scaphiopus 
couchi  may  be  distinguished  from  5.  hammondi  (Western  spadefoot  toad)  by  the  fol- 
lowing characters:  1)  the  area  from  the  spina  pelvis  anterior  to  the  acetabulum  is 
relatively  flat  on  S.  hammondi  but  is  raised  at  the  acetabulum  on  S.  couchi;  2)  the 
articular  surface  at  the  acetabulum  is  curved  on  S.  hammondi,  but  straight  on  5'.  couchi; 
and  3)  the  shape  of  the  Spina  pelvis  posterior  is  angular  on  S.  hammondi,  but  curved 
on  5".  couchi.  S.  couchi  lives  in  a  wide  variety  of  desertscrub,  grassland,  and  subtropical 
habitats  including  the  Santa  Catalina  Mountains. 

Amphibian  subfossil  and  fossil  remains  are  very  inadequately  represented  in  Ar- 
izona (Van  Devender  and  Mead  1978,  Mead  1981).  Besides  Deadman  Cave,  S.  couchi 
is  known  from  an  Early  Holocene  age  wood  rat  midden  in  Arizona  (Van  Devender 
and  Mead  1978),  Late  Wisconsinan  and  Holocene  cave  deposits  in  southwestern  New 
Mexico  (Van  Devender  and  Worthington  1977,  Holman  1970,  Brattstrom  1964),  and 
from  Rancho  la  Brisca,  Sonora,  Mexico  (Van  Devender  et  al.  in  press). 

Scaphiopus  cf.  S.  hammondi— WestQm  spadefoot  toad 
Material.  —  Vertebra  (11410). 


251 


Table  1 .  Late  Pleistocene,  Holocene  and  present  fauna  from  Deadman  Cave  and  other  localities  in  southern 
Arizona.  Sequence  and  nomenclature  is  as  follows  (exceptions— see  text):  the  amphibians  and  reptiles 
follow  Stebbins  (1966),  the  birds  follow  Phillips  et  al.  (1964),  and  the  mammals  follow  Jones  et  al.  (1982) 
for  the  extant  species  and  Kurten  and  Anderson  (1980)  for  the  extinct  species.  1  =  Van  Devender  and 
Mead  (1978);  Van  Devender  (1973);  Mead  et  al.  (1983).  2  =  Haury  (1950).  3  =  Skinner  (1942)  and  Rea 
(1980).  *  =  occurs  in  category.  !  =  extinct  species.  S.C.Mt.  =  Santa  Catalina  Mountains.  ?  =  Questionable 
association. 


Present  fauna 


Wood  rat  middens' 


Ventana  Cave 
<  10,000  B.P.2 


S.C.Mt. 

or  nearby  Southern  Deadman  <  10,000  >  10,000  Volcanic  Conglom- 
valley      Arizona       Cave  B.P.  B.P.         unit  erate 


Papago 
Springs 
Cave' 
Late 
Pleisto- 
cene 


AMPHIBIANS 

Scaphiopus  couchi 
S.  cf  5'.  hammondi 
Bufo  cf  B.  wood- 
housei 

B.  punctatus 
Rana  sp. 

REPTILES 

Gopherus  agassizi 
Coleonyx  variegatus 
Sauromalus  obesus 
Holbrookia  maculata 
H.  texana 

Callisaurus  draconoides 
Crotaphytus  collaris 

C.  wislizeni 
Sceloporus  cf.  5". 

magister 
S.  cf.  5'.  clarkii 
S.  cf.  5".  undulatus 
Uta  stansburiana 
Urosaurus  cf  U. 

graciosus 
U.  ornatus 

Phrynosoma  douglassi 
P.  modestum 
P.  solare 
Cnemidophorus  cf. 

C  tigris 
Cnemidophorus  sp. 
Heloderma  suspectum 
Lichanura  trivirgata 
Phyllorhynchus  decur- 

tatus 
Masticophis  sp. 
Salvadora  sp. 
Arizona  elegans 
Pituophis  melano- 

leucus 
Lampropeltis  getulus 
L.  pyromelana 
Rhinocheilus  lecontei 
Sonora  semiamdata 
Chionactis  occipitalis 
Gyalopium  canum 
Trimorphodon  biscu- 

tatus 
Hysiglena  torquata 
Crotalus  atrox 
C.  cerastes 
C.  scutulatus 


* 
* 

* 
* 


7 
? 

* 


* 


* 
* 
* 

« 

* 

He 
* 

* 
* 

* 
* 

* 
* 

« 


252 


Table  1.     Continued. 


Papago 
Present  fauna  Ventana  Cave        Spnngs 

— — — Wood  rat  middens'      <  10,000  B.P.=  *       ,     ^ 

S.C.Mt. : Late 

or  nearby  Southern  Deadman  <  10,000   >  10,000  Volcanic  Conglom-  Pleisto- 

valley      Arizona       Cave  B.P.  B.P.         unit  erate  cene 

BIRDS 

Ibis-like  * 

Colinus  gambelii 
Colinus  sp. 

Cyrtonyx  montezumae 
Meleagris  crassipes 
Zenaida  cf.  Z. 

macroura 
Olus  sp. 

Micrathene  whitneyi 
Asio  otus 
Caprimulgidae 
Colaptes  auratus 
Turdus  cf.  T.  migra- 

torius 
Catharus  guttatus 
Icterinae  (probably 

extinct  species) 
Emberizinae  »  *  * 

MAMMALS 


*  *  « 

if  *  * 

itC  *  3tt 


*  *  * 

*  *  * 

*  *  * 

*  m  * 

«  He  * 

*  *  « 

*  *  * 

*  *  * 


Notiosorex  crawfordi 
Myotis  cf  M.  velifer 
M.  cf  M.  evotis 
M.  cf  M.  thysanodes 
cf  Myotis 
Plecotus  cf  P.  rafin- 

esquii 
Antrozous  pallidas 
Tadarida  cf.  T.  bra- 

siliensis 
\Nothrotheriops  shas- 

tensis 
Sylvilagus  auduboni 
Sylvilagus  sp. 
L.  californicus 
Lepus  sp. 
Eutamias  dorsalis 
Eutamias  sp. 
Marmota  flaviventris 
cf  Ammospermophilus 
Spermophilus  varie- 

gatus 
S.  tereticaudus 
S.  lateralis 

Cynomys  ludovicianus 
Thomomys  bottae  or 

umbrinus 
T.  cf  T.  bottae 
Perognathus  cf  P. 

flavescens 
P.  cf  P.  flavus 
P.  baileyi 
Perognathus  sp. 
Dipodomys  spectabilis 
D.  cf  D.  deserti 
D.  merriami 
Reithrodontomys  mon- 

tanus 


***** 

*  *  * 

*  *  * 

* 


*  *  * 

«  *  *  *  * 


*  * 

*  * 

*  * 

*  * 

*  * 

*  * 


*  * 

*  * 


*  * 

*  * 


*  *  * 

*  *  * 

*  *  * 

*  *  * 

*  * 

*  *  * 


Table  1.     Continued. 


253 


Present  fauna 
S.C.Mt. 


Ventana  Cave 
Wood  rat  middens'       <  10,000  B.P.^ 


Papago 

Springs 

Cave' 

Late 


or  nearby  Southern  Deadman  <  10,000   >  10,000  Volcanic  Conglom-  Pleisto- 
valley      Arizona       Cave  B.P.  B.P.         unit  erate  cene 


Reithrodontomys  sp. 

« 

* 

Peromyscus  manicu- 

latus 

* 

* 

P.  boylii  or  trueP 

*/ 

*/* 

Peromyscus  sp. 

* 

* 

* 

Onychomys  torridus 

* 

* 

O.  leucogaster 

4> 

Sigmodon  cf.  S.  ari- 

zonae 

* 

* 

* 

S.  ochrognathus 

* 

Neotoma  albigula 

* 

* 

* 

N.  lepida 

* 

N.  mexicana  or  albi- 

gula^ 

*/* 

*/* 

Microtus  sp. 

* 

« 

M.  cf.  M.  mexicana 

* 

Erethizon  dorsatum 

« 

* 

Canis  latrans 

* 

* 

C.  lupus 

7 

!C.  dirus 

Vulpes  macrotis 

* 

* 

Urocyon  cinereoar- 

genteus 

* 

* 

Ursus  americanus 

* 

* 

Bassariscus  astutus 

* 

* 

* 

\B.  sonoitensis 

Taxidea  taxus 

* 

* 

Spilogale  putorius 

* 

m 

* 

Mephitis  mephitis 

* 

* 

M.  macroura 

* 

* 

* 

Felis  concolor 

* 

* 

* 

\Panthera  leo  atrox 

\Equus  tau 

\E.  occidentalis 

\E.  conversidens 

\Equus  sp. 

* 

!  Tapirus  sp. 

\Platygonus  comp- 

ressus 

Dicotyles  (=Tay- 

assu)  sp. 

* 

m 

\Camelops  sp. 

Odocoileus  sp. 

* 

« 

* 

Cen'us  sp. 

IStockoceros  cf.  S. 

conklingi 

\S.  onusrosagris 

\Euceratherium  col- 

linum 

* 

Bison  bison 

Bison  sp. 

* 
* 

* 

* 


* 


Remarks.  — The  shape  of  centrum  and  the  size  of  the  fossil  vertebra  was  indistin- 
guishable with  that  of  the  Western  spadefoot  toad.  S.  hammondi  is  widely  distributed 
in  southeastern  Arizona  where  it  lives  in  desert,  grassland,  chaparral,  woodland,  and 
pine  forest  habitats  (Lowe  1964).  The  fossil  referred  to  as  cf.  S.  hammondi  represents 


254 


the  first  fossil  occurrence  for  the  species  in  Arizona.  Late  Pleistocene  and/or  Early 
Holocene  age  occurrences  outside  Arizona  for  this  species  are  known  from  Nevada 
(Brattstrom  1976)  and  New  Mexico  (Holman  1970). 

Family  Bufonidae— Toads  ♦ 

Bufo  cf.  B.  woodhousei—V\l oodhoxxst's,  toad 

Material. -Y^nthrdi  (1 1408). 

Remarks.— Bufo  woodehousei  is  a  large  toad  in  relation  to  the  other  species  found 
today  in  Arizona,  although  distinctly  smaller  than  B.  alvarius  (Colorado  River  toad). 
Bufo  woodhousei  can  be  identified  by:  1)  the  size  is  relatively  larger  at  all  stages  of 
growth  than  most  other  Bufo,  2)  the  neural  arch  is  higher  making  the  centrum  more 
pronounced,  and  3)  the  articular  facets  are  larger.  Woodhouse's  toad  occurs  in  eastern 
and  central  Arizona  and  in  isolated  populations  in  the  Yuma  area  of  southwestern 
Arizona.  In  southern  Arizona  B.  w.  australis  is  primarily  a  riparian  species  restricted 
to  permanent  or  semi-permanent  streams.  Late  Pleistocene  or  Early  Holocene  age 
remains  of  Bufo  woodhousei  have  not  been  previously  reported  from  Arizona.  Outside 
Arizona,  fossil  remains  of  this  toad  are  known  from  Nevada  (Mead  et  al.  1982)  and 
New  Mexico  (Holman  1970). 

Bufo  punctatus—Ktd-spoUQd.  toad 

Material.  - Urostyle  ( 1 5002). 

Remarks.— Bufo  punctatus  is  a  small  toad  with  many  easily  identifiable  skeletal 
elements.  The  paired  anterior  condyles  of  the  urostyle  are  relatively  broad,  flat  ovals 
as  in  B.  punctatus.  Juveniles  of  the  larger  species  of  Bufo,  do  not  have  as  flattened  an 
anterior  end  to  the  urostyle.  This  xeric-adapted  toad  occurs  throughout  most  of  the 
Southwest,  living  in  habitats  ranging  from  desertscrub  to  Mexican  pine-oak  woodland. 
Late  Pleistocene  remains  of  B.  punctatus  occur  in  three  Arizona  localities  (Van  De- 
vender  and  Mead  1978),  New  Mexico  (Holman  1970,  Van  Devender  and  Worthington 
1977),  and  Rancho  la  Brisca,  Sonora,  Mexico  (Van  Devender  et  al.  in  press). 

Family  Ranidae— Frogs 
Rana  species— Frog 

Material  — WnmcYViS  (11412). 

Remarks.  — T\\Q  long,  slender  humerus  is  identifiable  to  Rana,  but  we  were  only 
able  to  identify  the  fossil  to  a  small  species.  Rana  pipiens  (Leopard  frog)  and  R. 
tarahumarae  (Tarahumara  frog)  along  with  the  introduced  R.  catesbeiana  (Bull  frog) 
occur  in  southern  Arizona  today.  Only  R.  pipiens  and  the  introduced  species  occur 
near  Deadman  Cave  where  they  are  restricted  to  permanent  water  habitats  along  streams. 
Rana  species  have  been  recovered  as  Quaternary  fossils  in  California  (Brattstrom 
1953<2,  b,  Hudson  and  Brattstrom  1977),  Nevada  (Brattstrom  1954),  New  Mexico 
(Holman  1970,  Van  Devender  and  Worthington  1977),  and  Rancho  la  Brisca,  Sonora, 
Mexico  (Van  Devender  et  al.  in  press). 

Class  REPTILIA- Reptiles 

Order  SQU  AM  AT  A— Lizards  and  Snakes 

Suborder  Sauria  — Lizards 

Family  Iguanidae— Iguanid  lizards 

Holbrookia  maculata— Lesser  earless  lizard 

Material.  —  Dentary  ( 1 1 394). 

Remarks.— The  Lesser  earless  lizard  is  a  small  ground-dwelling  lizard  common  in 
open  habitats  of  desertscrub,  desert  grassland,  and  oak  woodlands.  It  lives  in  the  lower 
elevations  of  the  Santa  Catalina  Mountains  and  occurs  over  most  of  eastern  Arizona 
(Lowe  1964).  The  only  previous  Late  Pleistocene  and  Early  Holocene  records  of  this 
lizard  are  from  New  Mexico  (Van  Devender  and  Worthington  1977). 


255 


Holbrookia  texana— Greater  earless  lizard 

Material. -R  &  L  dentaries  (2;  11401);  R  maxilla  (1 1395). 

Remarks.  —  Teeth  of//,  texana  are  relatively  taller  on  a  deeper  dentary  than  those 
of  the  smaller  //.  maculata.  Dentaries  of  Holbrookia  have  a  closed  but  not  fused 
Meckel's  canal.  Today  this  lizard  is  found  in  open  habitats  on  the  south  side  of  the 
Santa  Catalina  Mountains,  but  not  near  the  cave  at  present.  This  insectivorous  lizard 
lives  at  middle  elevations  in  west  central  and  southern  Arizona,  avoiding  extreme 
desert  lowlands  (Stebbins  1966,  Lowe  1964).  The  only  other  known  Late  Pleistocene 
or  Early  Holocene  records  of  this  lizard  are  from  New  Mexico  (Van  Devender  and 
Worthington  1977). 

Callisaurus  draconoides— Zebra-tailed  lizard 

Material.  — L  dentaries  (2;  1 1386). 

Remarks.  — Teeth  and  dentaries  of  Callisaurus  draconoides  are  similar  to  those  of 
most  medium-sized  sceloporine  lizards  and  to  Holbrookia  in  particular,  but  they  can 
be  differentiated  using  an  ontogenetic  size  series  of  specimens.  Dentaries  and  teeth  of 
Callisaurus  are  much  larger  and  more  robust  than  those  of  species  of  Holbrookia; 
osteologically  Callisaurus  is  most  similar  to  //.  texana.  The  anterior  one  third  of  the 
Callisaurus  dentary  is  very  slender  and  has  a  more  medial,  internal,  orientation  to 
Meckel's  canal,  as  compared  to  the  more  ventral  orientation  of  either  Holbrookia  or 
Sceloporus.  The  Zebra-tailed  lizard  lives  in  regions  of  fairly  open  sandy  or  gravelly, 
low-elevation,  desertscrub  communities.  The  nearest  population  to  Deadman  Cave  is 
in  the  low  areas  near  Florence  Junction  and  along  the  San  Pedro  River.  This  is  the  first 
Late  Pleistocene-Early  Holocene  record  of  C  draconoides. 

Crotaphytus  col laris— Collared  lizard 

Material.— R  &  L  dentaries  (6;  11390);  R  &  L  maxillae  (8;  11391);  pterygoid 
(11392);  frontal  (15003). 

/^^war/c^.  — Specimens  of  C  collaris  and  C  wislizeni  (Leopard  lizard)  can  be  sep- 
arated from  most  other  iguanid  lizards  by  their  overall  larger  size,  and  the  tendency 
for  the  teeth  to  be  pointed  and  recurved,  an  adaptation  for  their  carnivorous  habits. 
The  teeth  of  C  collaris  are  relatively  wider  anteroposteriorly  than  those  of  C  wislizeni, 
with  the  posterior  teeth  strongly  tricuspid  and  the  anterior  teeth  being  more  like  blunt 
cones  with  a  slight  posterior  curve.  Both  the  pterygoid  and  the  frontal  are  more  rugose 
on  C.  collaris  than  they  are  on  C.  wislizeni. 

We  use  the  name  Crotaphytus  collaris  (sensu  lato)  and  have  not  tried  to  separate 
C.  collaris  from  C.  insularis  (Smith  and  Tanner  1972,  Montanucci  et  al.  1975).  Collared 
lizards  can  be  found  in  all  mountainous  regions  of  southern  Arizona  and  occasionally 
on  open  flat  terrain  (Lowe  1964).  It  presently  lives  near  Deadman  Cave. 

The  Collared  lizard  is  known  from  fossil  sites  in  Arizona  (Van  Devender  and  Mead 
1978,  Mead  1981,  Cole  and  Mead  1981),  Nevada  (Brattstrom  1 954a,  Mead  et  al.  1 982). 
New  Mexico  (Holman  1970,  Gehlbach  and  Holman  1974,  Van  Devender  and  Worth- 
ington 1977). 

Sceloporus  cf.  5.  clarkii— Clark's  spiny  lizard 
Material. -"L  dentaries  (2;  1 1396);  R  &  L  maxillae  (2;  1 1397). 

Sceloporus  cf.  S.  magister— Desert  spiny  lizard 
Material.  — R  &  L  dentaries  (2;  1 1398). 

Sceloporus  clarkii  or  magister— Clark's  or  Desert  spiny  lizard 

Material. -R  &  L  dentaries  (10;  1 1400);  R  &  L  maxillae  (9;  1 1399). 

Remarks.  — Osteologically  it  is  difficult  to  distinguish  these  two  moderately  large 


256 


spiny  lizards  in  their  southeastern  Arizona  range.  Sceloporus  magister  can  have  more 
robust  dental  characters.  It  usually  inhabits  the  low  deserts  but  will  occur  up  into  the 
desert-grassland.  Sceloporus  clarkii  lives  in  woodlands  in  Arizona,  but  is  a  common 
inhabitant  in  the  subtropical  thomscrub  in  Sonora,  Mexico.  Both  species  ^re  found 
today  in  the  Santa  Catalina  Mountains.  Other  Late  Pleistocene  and/or  Early  Holocene 
records  of  Clark's  spiny  lizard  are  in  New  Mexico  (Van  Devender  and  Worthington 
1977)  and  Rancho  la  Brisca,  Sonora,  Mexico  (Van  Devender  et  al.  in  press).  Sceloporus 
magister  is  fairly  common  in  the  fossil  record,  including  Arizona  (Van  Devender  and 
Mead  1978,  Mead  1981),  California  (Brattstrom  1953a,  b),  and  New  Mexico  (Van 
Devender  and  Worthington  1977). 

Sceloporus  cf.  S.  undulatus—EasXem  fence  lizard 

Material.  — R  &  L  maxillae  (3;  1 1387);  R  dentaries  (2). 

Remarks.— These  specimens  are  from  a  small  species  of  Sceloporus  similar  to 
either  S.  undulatus  or  S.  occidentalis  (Western  fence  lizard).  They  can  be  distinguished 
from  juvenile  S.  magister  or  5".  clarkii  by  their  more  slender,  taller  teeth.  Maxillae  and 
dentaries  are  less  rugose  in  the  Fence  lizard,  but  are  larger  in  all  aspects  than  the  S. 
graciosus  (Sagebrush  lizard).  We  are  not  convinced  that  5".  undulatus,  S.  occidentalis, 
or  S.  virgatus  (Striped  Plateau  lizard)  can  be  reliably  separated  satisfactorily  on  skeletal 
fragments. 

Sceloporus  undulatus  presently  occurs  near  the  cave,  while  5.  occidentalis  and  S. 
graciosus  occur  farther  north  and  S.  virgatus  occurs  in  southeastemmost  Arizona.  For 
this  reason  the  material  may  be  referred  to  S*.  undulatus.  The  Eastern  fence  lizard 
habitat  in  Arizona  ranges  from  forested  mountains  down  into  the  desert-grassland. 

Remains  of  S.  undulatus  are  known  from  Late  Pleistocene  and  Early  Holocene 
deposits  in  Arizona  (Van  Devender  and  Mead  1978,  Mead  1981,  Cole  and  Mead  1981) 
and  New  Mexico  (Holman  1970,  Van  Devender  and  Worthington  1977). 

Urosaurus  ornatus— Tree  lizard 

Material.  — L  dentary  (1 1402). 

Remarks.  — Urosaurus  ornatus  may  be  differentiated  from  most  small  iguanids 
including  U.  graciosus  (Long-tailed  Brush  lizard)  by  its  more  slender  teeth  and  the 
presence  of  a  small  fused  area  of  the  Meckel's  canal.  The  Tree  lizard  in  Arizona  occurs 
in  a  wide  variety  of  habitats  from  low,  hot  deserts  up  to  open  pine-oak  woodlands. 
Late  Pleistocene-Early  Holocene  remains  of  the  Tree  lizard  have  been  found  in  south- 
western New  Mexico  (Van  Devender  and  Worthington  1977).  The  Deadman  Cave 
specimen  is  the  first  fossil  record  for  the  species  in  Arizona. 

Phrynosoma  douglassi—Shon-homed  lizard 
Material. -R  &  L  dentaries  (6;  1 1385);  R  &  L  maxillae  (8;  1 1384). 

Phrynosoma  modestum— Round-tailed  homed  lizard 
Material. -R  dentary  (1 1382);  L  maxilla  (1 1383);  parietal  (3;  15004-15006). 

Phrynosoma  solare— Regal  homed  lizard 

Material. -Parietal  (11 381);  angular  (15007);  squamosal  (15008). 

/^^mar/c^.  —  Species  of  Phrynosoma  can  be  differentiated  from  one  another  by  most 
bones  of  the  skull,  especially  those  which  bear  horns  (see  Figs.  1-8  in  Reeve  1952). 
The  dentary,  maxilla,  and  parietal  are  very  mgose  in  P.  modestum  and  are  easily 
differentiated  from  the  similar  species,  P.  platyrhinos  (Desert  homed  lizard),  which 
lacks  rugosity.  Size,  shape,  and  omateness  will  differentiate  P.  solare  from  other  species 
(^fe  also  Reeve  1952). 

Phrynosoma  douglassi  presently  occurs  in  the  higher  forests,  woodlands,  and  grass- 


257 


land  habitats  in  eastern  Arizona  and  the  Santa  CataHna  Mountains,  whereas,  P.  solare 
Hves  in  the  Sonoran  Desert  valleys  and  bajadas,  and  adjacent  desert-grasslands.  Phry- 
nosoma  modestum  is  a  characteristic  Chihuahuan  Desert  animal  found  in  desertscrub 
and  desert-grassland  habitats.  Today  it  occurs  no  further  west  than  Sulphur  Springs 
Valley,  95  km  east  of  the  San  Pedro  River  Valley.  This  is  the  first  fossil  record  for  P. 
solare;  Phrynosoma  modestum  is  recorded  from  New  Mexico  (Van  Devender  and 
Worthington  1977);  P.  douglassi  is  recorded  from  New  Mexico  (Gehlbach  and  Holman 
1974,  Van  Devender  and  Worthington  1977)  and  Nevada  (Mead  et  al.  1982). 

Family  Teiidae— Teiid  lizards 
Cnemidophorus  species— Whiptail  lizard 

Material. -I.  dentary  (1 1389);  L  maxilla  (1 1388). 

/^6'mar/c.s'.  —  Neither  the  dentary  nor  the  maxilla  allowed  for  specific  identification. 
Five  species  of  Whiptail  lizards  occur  in  southern  Arizona  (C  burti,  C.  exanguis,  C. 
arizonae,  C.  inornatus,  and  C  tigris). 

Family  Helodermatidae— Beaded  lizards 
Heloderma  suspectum— Gila,  monster 

Material.  — Venehra  (1 1393). 

Remarks.  —  Vertebrae  of  Heloderma  can  be  separated  from  the  only  other  large 
lizard  of  comparable  size  in  Arizona,  Sauromalus  obesus  (Chuckwalla),  because  they 
lack  zygantra  and  zygosphenes  and  the  dorsal  half  of  the  cotyle  is  oval  rather  than 
subsquare  to  orbicular. 

The  Gila  monster  occurs  in  Arizona  from  the  southern  half  of  the  state  north  into 
the  extreme  northwestern  comer.  Living  primarily  in  the  lowlands  of  the  Sonoran 
Desert  and  portions  of  the  Mohave  Desert,  the  venomous  Gila  monster  also  occurs 
less  commonly  in  desert-grasslands,  and  rarely  in  the  oak  woodlands.  Heloderma  sus- 
pectum is  common  along  the  lower  portions  of  the  Santa  Catalina  Mountains  but 
probably  does  not  occur  today  at  Deadman  Cave. 

It  is  not  known  whether  the  Gila  monster  occurred  in  Arizona,  California,  and 
Nevada  during  the  Late  Wisconsinan  glacial  or  if  it  was  a  Holocene  immigrant  from 
the  Sonoran  Desert  lowlands  in  the  Lower  Colorado  River  Valley  around  the  head  of 
the  Gulf  of  California  in  Sonora,  Mexico.  Inadequately  dated  Late  Pleistocene-Holocene 
remains  occur  at  Vulture  Cave,  Arizona,  and  Gypsum  Cave,  Nevada  (Mead  and  Phillips 
1981,  Brattstrom  1954a). 

Suborder  Serpentes— Snakes 

Family  Colubridae— Colubrid  snakes 

Masticophis  species— Racer 

Material.  — W^nebvat  (1 1;  1 1404). 

Remarks.— The  vertebrae  of  Masticophis  are  similar  to  those  of  Coluber  (Racer) 
and  Salvadora  (Patch-nosed  snake)  {see  the  remarks  under  the  latter  species).  Masti- 
cophis may  be  identified  by  the  following:  1)  the  cotyle-condyle  length  (cl)  is  up  to  6.5 
mm,  occasionally  to  8.2  mm,  2)  the  ratio  of  the  cotyle-condyle  length  in  relation  to 
the  neural  arch  width  (NAW)  is  between  1.48  and  1.75,  3)  the  accessory  process  is 
long,  pointed,  and  mostly  oblique  to  anterior,  and  4)  the  ratio  of  the  distance  between 
the  prezygapophyses  (PR-PR)  and  that  distance  between  the  prezygapophysis  and  the 
postzygapophysis  (PR-PO)  is  between  0.87  and  1.00.  This  same  ratio  for  Coluber  is 
0.98  to  1.25  (Auffenberg  1963). 

Snakes  of  the  genus  Masticophis  are  large,  active,  diurnal  predators.  Within  the 
genus,  identification  to  species  is  difficult;  the  vertebrae  of  Coluber  constrictor  (Blue 
Racer)  are  similar  as  well.  Masticophis  flagellum  (Coachwhip)  and  M.  bilineatus  (So- 
noran whipsnake)  occur  near  Deadman  Cave,  while  M.  taeniatus  (Striped  whipsnake) 
occurs  in  the  mountains  to  the  north  and  northeast.  Late  Wisconsinan  and  Early 


258 


Holocene  remains  of  Masticophis  species  are  known  from  New  Mexico  (Van  Devender 
and  Worthington  1977),  Arizona,  and  California  (Van  Devender  and  Mead  1978), 
Nevada  (Mead  et  al.  1982)  and  from  the  interglacial  age  Rancho  la  Brisca,  Sonora, 
Mexico  (Van  Devender  et  al.  in  press). 

Salvador  a  species— Patch-nosed  snake 

Ma/^r/a/.— Vertebrae  (3;  1 1405). 

Remarks.— T\it  vertebrae  of  Salvadora  are  similar  to  those  of  Coluber  and  Mas- 
ticophis. All  generally  have  thin  dorsal  spines,  a  well-defined,  thin  haemal  keel,  and  a 
tendency  for  epizygapophyseal  spines.  Salvadora  is  different  from  the  latter  two  species 
in  having  a  relatively  smaller  neural  canal  and  a  smaller  condyle  (Holman  1962).  We 
do  not  know  of  any  vertebral  characters  that  unequivocally  separate  the  two  species 
within  this  genus.  Snakes  of  the  genus  Salvadora  are  small  ground  dwellers.  S.  gra- 
hamiae  (Mountain  patch-nosed  snake)  occurs  in  the  mountains  of  southeastern  Arizona 
in  oak  woodlands  and  above,  whereas  S.  hexalepis  (Desert  patch-nosed  snake)  is  widely 
distributed  in  southern  and  western  Arizona,  living  below  the  chaparral  and  woodland 
edge.  Both  species  live  in  the  Santa  Catalina  Mountains.  The  genus  was  recovered  from 
the  inter-glacial  deposit  at  Rancho  la  Brisca,  Sonora,  Mexico  (Van  Devender  et  al. 
in  press)  and  from  a  Late  Wisconsinan-Early  Holocene  cave  deposit  in  New  Mexico 
(Van  Devender  and  Worthington  1977). 

Arizona  elegans— Glossy  snake 

Material.— Ytnobvae  {9;  11413). 

/^^marfo.  —  Characters  used  to  identify  the  vertebrae  of  ^.  elegans  are:  1)  the  cl  is 
up  to  3.5  mm,  2)  the  ratio  of  cl  and  NAW  is  between  1.08  and  1.25,  3)  the  high  neural 
arch,  4)  the  neural  spine  is  high  and  moderately  thin,  5)  there  is  a  long  thin  accessory 
process  which  is  rounded  and  oblique  to  the  anterior,  and  6)  the  cotyle  is  oval  to 
subround  (Van  Devender  and  Mead  1978).  This  medium-sized  nocturnal  snake  lives 
in  deserts  and  grasslands  of  most  of  the  Southwest  as  well  as  in  northern  Mexico.  This 
snake  is  known  from  Late  Wisconsinan  and  Early  Holocene  remains  in  New  Mexico 
(Van  Devender  and  Worthington  1977)  and  Arizona  (Van  Devender  and  Mead  1978). 

Pituophis  melanoleucus— Bull  or  Gopher  snake 

Material.  -  Vertebrae  (10;  11418). 

Remarks.— Criteria,  for  identification  are  discussed  in  Auffenberg  (1963),  but  those 
used  here  are:  1)  the  cl  is  up  to  7.5  mm,  2)  the  cl/NAW  ratio  between  1.07  and  1.17, 
3)  the  neural  arch  is  very  high  with  the  neural  spine  being  high  and  thick,  4)  the 
zygosphene  is  moderately  or  strongly  convex  from  the  anterior,  5)  the  accessory  pro- 
cesses are  short,  pointed  or  blade-like,  and  6)  the  cotyle  is  round,  relatively  large,  and 
only  slightly  oblique  (Van  Devender  and  Mead  1978).  Pituophis  melanoleucus  is  a 
widespread  North  American  snake  that  lives  in  a  wide  variety  of  habitats  in  Arizona 
up  to  about  3000  m  (9900  ft).  Fossils  of  the  species  are  found  in  Arizona  (Van  Devender 
et  al.  1977,  Van  Devender  and  Mead  1978,  Mead  1981,  Cole  and  Mead  1981),  Cali- 
fornia (Brattstrom  1953a),  Nevada  (Brattstrom  1958,  1976,  Mead  et  al.  1982),  and 
New  Mexico  (Van  Devender  and  Worthington  1977). 

Lampropeltis  getulus— Common  king  snake 

Material-WertehvaQ  (7;  11416). 

Remarks.  — ¥oT  the  species  identification  characters,  see  L.  pyromelana.  Fossils  of 
L.  getulus  are  known  from  Arizona  (Van  Devender  et  al.  1977,  Van  Devender  and 
Mead  1978,  Mead  and  Phillips  1981,  Mead  1981),  California  (Brattstrom  1976,  Van 
Devender  and  Worthington  1977). 


259 


Lampropeltis  pyromelana—Sonoran  mountain  kingsnake 

Material.  — WeriehTae  (8;  11417). 

Remarks.— The  haemal  keel  and  subcentral  ridges  are  well  developed  in  the  king- 
snakes.  Lampropeltis  getulus  is  a  large  species  and  has  a  sharp  neural  spine  with  rel- 
atively blunt  accessory  processes.  Lampropeltis  pyromelana  is  a  small  species  and  has 
a  thin,  low  neural  spine  and  has  short,  pointed  accessory  processes.  Both  species  occur 
in  the  Santa  Catalina  Mountains.  L.  getulus  is  common  over  most  of  North  America, 
whereas  L.  pyromelana  is  found  in  montane  habits  in  Nevada,  Utah,  and  south  into 
Mexico.  Fossil  remains  have  been  reported  from  New  Mexico  (Van  Devender  and 
Worthington  1977)  and  Nevada  (Mead  et  al.  1982). 

Rhinocheilus  /^con/^/— Long-nosed  snake 

Material. -MQnobraQ  (22;  11419). 

Remarks.— A\\h.o\xg\\  vertebrae  of  i?.  lecontei  superficially  resemble  those  o^ Lam- 
propeltis getulus,  they  are  readily  distinguished  using  the  following  criteria:  1 )  the  cl  is 
up  to  3.0  mm,  2)  the  ratio  of  the  cl  and  the  NAW  is  between  1.07  and  1.21,  3)  the 
neural  spine  is  often  flat-topped,  4)  the  zygosphene  is  often  flat  from  the  anterior,  5) 
the  accessory  process  is  blunt,  lateral  or  dorsal  from  the  anterior,  6)  the  cotyle  is  round 
and  narrower  than  the  zygosphene,  and  7)  the  subcentral  ridges  are  well-developed, 
but  less  so  than  in  Lampropletis  getulus  (Auffenberg  1963,  Hill  1971,  and  Van  Devender 
and  Mead  1 978).  This  medium-sized,  nocturnal  snake  is  widespread  in  desert,  grassland, 
subtropical  thomscrub  habitats  in  the  Southwestern  U.S.  and  northern  Mexico.  The 
snake  probably  occurs  near  the  cave  today.  Late  Pleistocene-Holocene  fossils  occur  in 
New  Mexico  (Van  Devender  and  Worthington  1977),  Arizona  and  California  (Van 
Devender  and  Mead  1978,  Mead  1981)  and  Nevada  (Mead  et  al.  1982). 

Gyalopium  ca«wm— Western  hook-nosed  snake 

Material. — Vertebra  (11414). 

Remarks.  — VtrXehrae  of  Gyalopium  canum  are  small  but  very  broad  for  their 
length.  The  haemel  keel  is  poorly  developed  and  the  cotyle  and  condyle  are  relatively 
large  compared  to  those  of  Sonora  semiannulata  (Ground  snake)  and  Chionactis  oc- 
cipitalis (Banded  sand  snake). 

The  Western  hook-nosed  snake  is  a  small  snake  that  lives  in  the  desertscrub  and 
desert-grasslands  from  southeastern  Arizona  to  Trans-Pecos,  Texas  and  south  into  the 
Chihuahuan  Desert  of  Mexico.  Presently  it  occurs  no  closer  to  Deadman  Cave  than 
the  Santa  Rita  Mountains,  80  km  to  the  south.  The  only  previous  fossil  record  of  the 
species  (as  Ficimia  cana)  was  from  New  Mexico  (Van  Devender  and  Worthington 
1977). 

Trimorphodon  biscutatus— Lyre  snake 

Material. -YeTXebrae  (105;  1 1420). 

Remarks.  — Chteha  for  the  identification  of  T.  biscutatus  are  as  follows:  1)  the  cl 
is  up  to  4.5  mm,  2)  the  cl/NAW  ratio  is  between  1.08  and  1.25,  3)  the  neural  arch  is 
flattened,  4)  the  neural  canal  is  relatively  small,  5)  the  zygosphene  is  relatively  small. 
6)  the  accessory  process  is  short  and  pointed,  7)  the  cotyle  is  oval  to  subround,  strongly 
oblique,  narrower  than  the  zygosphene,  and  8)  the  haemal  keel  is  well-developed  but 
low  (Van  Devender  and  Mead  1978).  The  Lyre  snake  is  a  medium-sized  species  that 
lives  in  desertscrub  habitats  in  the  Southwest  and  northern  Mexico.  It  is  found  near 
Deadman  Cave  today.  Fossils  occur  in  New  Mexico  (Van  Devender  and  Worthington 
1977),  Arizona,  and  California  (Van  Devender  and  Mead  1978). 

Hypsiglena  torquata—Nighl  snake 
MateriaL-Yenebrae  {\2-  11403,  11415). 


260 


Remarks.— Criieria  for  identification  are  as  follows:  1)  the  cl  is  between  1.65  and 
2.75  mm,  2)  the  cl/NAW  ratio  is  1 . 1 8  to  1 .3 1 ,  3)  the  neural  arch  is  moderately  depressed 
from  the  posterior,  4)  the  neural  spine  is  low,  usually  with  the  dorsal  edge  thickened 
and  the  anterior  comer  is  bifurcate,  5)  the  accessory  process  is  lateral  from  the  anterior, 
and  6)  the  cotyle  is  relatively  small  (Van  Devender  and  Mead  1978).  The  Night  snake 
is  widespread  in  desert,  grassland,  and  woodland  habitats  in  the  Southwestern  U.S. 
and  northern  Mexico.  It  occurs  today  near  Deadman  Cave.  Fossils  are  known  from 
New  Mexico  (Van  Devender  and  Worthington  1977),  Arizona  and  California  (Van 
Devender  and  Mead  1978,  Mead  1981),  Nevada  (Mead  et  al.  1982),  and  Rancho  la 
Brisca,  Sonora,  Mexico  (Van  Devender  et  al.  in  press). 

Family  Viperidae  (=  Crotalidae)— Pit  vipers 
Crotalus  a^rox— Western  diamondback  rattlesnake 

Material  — YtnobraQ  (5;  1 1406). 

Remarks.— Tht  thoracic  vertebrae  of  the  Viperidae  are  distinct  from  those  of  the 
Colubridae  and  Boidae  in  that  they  have  a  long,  pointed  hypophysis.  The  fossil  vertebrae 
are  from  a  large  rattlesnake  resembling  C.  atrox;  other  rattlesnakes  in  Arizona  generally 
do  not  attain  its  size,  except  for  some  C.  molossus  (Blacktailed  rattlesnake).  Both  species 
occur  near  Deadman  Cave.  Crotalus  atrox  is  a  large  desert  species  usually  occuring  in 
the  lower  valleys,  whereas,  C  molossus  is  a  woodland-dwelling  species  that  is  occa- 
sionally found  in  rocky  habitats  in  more  xeric  desert  mountain  ranges.  Late  Pleistocene 
fossils  of  C.  atrox  have  been  reported  from  Gypsum  Cave,  Nevada  (Brattstrom  1 954(2, 
b)  and  Conkling  Cavern,  Shelter,  Fosbert  (Brattstrom  1964)  and  Dry  caves.  New 
Mexico  (Holman  1970).  There  are  no  unequivocal  records  of  fossil  C.  atrox  from 
Arizona;  however,  there  is  the  interglacial  record  from  Rancho  la  Brisca,  Sonora,  Mexico 
(Van  Devender  et  al.  in  press). 

Crotalus  scutulatus— Mohave  rattlesnake 

Material.  — Vertehme  (3;  1 1407). 

Remarks.— The  fossil  vertebrae  are  from  a  medium-sized  rattlesnake  that  is  smaller 
than  Crotalus  atrox  or  C.  molossus.  The  vertebrae  of  C.  viridus  cerberus  (Arizona  black 
rattlesnake),  a  common  snake  in  the  oak  woodland,  differ  from  those  of  C.  scutulatus 
in  their  relative  size  of  the  hypophysis  {see  also  Brattstrom  \96Ab).  The  vertebrae  of 
C.  cerastes  (Sidewinder),  C  lepidus  (Rock  rattlesnake),  C.  pricei  (Twin  spotted  rattle- 
snake), and  C.  willardi  (Ridgenosed  rattlesnake)  are  smaller  and  differ  in  various  mor- 
phological characters.  The  Mohave  rattlesnake  is  a  common  desert-grassland  and  des- 
ertscrub  snake  in  southern  Arizona  and  near  Deadman  Cave  today.  Fossils  of  C 
scutulatus  have  not  been  reported  previously. 

Class  AVES- Birds 

Order  GALLIFORMES- Gallinaceous  birds 

Family  Phasianidae—  Pheasants,  quails,  etc. 

Colinus  gambelii—GamheVs  quail 

Material. -CormplelQ  carpometacarpus  (15313). 

Remarks.— This  specimen  is  much  smaller  than  the  carpometacarpi  of  C  ("'Or- 
eortyx"")  pictus  (Mountain  quail)  or  Cyrtonyx  montezumae  (Harlequin  quail),  and  is 
slightly  smaller  than  that  in  C.  {"'Callipepla'")  squamata  (Scaled  quail).  It  differs  from 
C.  squamata  and  C.  virginianus  (Bobwhite)  in  having  the  Os  metacarpale  minus  (meta- 
carpal III)  more  slender  and  more  curved  in  caudal  aspect,  and  in  having  a  slightly 
smaller  Processes  extensorius.  This  is  only  the  second  fossil  occurrence  of  C.  gambelii, 
the  other  being  from  the  Early  Pleistocene  (Irvingtonian)  of  Vallecito  Creek,  California 
(Howard  1 963).  Brodkorb  ( 1 964)  listed  C.  gambelii  questionably  from  Conkling  Cavern 
and  Shelter  Cave,  New  Mexico.  These  assignments  conflict,  however,  with  the  original 
references,  as  Howard  and  A.   H.   Miller  (1933)  reported  ""Lophortyx  sp.  Quail" 


261 


(sic)  from  these  two  sites.  Gambel's  quail  lives  today  in  the  vicinity  of  Deadman  Cave, 
but  is  approximately  at  its  upper  elevational  limit. 

Colinus  species— quail 

Material.  — VroxxmaX  and  distal  ends  of  humeri  (15009). 

Remarks.— Among  quail  of  the  Southwest,  these  specimens  are  smaller  than  the 
humeri  of  Colinus  {""Oreortyx")  pictus  or  Cyrtonyx  montezumae.  We  cannot,  however, 
distinguish  them  from  humeri  of  Colinus  virginianus,  C.  {"" Lophortyx")  gambelii,  or 
C.  C'Callipepla"")  squamata,  any  of  which  could  have  occurred  at  Deadman  Cave. 
Although  species-level  identifications  are  often  very  difficult,  quail  of  the  genus  Colinus 
are  common  as  Pleistocene  fossils  in  southern  North  America,  especially  Florida,  New 
Mexico,  and  California. 

Cyrtonyx  montezumae— HarlQquin  quail 

Material.  — PTOximal  end  with  partial  shaft  of  radius  (15010). 

Remarks.— This  specimen  is  larger  than  the  radius  in  all  other  southwestern  quail 
except  Colinus  pictus.  It  is  referable  to  Cyrtonyx  by  the  less  expanded  articulating 
surface  of  the  proximal  end  relative  to  the  width  of  the  shaft.  The  only  other  fossil 
occurrence  of  C  montezumae  is  at  San  Josecito  Cave,  Nuevo  Leon,  Mexico  (L.  Miller 
1 943),  also  of  Late  Pleistocene  age.  Today  the  Harlequin  quail  occurs  in  grassy  mountain 
woodlands  of  central  and  southeastern  Arizona,  thence  ranging  south  well  into  Mexico. 
This  species  is  very  characteristic  of  evergreen  oak  grassland  and  is  at  its  lower  ele- 
vational limit  near  Deadman  Cave  today.  With  the  historical  reduction  of  grass  and 
increase  in  brush  at  mid-elevations  in  Arizona  mountains,  this  once  common  bird  has 
decreased  in  abundance. 

Order  COLUMBIFORMES- Pigeons  and  doves 

Family  Columbidae— Pigeons  and  doves 

Zenaida  cf  Z.  macrawra— Mourning  dove 

Marma/.  — Proximal  end  of  carpometacarpus  (1501 1). 

Remarks.— This  fossil  differs  markedly  in  size  from  the  carpometacarpi  of  all 
Arizona  columbids  except  Zenaida  macroura  and  Z.  asiatica  (White-winged  dove).  It 
is  tentatively  assigned  to  Z.  macroura  in  being  slightly  smaller  than  all  available  spec- 
imens of  Z.  asiatica.  This  is  the  first  Pleistocene  record  of  Z.  macroura  in  Arizona, 
although  this  species  is  a  fairly  common  Late  Pleistocene  fossil  elsewhere  in  North 
America.  The  Mourning  dove  is  very  widespread  in  Arizona,  both  geographically  and 
altitudinally,  and  thus  is  of  little  paleoecological  interest. 

Order  STRIGIFORMES-Owls 

Family  Strigidae— Typical  owls 
Otus  species— Screech-owl 

Material.— Tv^o  proximal  ends  and  one  distal  end  of  humeri  (3;  15012),  proximal 
end  of  carpometacarpus  (15013). 

Remarks.  -These  specimens  all  agree  in  size  and  morphology  with  Otus  asio  (Com- 
mon screech-owl),  and  are  either  larger  or  smaller  than  in  all  Arizonan  owls  outside  of 
the  genus  Otus.  The  carpometacarpus  and  one  proximal  end  of  humerus  are  slightly 
larger  than  in  O.  Jlammeolus  (Flammulated  screech-owl),  but  the  other  elements  re- 
semble both  O.  asio  and  O.  jlammeolus.  No  skeleton  was  available  for  O.  trichopsis 
(Spotted  screech-owl),  so  identification  beyond  generic  level  is  not  possible.  These  three 
species  of  Otus  in  Arizona  are  largely  separated  from  each  other  today  by  habitat  and 
elevation,  and  it  seems  most  likely  that  O.  asio  or  O.  trichopsis  would  have  lived  near 
Deadman  Cave  in  the  Late  Pleistocene.  Probably  only  O.  asio  occurs  in  the  immediate 
vicinity  of  Deadman  Cave  today.  Otus  asio  is  a  common  Late  Pleistocene  fossil  in 


262 


North  America,  while  O.  flammeolus  and  O.  thchopsis  have  only  two  and  one  Pleis- 
tocene records,  respectively.  This  is  the  first  fossil  record  of  Otus  in  Arizona. 

Micrathene  Whitney i— Elf  o'wl  * 

Material.  —  Proximal  end  of  humerus  (1501 4),  distal  end  of  tarsometatarsus  (1501 5). 

Remarks.— ThQ  Elf  owl  is  readily  separated  from  all  other  owls  by  its  extremely 
small  size.  This  is  the  first  Pleistocene  record  for  M.  whitneyi.  It  occurs  today  in  the 
region  of  Deadman  Cave,  nesting  in  holes  in  trees  at  any  elevation  "below  the  heavy 
pine  forest"  (Phillips  et  al.  1964). 


Asio  o?W5— Long-eared  owl 

Material  — 'D\s\.3\  end  of  humerus  (15016). 

Remarks.  — This  fossil  agrees  with  the  humerus  of  Asio  otus  versus  A.  flammeus 
(Short-eared  owl)  in  having  a  distinctive  knot-like  ectepicondylar  prominence  (Pro- 
cesses supracondylaris  dorsalis).  Asio  otus  is  a  fairly  common  Late  Pleistocene  fossil 
in  western  North  America,  but  this  is  the  first  such  record  in  Arizona.  The  Long-eared 
owl  is  not  unexpected  at  Deadman  Cave,  as  it  occurs  today  in  Arizona  in  a  variety  of 
habitats,  both  as  a  nesting  bird  and  a  winter  visitor.  As  mentioned  above,  A.  otus 
probably  was  involved  in  the  accumulation  of  small  vertebrates  in  Deadman  Cave. 

Order  CAPRIMULGIFORMES- Goatsuckers,  etc. 

Family  Caprimulgidae— Night  jars 

Genus  and  species  indeterminate 

Ma?m<3/.  —  Carpometacarpus  lacking  distal  end  and  much  of  metacarpal  III  (1 50 1 7). 

Remarks.  — This  carpometacarpus  is  distinguished  from  that  of  Chordeiles  minor 
(Common  nighthawk)  and  C.  acutipennis  (Lesser  nighthawk)  by  its  much  smaller  size, 
and  from  Caprimulgus  vociferus  (Whip-poor-will)  by  its  slightly  smaller  overall  size 
with  a  more  slender  metacarpal  III.  It  resembles  that  of  Phalaenoptilus  nuttallii  (Poor- 
will)  very  closely,  but  the  shape  of  metacarpal  I  is  somewhat  more  similar  to  that  in 
Caprimulgus.  In  the  absence  of  a  comparative  skeleton  of  Caprimulgus  ridgwayi  (Ridge- 
way's  whip-poor-will),  the  only  other  caprimulgid  living  in  Arizona,  precise  identifi- 
cation of  this  fossil  is  impossible.  The  Pleistocene  record  of  caprimulgids  is  poorly 
known  everywhere.  This  is  the  first  fossil  record  for  the  family  in  Arizona. 

Order  PICIFORMES- Woodpeckers,  etc. 

Family  Picidae— Woodpeckers 

Colaptes  auratus—¥hc\<.tr 

Material.  —  Distal  end  of  tarsometatarsus  (1501 8). 

Remarks.— Among  Arizonan  woodpeckers,  the  tarsometatarsus  of  Colaptes  au- 
ratus  is  similar  in  size  only  to  that  of  Melanerpes  {^" Asyndesmus"")  lewis  (Lewis'  wood- 
pecker). The  fossil  agrees  with  C.  auratus  versus  M.  lewis  in  its  larger,  less  deeply 
sculptured  middle  trochlea.  The  distal  end  of  the  tarsometatarsus  in  the  "Red-shafted" 
flicker  (C  a.  collaris)  appears  to  be  indistinguishable  from  that  in  the  "Gilded"  flicker 
(C  a.  mearnsi).  Thus  the  fossil  provides  no  evidence  of  paleohabitats  near  Deadman 
Cave.  C.  a.  collaris  is  a  bird  of  mountain  woodland  and  forest,  ranging  upward  from 
approximately  1220  m  elevation,  whereas  C.  a.  mearnsi  occurs  in  desertscrub,  generally 
below  1 370  m  elevation.  Based  on  the  remainder  of  the  avifauna,  one  would  guess  that 
the  Late  Pleistocene  flicker  at  Deadman  Cave  was  C  a.  collaris,  although  both  forms 
occur  in  the  general  region  of  the  site  today.  Flickers  are  very  common  Late  Pleistocene 
fossils,  yet  once  again  this  is  the  first  such  record  for  Arizona. 


263 

Order  PASSERIFORMES- Perching  birds 

Family  Turdidae  — Thrushes 
Turdus  cf.  T.  migratorius— American  robin 

Material.  — Proximal  end  of  humerus  (15019). 

Remarks.— The  humerus  of  Turdus  migratorius  can  be  told  from  that  of  most 
other  North  American  turdids  by  its  larger  size.  It  can  be  recognized  from  that  in 
Ixoreus  naevius  (Varied  thrush)  by  its  stouter  Crus  dorsalis  fossae  which,  along  with 
the  deeper  dorsal  Fossa  pneumotricipitalis,  also  distinguishes  it  from  the  humeri  of 
mimids  (thrashers,  mockingbirds).  The  humerus  of  T.  migratorius  can  be  separated 
from  that  in  the  neotropical  T.  grayi  (Clay-colored  robin)  reported  from  Stanton's 
Cave,  Coconino  County,  Arizona  (Rea  and  Hargrave,  ms)  by  its  lesser  degree  of  pneu- 
maticity  in  both  the  dorsal  and  ventral  Fossa  pneumotricipitalis,  the  former  also  being 
larger  in  T.  grayi.  The  fossil  differs  from  the  only  available  humerus  of  T.  rufopalliatus 
(Rufous  backed  robin,  a  vagrant  to  Arizona  today;  resident  in  Sonora)  in  having  a 
larger  and  more  oblong  (less  circular)  opening  of  the  ventral  Fossa  pneumotricipitalis. 
Lacking  additional  specimens  of  T.  rufopalliatus  to  confirm  this  character,  no  more 
than  a  tentative  assignment  of  the  fossil  to  T.  migratorius  is  warranted.  T.  migratorius 
is  a  common  Late  Pleistocene  species  in  much  of  North  America,  and  has  been  reported 
in  Arizona  from  Stanton's  Cave  (Rea  and  Hargrave,  ms).  The  American  robin  is 
common  in  Arizona  today,  nesting  throughout  the  state  in  wooded  regions  above 
approximately  1220  m  elevation. 

Catharus  guttatus-^HermiX  thrush 

Material.  — 'DisXaX  end  of  humerus  with  most  of  shaft  (19020). 

Remarks.— The  humerus  of  Catharus  guttatus  is  smaller  than  in  mimids  and  in 
the  following  species  of  turdids:  Hylocichla  mustelina  (Wood  thrush),  Ixoreus  naevius, 
Myadestes  townsendi  (Townsend's  solitaire),  and  all  species  of  Turdus.  It  is  larger  and 
has  a  stouter  Corpus  humeri  (shaft)  than  in  T.  ustulatus  (Swainson's  thrush).  It  is 
approximately  equal  in  size  to  that  of  C.  fuscescens  (Veery),  C.  minimus  (Gray-cheeked 
thrush),  and  the  species  of  Sialia  (bluebirds),  but  is  told  from  these  and  all  other  Arizona 
turdids  by  having  a  relatively  smaller  Processus  supracondylaris  dorsalis,  this  being 
particularly  evident  in  dorsal  aspect,  where  P  dorsalis  is  seen  not  to  extend  as  far 
proximally  in  C  guttatus  as  in  other  species. 

This  is  the  first  fossil  record  anywhere  for  C.  guttatus.  Brodkorb  (1978)  listed  C. 
guttatus  from  the  Late  Pleistocene  site  of  Carpinteria,  California,  citing  A.  H.  Miller 
(\932b)  as  the  authority.  However,  A.  H.  Miller  (1932^^)  clearly  did  not  refer  the 
specimen  in  question,  a  humerus,  to  any  species.  Miller  listed  the  specimen  as  "HY- 
LOCICHLA? Thrush"  (sic),  stating  that  the  fossil  resembled  Hylocichla  {=Catharus) 
guttata  in  certain  aspects,  H.  mustelina  in  others,  and  probably  represented  an  extinct 
taxon  of  thrushes.  The  Hermit  thrush  is  widespread  in  Arizona  today,  and  is  common 
in  the  Santa  Catalina  Mountains,  nesting  at  high  elevations  and  occurring  elsewhere 
as  a  migrant  or  wintering  bird. 

Family  Fringillidae— Sparrows,  finches,  tanagers,  blackbirds,  warblers,  etc. 

Subfamily  Icterinae— Blackbirds,  etc. 
Genus  and  species  indeterminate 

Material.  — Distal  end  of  tarsometatarsus  (15021). 

Remarks.— This  specimen  is  larger  than  the  tarsometatarsus  in  all  non-icterine, 
nine-primaried  oscines  of  the  Southwest.  Among  southwestern  icterines,  it  is  smaller 
than  in  Sturnella  magna  (Eastern  meadowlark),  S.  neglecta  (Western  meadowlark), 
and  Cassadix  mexicanus  (Boat-tailed  grackle),  and  larger  than  in  Molothrus  ater  (Brown- 
headed  cowbird)  or  any  species  of  Icterus  (orioles).  Of  the  species  that  it  approximates 
in  size,  the  fossil  may  be  distinguished:  from  Agelaius  phoeniceus  (Red-winged  black- 


264 


bird)  and  Xanthocephalus  xanthocephalus  (Yellow-headed  blackbird)  by  its  more  dorso- 
plantar  expansion  of  the  middle  and  inner  trochleae;  from  Euphagus  cyanocephalus 
(Brewer's  blackbird)  and  E.  carolinus  (Rusty  blackbird)  by  its  larger  intertrochlear 
spaces,  the  more  dorso-plantar  expansion  of  the  inner  trochlea,  and  the  more  proximo- 
distally  expanded  outer  trochlea;  and  from  Molothrus  aeneus  (Bronzed  coWbird)  by  its 
slightly  wider  outer  intertrochlear  space  and  slightly  more  laterally  compressed  inner 
trochlea.  Overall,  this  specimen  seems  to  be  more  similar  to  the  tarsometatarsus  of 
Molothrus  aeneus  than  to  any  other  living  icterine,  but  is  not  similar  enough  to  be 
referred  confidently  to  that  species. 

The  fossil  icterine  from  Deadman  Cave  may  represent  an  extinct  taxon.  Four 
species  of  extinct  icterines  have  been  described  from  rostra  and  mandibles  from  Late 
Pleistocene  sites  in  North  America.  These  are  Pandanaris  convexa  (A.  H.  Miller  1947), 
and  Euphagus  magnirostris  (A.  H.  Miller  1929),  both  from  Rancho  La  Brea,  California; 
Pandanaris  floridana  (Brodkorb  1957)  from  Reddick  and  Haile  XIB,  Florida;  and 
Pyeloramphus  molothroides  (A.  H.  Miller  1932a)  from  Shelter  Cave,  New  Mexico. 
Referred  post-cranial  elements  have  been  reported  for  Pandanaris  floridana  and  Eu- 
phagus magnirostris,  but  we  have  not  examined  this  material.  All  Late  Pleistocene 
icterines  are  in  need  of  re-study  (Steadman  and  Martin,  in  press),  and  pending  such 
work  the  specimen  from  Deadman  Cave  is  best  left  unidentified.  Nevertheless,  it  may 
represent  a  new  faunal  element  for  Arizona. 

Subfamily  Emberizinae— "New  World"  sparrows,  finches,  etc. 
Genus  and  species  indeterminate 

Afafm^/.— Tarsometatarsus  lacking  proximal  end  (15022). 

i^^mar/c^.— Postcranial  emberizine  fossils  are  often  very  difficult  or  impossible  to 
identify  to  genus  or  species.  The  present  specimen  is  smaller  than  the  tarsometatarsi 
of  any  icterine  (blackbirds)  or  North  American  thraupine  (tanagers),  and  is  smaller  than 
in  most  parulines  (New  World  warblers).  It  differs  from  the  tarsometatarsi  of  vireonids 
(vireos)  in  its  more  slender  middle  trochlea  and  broader  inner  trochlea,  this  last  char- 
acter also  distinguishing  it  from  the  tarsometatarsi  of  parulines.  Within  the  emberizines, 
no  readily  apparent  patterns  of  tarsometatarsal  variation  are  discernible.  When  com- 
pared to  all  species  of  North  American  emberizines,  the  fossil  was  found  to  be  indis- 
tinguishable, both  in  size  and  quality,  from  the  following  species  of  medium-sized 
sparrows:  Ammodramus  sandwichensis  (Savannah  sparrow),  Melospiza  lincolnii  (Lin- 
coln's sparrow),  M.  georgiana  (Swamp  sparrow),  Junco  hyemalis  (Dark-eyed  junco), 
and  J.  phaeonotus  (Mexican  junco).  Geographical  and  sexual  variation  combine  to 
render  the  tarsometatarsus  of  these  five  species  inseparable  in  many  instances.  Certain 
individuals  of  each  species  appear  to  be  distinct,  but  no  consistent  variation  is  seen. 
Each  of  these  species  occurs  today  in  southern  Arizona,  although  in  different  habitats 
and  in  very  different  frequencies. 

Class  MAMMALIA— Mammals 
Order  INSECTIVORA-Insectivores 

Family  Soricidae— Shrews 
Notiosorex  crawfordi—DQseri  shrew 

Material. -L  mandible  (3;  15023);  R  mandible  (2;  15024). 

Remarks.  — The  shape  of  the  mandibles  and  the  presence  of  three  unicusps  on  each 
jaw  were  the  identifying  characters.  Notiosorex  crawfordi  occurs  fairly  commonly  as 
fossils  in  Arizona  (Mead  and  Phillips  1981,  Mead  et  al.  1 983),  New  Mexico,  and  Texas 
(Harris  1 977).  Presently  the  Desert  shrew  occupies  a  wide  variety  of  ecological  situations 
from  semi-desertscrub  to  woodland  (Armstrong  and  Jones  1972).  It  is  not  known  to 
occur  at  present  in  the  Santa  Catalina  Mountains,  but  it  does  live  nearby  (Cockrum 
1960). 


265 


Order  CHIROPTERA-Bats 

Family  Vespertilionidae— Vesperlilionid  bats 

cf.  Mvo//5— Mouse-eared  bat 

Material. -h  mandible  with  M.  (15025);  R  mandible  (3;  15027). 

Remarks.— T\iQSQ  specimens,  clearly  a  vespertilionid  based  on  the  shape  of  the 
jaw,  could  not  be  identified  unequivocally  to  genus  because  of  fragmentation  and/or 
for  loss  of  teeth.  Skinner  ( 1 942)  reported  Myotis  cf.  M.  velifer  (Cave  myotis),  M.  cf. 
M.  thysanodes  (Fringed  myotis),  and  M.  cf.  M.  evoltis  (Long-eared  myotis)  from  Papago 
Springs  Cave,  Arizona. 

Antrozous  pallidus— Pallid  bat 

Material. -L  femur  (15025). 

Remarks.— The  greatest  length  of  the  fossil  femur  is  19.0  mm  and  the  width  of 
the  proximal  end  is  3.0  mm  (lesser  trochanter  to  greater  trochanter).  A  blade-like  third 
trochanter  is  present.  The  Deadman  Cave  specimen  was  compared  to  Myotis  thysan- 
odes, M.  californicus  (California  myotis),  M.  velifer,  Plecotus  townsendii  (Townsend's 
big-eared  bat),  Lasiurus  borealis  (Red  bat),  L.  cinereus  (Hoary  bat),  Macrotus  water- 
housei  (Leaf-nosed  bat),  Tadarida  brasiliensis  (Brazilian  free-tailed  bat),  T.  femorosacca 
(Pocketed  free-tailed  bat),  Mormoops  megalophylla  (Ghost-faced  bat),  Antrozous  pal- 
lidus,  and  Eptesicus  fuscus  (Big  brown  bat).  Only  the  last  two  species  were  similar  to 
the  fossil  in  having  a  femur  of  total  length  averaging  near  19.0  mm  and  a  proximal 
width  of  2.8  to  3.0  mm,  along  with  the  lesser  trochanter  as  pronounced  as  the  greater 
trochanter;  but  of  these  two,  only  A.  pallidus  had  the  third  trochanter.  The  other  species 
of  bats  lacked  two  or  all  of  the  criteria  used  to  differentiate  the  fossil  specimen. 

The  Pallid  bat  occurs  throughout  Arizona  and  can  be  found  near  Deadman  Cave 
today  (Cockrum  1960,  Barbour  and  Davis  1969).  Bats  of  the  genus  Antrozous  have 
been  recovered  as  fossils  in  a  wood  rat  midden  in  the  Sonoran  Desert  (Mead  et  al. 
1983)  and  from  Papago  Springs  Cave  (Skinner  1942). 


Order  EDENTATA -Edentates 

Family  Megatheriidae— Megathere  ground  sloths 

Nothrotheriops  shastensis —Shasta,  ground  sloth 

Material.  — Molar  (15314). 

Remarks.— Greg  McDonald  (Royal  Ontario  Museum,  1982,  personal  communi- 
cation) confirmed  this  identification  of  A^.  shastensis  and  indicated  that  because  the 
small  molar  contained  a  high  percentage  of  hollow  pulp  cavity  and  lacked  wear  stria- 
tions,  it  must  have  been  from  a  fetal  or  new  bom  sloth.  Remains  of  the  extinct  Shasta 
ground  sloth  are  very  common  in  the  Southwest,  especially  in  Arizona  (Long  and  Martin 
1974,  Thompson  et  al.  1980).  A  typographical  error  in  Lindsay  and  Tessman  (1974) 
has  the  sloth  incorrectly  located  in  Stanton's  Cave,  Grand  Canyon. 


Order  LAGOMORPHA-Lagomorphs 

Family  Leporidae— Hares  and  rabbits 

Sylvilagus  species— Cottontail 

Material  — 'L  mandible;  R  mandibles  (2);  maxilla. 

Remarks.  — VosXcramal  remains  of  leporids  were  the  second  most  common  ele- 
ments in  the  fossil  deposit.  The  mandibles  and  the  maxilla  are  not  identified  to  species 
at  this  time  because  a  more  detailed  study  of  all  Late  Pleistocene  leporid  remains  of 
Arizona  is  in  order  and  will  be  appearing  in  the  near  future  (JIM).  The  genus  is  recovered 
from  a  number  of  Late  Pleistocene  localities  in  Arizona  (Lindsay  and  Tessman  1974, 
Mead  et  al.  1983)  and  New  Mexico  (Harris  1977). 


266 

Lepus  species— Jackrabbit 

Material  — l.  mandibles  (6);  R  mandibles  (4);  L  maxillae  (2);  R  maxillae  (2);  pre- 
maxilla;  isolated  molars  (4);  L  femur  proximal  half. 

Remarks.— See  the  remarks  under  Sylvilagus.  Lepus  alleni  (Antelope  jackrabbit) 
and  L.  callotis  (White-sided  jackrabbit)  both  presently  occur  in  Arizona,  but  not  near 
Deadman  Cave.  L.  californicus  occurs  near  Deadman  Cave  today.  Lepus  californicus 
(Black-tailed  jackrabbit)  was  recovered  from  Papago  Springs  Cave  (Skinner  1 942)  and 
other  Arizona  localities  (Lindsay  and  Tessman  1974,  Mead  et  al.  1983). 

Order  RODENTI A— Rodents 

Family  Sciuridae— Squirrels 

Spermophilus  variegatus—Rock  squirrel 

Material.  — L  mandible  with  M,_2  (2;  15028);  R  mandible  (15029);  L  maxilla  with 
M'-M15030). 

Remarks.— Spermophilus  variegatus  can  be  differentiated  from  other  species  of 
ground  squirrels  by  its  larger  size  and  the  tendency  of  the  skeleton  to  be  slightly  more 
rugose.  The  only  other  squirrel  of  similar  size  is  Sciurus  aberti  (Abert's  squirrel).  The 
P'*  is  relatively  larger  in  Spermophilus  variegatus  as  compared  to  that  in  Sciurus.  The 
shape  and  medial  inflection  of  the  angle  on  the  mandible  is  greater  on  S.  variegatus. 
The  Rock  squirrel  is  a  common  ground  squirrel  and  the  largest  within  its  distribution. 
It  prefers  rocky  regions  and  is  found  throughout  the  Southwest,  including  the  vicinity 
of  Deadman  Cave.  Fossil  remains  of  the  Rock  squirrel  are  not  common  in  Late  Pleis- 
tocene localities  in  the  Southwest  (Harris  1977,  Mead  1981,  Kurten  and  Anderson 
1980),  although  Skinner  (1942)  identified  three  mandibular  rami  of  S.  (^Citellus) 
variegatus  from  Papago  Springs  Cave. 

Family  Geomyidae— Pocket  gophers 
Thomomys  cf.  T.  bottae— Bonne's  pocket  gopher 

Material. -L&  R  maxillae  (3;  1 5036);  isolated  teeth  (9;  1 5037);  R  humerus  (1 5038). 

Remarks.— The  upper  incisors  were  lacking  any  conspicuous  longitudinal  groove, 
and  the  maxillary  and  isolated  cheek  teeth  all  were  the  lobbed,  simple  hypsodont  molars 
and  premolars  of  Thomomys.  We  follow  Thaeler  (1968)  in  using  the  designation  T. 
bottae,  which  is  the  T.  umbrinus  of  Hall  (1981).  The  former  species  is  common  today 
in  the  Santa  Catalina  Mountains,  and  therefore,  the  reason  for  our  identification  of  the 
fossils.  The  genus  is  a  common  fossil  recovered  in  the  Southwest  (Mawby  1 967,  Lindsay 
and  Tessman  1974,  Harris  1977). 

Family  Heteromyidae— Pocket  mice  and  Kangaroo  rats 
Perognathus  cf.  P.  flavus SiWay  pocket  mouse 

Material. -K  maxillae  (3;  15032);  L  maxillae  (2;  15033);  R  mandibles  (3;  15034); 
L  mandibles  (4;  15035). 

Remarks.— The  pocket  mouse  specimens  from  Deadman  Cave  compare  well  with 
P.  flavus,  although  two  other  indistinguishable,  small  pocket  mice,  P.  parvus  (Great 
Basin  pocket  mouse)  and  P.  flavescens  (Plains  pocket  mouse)  could  also  be  in  the 
assemblage.  Our  tentative  identification  is  based  on  the  present  geographic  distributions. 
Complete  skulls  are  needed  for  unequivocal  identification.  Perognathus  flavus  lives 
near  Deadman  Cave  region.  The  only  other  Late  Pleistocene  occurrence  of  this  mouse 
is  from  Isleta  Cave,  New  Mexico  (Harris  and  Findley  1964). 

Dipodomys  5/?£'c/<3Z>///5— Banner- tailed  kangaroo  rat 

Material.  —  Bacculum  ( 1 503 1 ). 

Remarks.— The  shape  of  the  bacculum  o^  Dipodomys  spectabilis  is  distinct  from 
that  of  all  other  species.  Dipodomys  spectabilis  is  a  large  kangaroo  rat  that  inhabits  the 


267 


desert-grasslands  of  southeastern  Arizona,  including  the  valleys  below  the  Santa  Cat- 
alina  Mountains  (Cockrum  1960,  Hall  1981).  Harris  (1977)  has  reported  fossils  of  D. 
spectabilis  from  southern  New  Mexico. 

Family  Cricetidae— New  World  Rats  and  Mice 
Reithrodontomys  mo nt anus— Plains  harvest  mouse 

Material. -L  mandible  (15041);  R  mandibles  (3;  15042). 

Remarks.— The  mandibles  and  teeth  of  the  harvest  mice  from  Deadman  Cave 
compare  favorably  with  those  of  R.  montanus.  The  other  harvest  mice  in  Arizona,  R. 
megalotis  (Western  harvest  mouse)  and  R.  fulvescens  (Fulvous  harvest  mouse),  are  both 
larger  than  R.  montanus.  The  occlusal  pattern  of  the  molar  of  R.  fulvescens  is  an  "S" 
configuration  as  opposed  to  a  "C"  in  R.  montanus  (Hooper  1952). 

Reithrodontomys  montanus  occurs  today  in  the  grasslands  of  southeastern  Arizona 
but  not  in  the  Santa  Catalina  Mountains.  The  other  two  species  occur  in  a  wider  variety 
of  communities  (Cockrum  1960).  Fossil  occurrences  are  discussed  in  Kurten  and  An- 
derson (1980). 

Peromyscus  species— Deer  mouse 

Materia!.— h  mandibles  (4);  R  mandibles  (4). 

Remarks.  — Fragmenls  of  Peromyscus  can  be  confused  with  those  of  Reithrodon- 
tomys. The  following  characters  will  separate  the  two  genera:  1)  the  M3  is  relatively 
larger  on  Peromyscus,  2)  the  articular  condyle  of  the  mandible  extends  more  posteriorly 
than  does  the  angle,  on  Peromyscus,  and  3)  the  angle  of  the  mandible  has  a  less  medial 
inflection  on  Peromyscus.  We  were  unable  to  identify  these  specimens  to  species.  Eight 
species  of  Peromyscus  occur  in  southern  Arizona,  thus  species  level  identification  of 
fossils  is  extremely  difficult  if  not  impossible.  Fossils  of  the  genus  have  been  recovered 
from  all  over  the  Southwest  (Harris  1977,  Kurten  and  Anderson  1980,  Mead  et  al. 
1983). 

Sigmodon  species— Cotton  rat 

Material. -K  mandible  with  M,_3  (15039);  R  maxilla  with  M'--  (15040). 

Remarks.— The  occlusal  pattern  on  all  cheek  teeth  are  distinct  in  Sigmodon.  We 
are  not  able  to  identify  our  specimens  to  species.  Four  species  of  cotton  rat  now  inhabit 
southern  Arizona  (Hall  1981).  Sigmodon  hispidus  (Hispid  cotton  rat)  occurs  on  the 
western  and  eastern  borders  of  southern  Arizona  while  Sigmodon  arizonae  (Arizona 
cotton  rat)  lives  in  the  area  of  the  Santa  Catalina  Mountains.  Both  5. /w/v/'v^^f^T  (Tawny- 
bellied  cotton  rat)  and  S.  ochrognathus  (Yellow-nosed  cotton  rat)  occur  south  and  east 
of  Deadman  Cave  (Baker  and  Shump  1978a,  b.  Hall  1981).  S.  ochrognathus  was 
recovered  from  a  wood  rat  midden  near  the  Santa  Catalina  Mountains  (Mead  et  al. 
1983).  The  genus  has  a  rich  fossil  record  throughout  the  Southwest  (Lindsay  and 
Tessman  1974,  Harris  1977). 

Neotoma  a/Z7/gw/a— White-throated  wood  rat 

Material.  — I.  mandibles  (3);  R  mandibles  (4);  L  maxillae  (3);  R  maxillae  (7);  LM, 
(7);  RM,  (8);  LM"  (11);  RM'  (15);  MH55);  M:U15). 

Remarks.— A\\  the  Neotoma  remains  compare  well  with  N.  alhigula.  The  occlusal 
patterns  for  adult  teeth  are  well-rounded  as  in  A^.  alhigula  and  A^.  lepida  (Desert  wood 
rat),  but  the  teeth  are  much  larger  than  those  of  modem  A^.  lepida.  The  anterolingual 
re-entrant  on  the  M,  are  very  shallow  as  in  A^.  alhigula  compared  to  the  deep,  microtine- 
like  dental  characters  of  A^.  mexicana  (Mexican  wood  rat). 

Five  species  of  Neotoma  presently  live  in  Arizona  (Colorado  Plateau),  more  than 
in  any  other  state.  Only  A^.  lepida,  N.  alhigula,  and  A^.  mexicana  inhabit  southern 
Arizona  today,  and  only  the  last  two  presently  occur  in  the  vicinity  of  Deadman  Cave. 
A^.  alhigula  lives  in  desert-grassland  and  desertscrub  habitats  while  N.  mexicana  occurs 


268 


Table  2.     Measurements  of  modem  and  fossil  dentaries  (Deadman  and  Rampart  caves)  of  Bassariscus 
astutiis.  Measurements  are  rounded  to  the  nearest  0.5  mm. 


n 

OR 

A' 

Deadman  Cave 

Modem  Arizona 

Rampart  Cave,  Arizona  (northem) 

Alveolar  length  P4-M, 

1 
13 

1 

24.0 

21.0-23.0 

27.9 

24.0 
22.0 
27.9 

Deadman  Cave 

Modem  Arizona 

Rampart  Cave,  Arizona  (northem) 

Alveolar  length  Mi 

1 
13 

1 

8.0 

6.5-7.5 
6.5 

8.0 
7.0 
6.5 

in  higher  woodland  and  forest  areas.  Neotoma  stephensi  (Stephen's  wood  rat)  occurs 
in  the  northem  half  of  the  state  and  A'',  cinerea  (Bushy-tailed  wood  rat)  in  the  north- 
eastern sector  (Cockrum  1982).  The  midden  of  the  wood  rat  is  found  in  numerous  dry 
localities  throughout  the  Southwest  and  is  radiocarbon  dated  back  to  more  than  40  000 
B.P.  (Van  Devender  1977,  Van  Devender  and  Spaulding  1979). 

Microtus  species— Meadow  vole 

Material.-L  mandible  (15043);  LM,  (2:  15044);  RM,  (15045);  RM^  (3:  15046). 

Remarks.— We  have  not  identified  the  fossil  teeth  to  the  specific  level.  We  find  it 
difficult  to  differentiate  M.  longicaudus  (Long-tailed  vole)  from  M.  montanus  (Montane 
vole)  using  isolated  molars.  Of  the  three  complete  fossil  MjS  examined,  two  had  four 
closed  alternating  triangles  and  one  had  five  triangles. 

None  of  the  four  species  of  vole  found  in  Arizona  presently  occur  near  Deadman 
Cave.  Microtus  mexicanus  (Mexican  vole)  presently  occurs  in  the  mountainous  region 
of  eastern  Arizona  but  may  have  had  a  wider,  more  western  distribution,  based  upon 
the  present  isolated  occurrence  of  M  m.  hualpaiensis  in  northwestern  Arizona,  in  the 
Late  Pleistocene  and/or  Early  Holocene  (Hall  1981).  The  Mexican  vole  was  also  iden- 
tified in  the  fossil  remains  from  Papago  Springs  Cave  (Skinner  1 942).  Microtus  mon- 
tanus (Montane  vole)  has  a  predominantly  northwestern  distribution  in  the  United 
States.  Its  nearest  occurrence  to  Deadman  Cave  is  in  the  Arizona  Strip  region  of 
northernmost  Arizona  and  in  east-central  Arizona.  Microtus  pennsylvanicus  (Meadow 
vole)  lives  mainly  in  northem  and  eastem  North  America  but  approaches  Arizona  in 
northwestem  New  Mexico.  The  Meadow  vole  may  have  had  a  more  southem,  moun- 
tainous distribution  in  the  Late  Pleistocene  or  Early  Holocene  based  upon  an  isolated 
modem  population  in  northwestem  Chihuahua,  Mexico  (Bradley  and  Cockrum  1968). 
Microtus  longicaudus  occurs  through  much  of  westem  North  America,  including  north- 
eastem  Arizona.  An  isolated  population  presently  lives  in  the  Pinaleiio  Mountains  only 
60  km  east  of  Deadman  Cave.  Based  on  present  geographic  distributions,  this  species 
seems  most  likely  to  have  inhabited  the  mountains  of  the  Basin-and-Range  province 
of  southeastern  Arizona  during  the  Late  Pleistocene  and  Early  Holocene. 

Order  CARNIVORA-Camivores 

Family  Procyonidae— Racoons,  coatis,  and  ringtails 

Bassariscus  a^rw/w^— Ringtail 

Material. -K  mandible  P.-.,  (15062);  R  maxilla  (15315);  LP^  (15316). 

Remarks.— The  fossil  Ringtail  specimen  compares  well  with  modem  specimens 
except  that  the  M2  is  not  developed  in  the  fossil  specimen,  but  is  replaced  by  a  distinct 
depressional  scar  where  the  tooth  was  to  have  developed.  The  alveolar  length  from  P4 
to  M,  (Table  2)  is  slightly  longer  in  the  Deadman  Cave  specimen  than  in  modem 
specimens.  The  mandible  from  Deadman  Cave  does  not  seem  to  be  similar  to  the  rami 


269 


Table  3.     Measurements  of  dentaries  (modem  and  Deadman  Cave)  of  Spilogale  putorius.  Measurements 
are  rounded  to  nearest  0.5  mm. 


n 

OR 

.V 

Deadman  Cave 

Modem  Arizona 

Modem  Nevada  (northern) 

Alveolar  length  Pj-Mj 

5 
7 

1 

15.5-18.0 

12.5-14.5 

15.8 

16.5 
13.5 
15.8 

Deadman  Cave 

Modem  Arizona 

Modem  Nevada  (northern) 

Alveolar  length  P^-M' 

1 
8 

1 

14.0 

11.5-13.0 

12.0 

14.0 

12.5 
12.0 

described  as  B.  sonoitensis  from  Papago  Springs  Cave  (Skinner  1 942).  Late  Pleistocene 
and  Holocene  localities  of  the  Ringtail  are  shown  in  Mead  and  Van  Devender  (1981). 
The  Ringtail  is  widespread  in  rocky  habitats  in  the  desert  grassland,  and  woodlands 
of  the  Southwest,  and  lives  today  in  Deadman  Cave.  Numerous  modem  scats  are 
located  throughout  the  cave,  especially  near  the  entrance  as  demonstrated  by  the  ac- 
cumulations of  seeds,  insects,  and  bones. 

Family  Mustelidae— Weasels,  skunks,  and  badgers 
Spilogale  /7wror/"w5— Spotted  skunk 

Ma/ma/.  -L  mandibles  (3;  1 5047-1 5049);  R  mandibles  (6;  1 5050-1 5055);  L  max- 
illa (15056);  R  maxillae  (2;  15057-15058);  L  humerus  (15059). 

Remarks.— T\\Q  specimens  from  Deadman  Cave  are  consistently  larger  than  mod- 
em specimens  of  S.  putorius  (Table  3).  The  Spotted  skunk  is  common  throughout  the 
Southwest  and  can  be  found  in  the  Santa  Catalina  Mountains  down  to  the  lower  desert 
mountain  ranges.  Elsewhere,  fossil  remains  of  the  Spotted  skunk  have  been  recovered 
from  Arizona  (Skinner  1942),  Califomia  (Stock  1930),  and  New  Mexico  (Harris  1977). 

Mephitis  macroura— Hooded  skunk 

Material. -L  mandibles  (2;  15060);  L  maxilla  (15061);  L  humerus  (15317). 

Remarks.— The  left  mandible  compares  most  favorably  with  that  of  Mephitis 
macroura,  being  smaller  than  in  M.  mephitis  (Striped  skunk)  or  Conepatus  mesoleucus 
(Hog-nosed  skunk)  yet  definitely  larger  than  in  Spilogale.  All  three  skunks  occur  in 
southern  Arizona,  including  the  Santa  Catalina  Mountains.  Mephitis  macroura  has  its 
present  northem  distribution  in  southem  Arizona  and  New  Mexico  (Hall  1981).  We 
know  of  no  other  Late  Pleistocene-Early  Holocene  record  of  this  taxon. 

Family  Felidae— Cats 
Felis  coAzco/or— Mountain  lion 

Material.-La  (15063);  LC,  (15064);  RC,  (15065). 

Remarks.— AM  the  canines  compared  well  with  F.  concolor  rather  than  Panthera 
leo  atrox  (American  lion)  which  has  been  recovered  from  Late  Pleistocene  age  deposits 
of  the  nearby  Murray  Springs  site  (Haynes  1968,  J.  J.  Saunders  personal  communi- 
cation). The  canines  were  compared  in  size  to  those  of  modem  Arizona  Mountain  lions 
and  fossil  specimens  from  Rancho  La  Brea,  Califomia,  El  Durado,  Colorado,  Tule 
Springs,  Nevada  (Kurten  1973),  and  Rampart  Cave,  Arizona  (Table  4).  There  were  no 
discernible  differences  other  than  that  the  Late  Pleistocene  Felis  concolor  canines  from 
Rancho  La  Brea  may  have  been  slightly  larger  than  those  from  Deadman  Cave. 

The  Mountain  lion  occurs  historically  and  paleontologically  throughout  the  South- 
west, including  the  Santa  Catalina  Mountains  (Hall  1981).  Although  it  is  usually  found 
in  woodlands  and  forest  country,  it  also  lives  in  mgged  ranges  well  within  the  Sonoran 
Desert. 


270 


Table  4.  Length  (L;  anterior-posterior)  and  breadth  (B;  labial-lingual)  range  of  measurements  of  canines 
in  modem  and  Late  Pleistocene-Early  Holocene  Mountain  lion  (Felis  concolor).  Recent  specimens  from 
Arizona  (44,  654,  22785,  and  23444;  Department  of  Ecology  and  Evolutionary  Biology,  University  of 
Arizona,  Tucson).  Measurements  (in  millimeters)  of  fossil  specimens,  other  than  from  Deadman  and 
Rampart  caves,  from  Kurten  (1976). 


n  = 

C 

c, 

L 

B 

L 

B 

Recent  Arizona 
Deadman  Cave,  Arizona 
Rampart  Cave,  Arizona 
Rancho  La  Brea,  California 
El  Dorado,  Colorado 
Tule  Springs,  Nevada 

4 
3 
1 
3 

2 

1 

10.9-13.2 
11.3 
12.8 

12.6 

9.4-11.9 

10.3 

9.9 

11.0 
9.8 

10.8-13.2 
11.0-11.3 

12.4-16.4 
12.0 

8.0-9.8 
8.2-8.6 

11.2-12.5 
9.8 

Order  PERISSODACTYLA- Odd-toed  ungulates 

Family  Equidae— Horses 

Equus  species— Horse 

Material  — 2nd  phalanx  (15027). 

Remarks.  — This  single  element  of  a  horse  could  not  be  identified  to  species.  Re- 
mains of  extinct  species  of  Horse  are  common  throughout  the  Southwest  (Stock  1930, 
Mawby  1967,  Lindsay  and  Tessman  1974,  Harris  1977,  Harris  and  Porter  1980,  Kurten 
and  Anderson  1980).  Skinner  (1942)  identified  E.  conversidens  and  E.  tau  from  Papago 
Springs  Cave. 

Order  ARTIODACTYLA— Even-toed  ungulates 

Family  Cervidae— Cervids 

Odocoileus  species— Deer 

Material.-l.V2  {\5Q>66). 

Remarks.— The  Deadman  Cave  specimen  of  Odocoileus  could  not  be  identified  to 
species.  Today  in  southern  Arizona  O.  hemionus  (Black-tailed  deer)  lives  in  the  lowland 
habitats  while  O.  virginiana  (White-tailed  deer)  is  found  on  the  mountain  tops.  Fossil 
remains  of  both  species  of  deer  are  widespread  in  North  America  (Kurten  and  Anderson 
1980),  although  only  a  few  Late  Pleistocene  localities  in  Arizona  contain  remains  of 
deer  (Lindsay  and  Tessman  1974,  Mead  1981). 

Family  Bovidae— Bovids 

Euceratherium  collinum-Shvub-ox 

Material.  —  L  mandible  (15318). 

Remarks.— The  single  specimen  of  the  extinct  Shrub-ox  was  identified  by  Walter 
Dalquest  and  Ernest  Lundelius  (Dalquest  1981,  personal  communication).  The  occlusal 
surface  of  the  teeth  were  very  worn,  indicating  an  old  individual.  This  specimen  is  the 
first  record  of  the  Shrub-ox  in  Arizona.  Kurten  and  Anderson  (1980)  describe  this 
bovid  as  a  large,  specialized  grazer  that  probably  lived  in  the  lower  foothills  (like  at 
Deadman  Cave)  rather  than  in  the  high,  forested  mountains.  Euceratherium  collinum 
is  also  known  from  Burnet  Cave,  New  Mexico  (Schultz  and  Howard  1935),  but  is  not 
a  common  component  of  Late  Pleistocene  faunas  in  the  Southwest  (Kurten  and  An- 
derson 1980). 


Discussion 


Localities 


Although  there  are  numerous  Late  Pleistocene  age  localities  in  Arizona,  especially 
in  the  southern  portion  (Lindsay  and  Tessman  1974),  most  of  these  sites  are  isolated 


271 


finds  in  alluvial  deposits  containing  Mammuthus  jeffersoni  (Jefferson's  mammoth;  =M. 
columbi,  fide  Kurten  and  Anderson  1980),  Camelops  species  (camel),  Equus  species, 
or  Bison  species  (bison). 

The  most  current  work  in  Arizona  concerning  Late  Pleistocene  (Wisconsinan)  age 
deposits  comes  from  the  well-preserved,  radiocarbon  dated  wood  rat  middens,  but 
these  localities  rarely  contain  the  larger  animals  (Van  Devender  and  Mead  1978,  Mead 
198 1,  Mead  et  al.  1983).  The  only  cave  faunas  studied  in  southern  Arizona  are  Papago 
Springs  Cave  (Skinner  1942)  and  Ventana  Cave  (Haury  1950).  Ventana  Cave  (145  km 
west  of  Deadman  Cave,  Fig.  1)  was  excavated  primarily  for  its  abundant  archaeological 
remains.  Two  lower  units,  Volcanic  and  Conglomerate,  were  deposited  during  the 
Wisconsinan  glacial  episode  (Haury  1950).  Papago  Springs  Cave  (112  km  south  of 
Deadman  Cave,  Fig.  1)  was  excavated  solely  for  its  Late  Pleistocene  vertebrate  fossils, 
which  were  laborously  chiseled  from  brecciated  layers  along  the  walls  and  ceiling  of 
the  cave  (Skinner  1942). 

The  two  cave  faunas  are  very  different  in  composition  (Table  1).  The  fourteen  taxa 
recovered  from  the  lower  units  in  Ventana  Cave  were  all  mammals,  whereas  a  bird 
and  32  mammals  were  recovered  from  Papago  Springs  Cave.  Only  Canis  latrans  (=C. 
caneloensis\  Coyote),  Lepus  californicus,  and  Taxidea  taxus  (Badger)  are  shared  in  both 
faunas.  Of  the  64  taxa  from  Deadman  Cave  (5  amphibians,  25  reptiles,  12  birds,  and 
22  mammals),  only  Nothrotheriops  shastensis  is  shared  with  Ventana  Cave  and  five 
species  {Spermophilus  variegatus,  Thomomys  cf.  T.  bottae,  Neotoma  albigiila,  Antro- 
zous  pallidus,  and  Spilogale  putohus)  are  shared  with  Papago  Springs  Cave.  Each  cave 
is  in  a  different  physiographic  setting  in  southern  Arizona  and  had  different  modes  of 
fossil  accumulation.  Deadman  Cave  is  a  limestone  cave  on  a  large  mountain  mass 
between  two  major  river  valleys  surrounded  by  Sonoran  and  Chihuahuan  desertscrub 
communities;  its  fauna  was  collected  predominantly  by  small  carnivores  and  raptors. 
Ventana  Cave  is  a  volcanic  rockshelter  in  a  small,  low-elevation  desert  mountain  range 
presently  surrounded  by  the  hot,  dry  Sonoran  Desert.  The  open  cave  provided  some 
shelter,  easy  access,  as  well  as  water  from  a  spring.  Papago  Springs  Cave  was  an  open 
limestone  cave  in  rolling  oak  woodland  in  the  Canelo  Hills  of  southeastern  Arizona. 

The  Late  Wisconsinan  age  assignment  of  the  deposition  of  the  Volcanic  and  Con- 
glomerate units  in  Ventana  Cave  is  presumably  correct  (Haury  1950,  see  Long  and 
Muller  1981).  The  fossils  from  Papago  Springs  Cave  are  only  broadly  assigned  to  the 
Late  Pleistocene,  but  for  several  reasons  we  suggest  that  the  Papago  Springs  Cave  deposit 
was  formed  prior  to  the  last  glacial  maximum  in  the  Wisconsinan  (>22  000  B.P.).  The 
chamber  within  the  cave  has  changed  configuration  greatly  since  the  initial  deposition 
of  the  fauna.  A  great  abundance  of  rock  rubble  (10  m  thick)  with  bones  has  filled  the 
cavern  and  become  cemented.  According  to  Skinner  (1942)  the  cave  was  sealed  off 
from  the  outside  for  a  period  of  time,  allowing  settling  and  cementing  of  the  fossils  to 
take  place.  A  third  stage  in  the  history  of  the  cave  reopened  the  cavern  entrances 
permitting  recent  faunal  accumulations  and  partial  erosion  of  the  fossil  deposits.  We 
feel  that  such  an  accumulation  may  have  required  a  few  ten's  of  thousands  of  years, 
placing  the  time  of  deposition  sometime  in  the  Middle  or  Early  Wisconsinan  glacial 
episode. 

Fauna 

Five  species  of  amphibians  (all  anurans)  were  identified  from  Deadman  Cave 
(Table  1).  Of  these,  only  Bufo  punctatus  and  Scaphiopus  couchi  were  reported  prior  to 
this  report  from  the  Late  Pleistocene  and  Early  Holocene  of  Arizona.  We  report  Bufo 
cf.  B.  woodhousei,  Scaphiopus  cf.  5.  hammondi,  and  Rana  sp.  for  the  first  time  as 
fossils  in  Arizona,  although  they  have  been  previously  recorded  from  Late  Pleistocene 
localities  in  California,  Nevada,  and  New  Mexico.  Except  for  B.  woodhousei,  all  anurans 
from  the  Deadman  Cave  fauna  are  found  nearby  the  cave  today. 

Reptiles  are  better  known  than  the  amphibians  from  Late  Pleistocene-Early  Ho- 
locene deposits  in  Arizona.  We  report  25  reptiles,  including  13  species  of  lizards  and 
1 2  species  of  snakes,  from  Deadman  Cave.  Of  the  lizards,  Callisaurus  draconoides, 


272 


Holbrookia  maculata,  H.  texana,  Phrynosoma  douglassi,  P.  modestum,  P.  solare,  Sce- 
loporus  cf.  S.  clarkii,  and  Urosaurus  ornatus  have  not  been  reported  previously  from 
the  Late  Pleistocene  of  Arizona.  The  records  of  Heloderma  suspectum  from  Vulture 
Cave,  Arizona,  and  Gypsum  Cave,  Nevada,  may  be  Early  or  Middle  Holocene  in  age, 
and  therefore  this  species  perhaps  should  be  added  to  the  above  list.  Mdst  of  these 
lizards  have  previous  fossil  records  from  sites  in  California,  Nevada,  and/or  New 
Mexico.  Callisaurus  draconoides  is  reported  for  the  first  time  as  a  Late  Wisconsinan- 
Early  Holocene  fossil.  Phrynosoma  modestum  and  C.  draconoides  do  not  presently 
inhabit  the  region  of  Deadman  Cave  nor  the  immediate  valley.  The  closest  occurrence 
of/*,  modestum  is  in  Chihuahuan  desert-grassland  95  km  east  of  the  cave.  Callisaurus 
draconoides  occurs  just  west  of  Deadman  Cave  in  lower  Sonoran  desertscrub.  The  other 
lizards  from  Deadman  Cave,  as  well  as  most  of  the  snakes,  now  live  within  a  raptor's 
hunting  range  of  the  fossil  locality.  Of  the  1 2  species  of  snakes  identified  from  Deadman 
Cave,  only  Gyalopium  canum  has  not  been  previously  reported  as  a  fossil  from  Arizona, 
and  is  out  of  its  present  distributional  range  at  Deadman  Cave. 

The  birds  from  Deadman  Cave  represent  only  the  fifth  Late  Pleistocene  avifauna 
to  be  reported  from  Arizona.  This  contrasts  markedly  with  the  adjacent  states  of  New 
Mexico  and  California,  each  of  which  has  an  excellent  record  of  Late  Pleistocene  birds. 
The  Deadman  Cave  avifauna  is  especially  significant  in  that  the  diverse  Sonoran  Desert 
avifauna  has  yielded  very  few  Pleistocene  fossils. 

The  Deadman  Cave  fauna  contains  several  new  avian  fossil  records,  none  of  which 
is  unexpected.  Micrathene  whitneyi  and  Catharus  guttatus  are  reported  for  the  first 
time  as  fossils,  while  Cyrtonyx  montezumae  is  recorded  paleontologically  for  the  first 
time  anywhere  in  the  United  States.  New  Arizonan  fossil  records  include  Colinus 
gambelli,  Colinus  species,  Zenaida  cf.  Z.  macroura,  Otus  species,  Asio  otus,  the  in- 
determinate caprimulgid,  icterine,  and  Colaptes  auratus. 

Mammals  are  the  best  known  group  of  fossil  vertebrates  from  Arizona.  Twenty- 
two  mammals  are  identified  from  Deadman  Cave,  most  of  which  can  be  found  in  the 
Santa  Catalina  Mountains  today  or  in  the  surrounding  valleys.  Exceptions  are  the  three 
large  extinct  species  {Equus  species,  Nothrotheriops  shastensis,  and  Euceratherium  col- 
linum)  and  Microtus  species  which  lives  60  km  to  the  east  of  Deadman  Cave. 

Paleoenvironment  of  Deadman  Cave 

The  amphibian  remains  indicate  that  the  region  around  Deadman  Cave  was  at 
least  as  moist  at  the  time  of  deposition  as  it  is  today.  Certainly  pools  of  water  were 
nearby  as  they  are  today  in  portions  of  the  Santa  Catalina  Mountains  and  along  the 
major  streams  and  rivers.  The  lizard  fossils  argue  for  either  a  community  that  was  a 
composite  of  today's  vegetational  communities,  or,  a  few  vegetation  groups  abutting 
each  other  in  close  proximity  to  the  cave.  The  presence  of  Phrynosoma  modestum 
suggests  that  a  desert-grassland  area  may  have  occurred  on  the  broad  flat  hills  above 
the  large  river  valley,  at  the  base  of  the  mountain  mass.  Callisaurus  draconoides, 
Holbrookia  texana,  Sceloporus  magister,  and  Heloderma  suspectum  indicate  a  more- 
or-less  open  desertscrub  to  desert-grassland.  Such  areas  could  easily  abut  an  open 
woodland.  This  community  would  probably  be  in  the  lower  part  of  the  bajadas  between 
hills.  The  rest  of  the  lizard  fauna  indicates  several  vegetation  communities  ranging 
from  desert  and  desert-grassland  to  woodland.  Certainly  some  areas  of  talus  or  rock 
outcrops  were  nearby.  Most  of  the  snake  population  from  Deadman  Cave  suggests  little 
to  distinguish  the  region  then  from  what  it  is  today,  although  Gyalopium  canum  in- 
dicates a  cool  grassland  habitat. 

Except  for  the  possibly  extinct  icterid,  the  entire  avifauna  of  Deadman  Cave  con- 
sists of  living  taxa  that  occur  today  in  southern  Arizona.  No  birds  in  the  fauna  are 
restricted  to  coniferous  habitats.  In  fact,  Colinus  gambelli  and  Micranthene  whitneyi 
argue  strongly  against  any  sort  of  adjacent  coniferous  forest.  Colinus  gambelli  suggests 
the  presence  of  desertscrub,  while  Cyrtonyx  montezumae  is  characteristic  of  open  oak 
woodlands,  and  pine-oak  woodlands.  Cyrtonyx  is  the  only  bird  from  Deadman  Cave 


273 


that  is  not  found  at  least  occasionally  in  desertscrub  today,  although  Turdus  migratorius 
and  Catharus  guttatus  are  certainly  more  abundant  in  wooded  areas  than  in  desertscrub, 
and  never  nest  in  the  latter.  The  fossil  birds  from  Deadman  Cave  suggest  a  slightly 
more  grassy  and  wooded  condition  in  the  Latest  Pleistocene  and  Earliest  Holocene 
than  today,  such  as  an  evergreen  oak  grassland  mixed  with  desertscrub  and  desert- 
grassland  on  the  more  xeric  exposures,  and  oak  woodland  on  more  protected,  mesic 
areas. 

The  mammals  generally  indicate  an  open  woodland  with  some  areas  more  vege- 
tated and  other  areas  more  xeric  and  open.  None  of  the  mammals  are  restricted  to 
forested  habitats,  although  many  of  them  do  occur  today  in  the  forested  higher  ele- 
vations of  nearby  larger  mountains.  Notiosorex  crawfordi,  Microtus  sp.,  Reithro- 
dontomys  montanus,  and  Euceratherium  collinum  may  argue  for  an  open,  possibly 
grassy  woodland.  Today  this  sort  of  habitat  occurs  just  a  few  hundred  meters  upslope 
from  Deadman  Cave.  Perognathus  cf  P.  flavus,  Dipodomys  spectabilis,  Thomomys  cf 
T.  bottae,  and  Sigmodon  cf  S.  arizonae  indicate  that  a  desertscrub  to  desert-grassland 
was  nearby. 

No  distinctly  boreal  or  mesic  mammals  (e.g.,  Sorex  ornatus,  Ornate  shrew;  S. 
palustrus,  Water  shrew;  Ochotona  princeps,  Pika;  Marmota  flaviventris,  Yellow-bellied 
marmot;  Glaucomys  sabrinus,  Northern  flying  squirrel)  were  recovered  from  Deadman 
Cave.  This  may  be  because  the  faunal  assemblage  is  of  Late  Wisconsinan-Early  Ho- 
locene transition  in  age,  and  the  boreal-mesic  (full  glacial)  elements  had  already  become 
extirpated  locally.  This  may  be  the  case  with  the  Spermophilus  lateralis  from  Ventana 
Cave.  An  alternative  is  that  most  of  these  elements  never  did  occur  in  southern  Arizona 
during  the  last  Wisconsinan  full  and  late  glacial  (assuming  that  the  Papago  Spring  Cave 
deposit  is  Middle  Wisconsinan  or  older  in  age).  Presumably  if  any  geographically  or 
ecologically  extralocal  small  animals,  such  as  listed  above,  were  contemporaneous  with 
the  Shasta  ground  sloth,  Horse,  and  Shrub-ox,  they  too  would  have  been  observed  in 
the  Deadman  Cave  deposit. 

The  Deadman  Cave  faunal  record  is  very  similar  to  those  of  Early  Holocene-Late 
Pleistocene  wood  rat  faunas  in  the  Sonoran  Desert  (Van  Devender  and  Mead  1978, 
Mead  et  al.  1983)  and  Grand  Canyon  (Van  Devender  et  al.  1977,  Mead  and  Phillips 
1981)  in  that  the  small  vertebrates  were  conservative  with  few  animals  out  of  their 
present  range,  although  there  was  a  greater  change  in  the  local  vegetation.  The  pollen 
record  at  Willcox  Playa,  80  km  east  of  the  Santa  Catalina  Mountains,  recorded  a  pine 
forest  at  1 200  m  elevation  20  000  years  ago  in  an  area  that  now  supports  desert-grassland 
(Martin  1963).  This  is  an  estimated  lowering  of  the  vegetation  zones  by  about  1000  m 
elevation.  The  wood  rat  midden  record  for  lower  areas  in  the  Sonoran  Desert  in 
southwestern  Arizona  recorded  a  complex  elevational  lowering  of  600  m  or  less  of 
certain  woodland  plants  into  the  desert.  An  equivalent  lowering  of  vegetation  zones  in 
the  Santa  Catalina  Mountains  would  imply  a  shift  from  the  modem  desert-grassland- 
oak  woodland  vegetation  to  Pondersoa  pine  mixed  conifer  forest  or  a  Mexican  pine- 
oak  woodland,  depending  on  the  distribution  and  abundance  of  precipitation  in  the 
Late  Wisconsinan  (Whittaker  and  Niering  1968).  In  the  Early  Holocene  a  Mexican 
pine-oak  woodland  was  probably  near  the  site  until  about  8000  years  ago  with  the 
grassland  developing  more  recently.  Like  the  Microtus  sp.,  a  population  of  Abies 
lasiocarpa  (Corkbark  fir)  presently  on  the  top  of  the  Santa  Catalina  Mountains,  is  also 
isolated  from  its  nearest  population  in  the  spruce  forests  of  the  Pinalefio  Mountains. 
Most  of  the  remaining  Deadman  Cave  fauna  would  be  found  in  some  sort  of  open 
woodland  today.  Several  animals  including  Callisaurus  draconoides,  Heloderma  sus- 
pectum,  Dipodomys  spectabilis,  Phrynosoma  modestum,  and  Colinus  gambelli  do  not 
live  in  Mexican  pine-oak  woodland  today  and  are  more  likely  to  occur  in  open  desert- 
grassland  or  desertscrub  and  may  represent  the  Early  Holocene  portion  of  the  Deadman 
Cave  deposit.  Another  possibility  which  has  been  demonstrated  for  a  few  animals  in 
the  Sonoran  Desert  (Van  Devender  and  Mead  1978;  Mead  et  al.  1983)  is  that  under 
an  equable  Late  Wisconsinan  climate  certain  animals  now  confined  to  deserts  were 
able  to  live  in  more  diverse  woodland  communities. 


274 


Conclusions 

The  vertebrate  fauna  of  Deadman  Cave  includes  5  amphibians,  25  reptiles  (13 
lizards  and  12  snakes),  12  birds,  and  22  mammals  for  a  total  of  64  species.  The 
implication  from  this  faunal  assemblage  is  that  by  the  end  of  the  late  glacial  and  the 
beginning  of  the  post  glacial  (8000-12  000  B.P.),  most  of  the  local  fauna  was  essentially 
as  it  is  today— modem.  Only  one  amphibian  {Bufo  woodhousei),  three  reptiles  (Calli- 
saurus  draconoides,  Phrynosoma  modestum,  Gyalopium  canum),  and  one  mammal 
{Microtus  species)  are  locally  extirpated,  although  all  still  occur  in  southern  Arizona. 
Animals  unequivocally  extinct  are  the  Shrub-ox,  Horse,  and  Shasta  ground  sloth,  all 
large  mammals.  An  unidentified  icterine  bird  may  prove  to  be  an  extinct  species. 
Overall,  the  Deadman  Cave  fauna  suggests  that  the  vegetation  community  of  the  Late 
Pleistocene  and  Early  Holocene  were  rather  similar  to  those  found  today  in  the  same 
region. 

The  Deadman  Cave  bone  assemblage  has  expanded  our  knowledge  of  the  Late 
Pleistocene-Early  Holocene  fauna  of  southern  Arizona,  and  has  provided  new  questions 
on  Late  Pleistocene  zoogeography  of  the  hot  deserts.  Further  stratified  and  datable  cave 
deposits  and  wood  rat  middens  need  to  be  studied  to  refine  when  the  faunas  of  southern 
Arizona  became  "modem."  Upland  localities  need  to  be  studied  to  determine  whether 
boreal  and/or  mesic  faunal  elements  ever  existed  in  the  Late  Wisconsinan  of  southern 
Arizona,  and  whether  the  presently  extralocal  and  local  faunal  assemblages  ever  co- 
existed. 

Acknowledgments 

Foremost  we  thank  William  Peachey,  a  caver  from  Tucson,  Arizona,  for  finding 
the  bone  deposit  in  Deadman  Cave  and  for  realizing  the  cave's  importance  to  vertebrate 
paleontology.  Walter  Dalquest  and  Ernest  Lundelius  provided  the  identification  of  the 
Shrub-ox,  Greg  McDonald  verified  the  ground  sloth  identification;  we  appreciate  their 
help.  We  thank  Mary  C.  McKitrick  for  preliminary  identification  of  certain  avian  fossils. 
Storrs  L.  Olson  and  the  staff  of  the  Division  of  Birds,  United  States  National  Museum 
of  Natural  History,  Smithsonian  Institution,  made  available  their  skeleton  collection 
for  comparative  purposes.  Our  identification  of  the  mammals  was  aided  by  the  use  of 
the  mammalogy  collection  and  the  helpful  suggestions  provided  by  E.  Lendell  Cockrum 
and  Yar  Partzchian,  Department  of  Ecology  and  Evolutionary  Biology,  University  of 
Arizona,  Tucson.  The  Grand  Canyon  National  Park  provided  us  with  certain  skeletons 
from  their  Study  Collection  for  comparative  purposes.  Gene  Hall  helped  prepare  some 
of  the  Deadman  Cave  fossils.  Austin  Long  of  the  Laboratory  for  Isotope  Geochemistry, 
University  of  Arizona,  Tucson,  provided  the  radiocarbon  date.  Financial  support  was 
provided  by  grants  from  National  Science  Foundation  to  Paul  S.  Martin,  University 
of  Arizona  (DEB75- 13944)  and  Thomas  Van  Devender  (DEB76- 19784),  and  from  the 
Smithsonian  Institution  (Predoctoral  Fellowship,  Scholarly  Studies  Program)  to  David 
Steadman.  Emilee  M.  Mead  drafted  the  figures  and  helped  in  various  aspects  of  field- 
work.  We  thank  Donald  K.  Grayson,  two  anonymous  reviewers,  and  Gregory  Pregill 
for  critiquing  and  editing  our  manuscript.  The  Institute  for  Quaternary  Studies  provided 
final  typing  services. 

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■*<'. 


TRANSACTIONS 
\  OF  THE  SAN  DIEGO 

•  Oo         SOCIETY  OF 
^/^?V/i       NATURAL  HISTORY 


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nmf:A5\ 


Volume  20  Num6§i'^pp.  277-300         20  November  1984 


A  Pliocene  Flora  from  Chula  Vista, 
San  Diego  County,  California 

Daniel  I.  Axelrod 

Department  of  Botany,  University  of  California,  Davis,  California  95616  USA 

Thomas  A.  Demere 

Department  of  Geology,  San  Diego  Natural  History  Museum,  P.O.  Box.  1390,  San  Diego,  California  921 12 
USA 

Abstract.  A  small  fossil  flora  from  the  marine  Upper  Pliocene  San  Diego  Formation  suggests  that 
the  adjacent  coastal  plain  was  then  covered  with  an  avocado-Monterey  pine-live  oak  woodland  as- 
sociated with  palm,  cottonwood,  willow,  and  sycamore  along  streams.  Fossil  digger  pine  apparently 
was  confined  to  drier,  warmer  sites  away  from  the  coast.  At  higher,  cooler  levels  farther  inland  were 
stands  of  fossil  Jeffrey  pine.  Precipitation  was  near  50-58  cm  over  the  lowlands,  increasing  to  about 
65  cm  in  stands  of  fossil  Jeffrey  pine  near  450-600  m.  The  fossil  avocado,  palm,  and  pine  (afl!".  Pinus 
radiata  var.  binata)  indicate  summer  rainfall,  consistent  with  the  Late  Pliocene  higher-than-present  sea 
surface  temperature.  Mean  annual  temperature  on  the  coast  was  approximately  1 6°C,  and  annual  range 
was  about  7-8°C,  equability  was  near  M  70  with  frost  absent  along  the  coast,  light  in  the  interior. 

Comparison  with  Pliocene  floras  in  northern  California  shows  that  the  Chula  Vista  flora  lived  in 
a  separate  floristic  province,  one  corresponding  with  cismonlane  southern  California  which  has  been 
a  distinct  floristic  province  since  at  least  the  Middle  Miocene.  Two  new  species  of  fossil  pine  are 
described:  Pinus  diegensis  new  species  (allied  with  the  living  P.  radiata  var.  binata)  and  P.  jeffreyoides 
new  species  (similar  to  the  living  P.  jeffreyi). 

The  Chula  Vista  flora  provides  new  evidence  regarding  the  evolutionary  history  of  Pinus  radiata 
populations,  and  further  insight  into  the  disjunct  distribution  of  taxa  in  the  montane  conifer  forests  of 
southern  California  and  Baja  California. 

Introduction 

The  recent  discovery  of  a  small  flora  of  Late  Pliocene  age  (ca.  3  m.y.)  at  Chula 
Vista,  in  southwestern  San  Diego  County,  California  (Fig.  1),  provides  preliminary 
information  on  the  late  Tertiary  flora,  vegetation,  and  climate  of  the  region.  The  flora 
lived  at  a  critical  time  in  Neogene  environmental  history,  one  characterized  by  the 
cooler,  moister  climate  that  followed  the  warm,  dry  episode  of  the  latest  Miocene  (5- 
6  m.y.)  (Axelrod  1980Z?).  The  climatic  transition,  which  heralds  the  build-up  of  ice 
sheets  in  Alaska  and  border  areas,  resulted  in  the  displacement  southward  of  relatively 
xeric  sclerophyllous  vegetation  with  numerous  Madrean  taxa.  and  their  replacement 
by  a  more  mesic  flora.  The  nearest  floras  of  comparable  age  are  in  central  and  northern 
California  (Fig.  2).  Analysis  of  the  Chula  Vista  flora  thus  provides  a  general,  though 
tentative,  basis  for  interpreting  the  regional  floristic  differences  that  arose  as  cooler, 
moister  climate  spread  southward  and  into  lower  altitudes  in  the  Late  Pliocene. 

Geology 

The  geology  of  the  classical  marine  Pliocene  San  Diego  Formation  at  San  Diego 
has  been  summarized  recently  (Demere  1983).  This  rock  unit  crops  out  over  a  broad 
area  which  includes  much  of  the  southwestern  portion  of  San  Diego  County,  and  the 
extreme  northwestern  comer  of  Baja  California,  Mexico.  The  San  Diego  Formation 


278 


Late     Cenozoic     Floras 
of    California 


50 

_J 


t 1 1 T" 


0  50  100  150  200  Kilomelers 


Figure  1 .     Late  Cenozoic  floras  to  which  reference  is  made. 


was  deposited  during  a  marine  transgression  of  the  Neogene  San  Diego  Basin,  which 
hke  other  onshore  sedimentary  basins  in  southern  California  (e.g.,  Ventura  Basin,  Los 
Angeles  Basin)  is  structurally  related  to  the  wrench  and  extensional  tectonics  of  the 
continental  borderland.  Deposition  began  during  the  Late  Pliocene  and  apparently 
continued  into  Early  Pleistocene  time,  accumulating  at  least  75  m  of  marine  and  9  m 
of  nonmarine  sedimentary  rocks.  The  overall  stratigraphic  sequence  suggests  a  succes- 
sive shallowing  and  filling  of  this  basin.  It  is  now  apparent  that  extensional  tectonics 
have  controlled  both  the  initial  deposition  as  well  as  the  present  outcrop  distribution 
of  this  rock  unit.  Numerous  high-angle  normal  faults  striking  north  to  northwest  cut 
the  area  into  a  series  of  small  fault  blocks  which  expose  different  portions  of  the  Pliocene 
section. 

In  an  attempt  to  correlate  between  the  fault  blocks,  Demere  (1983)  informally 
subdivided  the  San  Diego  Formation  into  a  "lower"  and  an  "upper"  member  using 


279 


D- 

SOUTH 

CENTRAL 

NORTH 

L/ 

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Figure  2.     Ages  of  Late  Cenozoic  floras  in  California. 


both  lithologic  and  paleontologic  criteria.  The  "lower"  member  is  characterized  by 
yellowish,  very  fine-grained,  micaceous,  massive,  friable  sandstones  with  locally  well- 
bedded  sequences  of  laminated  and  cross-bedded  sandstones,  pebble-to-cobble  con- 
glomerates and  well-cemented  shell  beds.  This  "lower"  member  is  richly  fossiliferous 
and  has  produced  the  bulk  of  the  marine  invertebrate  fauna  so  well  known  through 
the  work  of  Grant  and  Gale  (1931)  and  Hertlein  and  Grant  (1944,  1960,  1972).  In 
addition,  the  diverse  avifauna  (Howard  1949,  Miller  1956)  and  cetacean  assemblage 
(Barnes  1973,  1 976)  reported  from  the  San  Diego  Formation  have  been  largely  collected 
from  the  "lower"  member. 

Lithologically  the  "upper"  member  is  characterized  by  well-bedded  sequences  of 
pebble-to-cobble  conglomerate  containing  reworked  "Poway"  clasts,  well-cemented 
fossiliferous  sandstones,  and  medium-  to  coarse-grained  friable  sandstones.  Marine 
invertebrate  fossils  are  locally  common  in  this  member. 

In  terms  of  paleoenvironment,  the  "lower"  member  contains  a  middle-to-outer- 
shelf  molluscan  fauna  characterized  by  Patinopecten  healeyi,  Pecten  stearnsii,  Luci- 
noma  annulata  and  Opalia  varicostata.  In  contrast,  molluscs  from  the  "upper"  member 
indicate  deposition  in  littoral-to-inner-shelf  depths.  Characteristic  species  include  Pec- 


280 


ten  bellus,  Argopecten  hakei,  Niicella  lamellosa  and  Dendraster  ashleyi.  Both  members 
reflect  normal  marine  deposition  in  a  broad  coastal  embayment  probably  similar  to 
present-day  Monterey  Bay  along  the  central  California  coast. 

Barnes  (1976)  correlated  the  San  Diego  Formation  ("lower"  member)  with  the 
Blancan  North  American  land  mammal  stage.  This  was  based  on  the  occurrence  of  the 
horse  Equus,  and  is  supported  by  the  recent  discovery  at  Chula  Vista  of  teeth  referable 
to  the  bunodont  mastodon  {Stegomastodon  rexroadensis  Woodbume).  Relying  on  the 
stratigraphic  ranges  of  moUuscan  species  the  "lower"  member  is  correlative  with  the 
upper  portion  of  the  San  Joaquin  Formation  in  the  San  Joaquin  Basin,  the  Careaga 
Formation  in  the  Santa  Maria  Basin,  and  the  Niguel  Formation  and  the  upper  Fernando 
Formation  in  the  southeastern  Los  Angeles  Basin,  all  Late  Pliocene  in  age  (Woodring 
and  Bramlette  1951,  Vedder  1972).  In  turn  the  "upper"  member  of  the  San  Diego 
Formation  is  correlated  with  the  lower  Santa  Barbara  Formation  in  the  Ventura  Basin 
which  is  considered  to  be  Late  Pliocene  to  Early  Pleistocene  in  age  (Keen  and  Bentson 
1944,  Lajoie  et  al.  1982).  The  meager  microfossil  record  available  in  the  San  Diego 
Formation  (Ingle  1967,  Mandel  1973)  suggests  that  it  is  apparently  no  older  than 
planktonic  foraminiferal  zone  N-21  (approximately  3.0  million  years  B.P.  Late  Plio- 
cene) and  is  perhaps  as  young  as  the  Emiliania  annula  calcareous  nannoplankton 
subzone  (approximately  1.5  million  years  B.P.,  Early  Pleistocene). 

The  section  at  Chula  Vista  consists  of  approximately  73  m  of  fossiliferous  San 
Diego  Formation.  Here  the  Pliocene  marine  rocks  rest  disconformably  on  fluvial  and 
lacustrine  sedimentary  rocks  of  the  Middle  Miocene  Sweetwater  Formation.  Overlying 
the  San  Diego  Formation  along  an  irregular  erosion  surface  are  fluvial  and  alluvial 
sedimentary  rocks  of  the  Lower  Pleistocene  Sweitzer  Formation  (=Lindavista  For- 
mation of  Kennedy  1975).  The  lower  46  m  of  the  Chula  Vista  Pliocene  section  contain 
marine  invertebrate  taxa  characteristic  of  the  "lower"  member  of  the  San  Diego  For- 
mation. This  part  of  the  section  is  also  characterized  by  locally  common  marine  ver- 
tebrate fossils  and  has  produced  all  of  the  paleobotanical  material  described  in  this 
report. 

In  terms  of  general  stratigraphic  position,  the  described  fossil  pine  cones  were 
found  betwen  24  and  30  m  above  the  base  of  the  San  Diego  Formation,  the  fossil  leaves 
between  12  and  19  m,  and  a  few  poorly  preserved  shipworm  {Teredo)-bovQ6.  logs 
between  12  and  21  m. 

The  fragmentary  cones  occur  in  large,  elongate  (up  to  40  cm  in  diameter),  limy 
concretions  which  formed  around  large  bones  of  fossil  baleen  whales.  Steinkems  of 
marine  invertebrates  also  occur  in  these  concretions.  The  cones  are  preserved  as  natural 
internal  and  external  molds  and  vary  from  incomplete  three-dimensional  specimens 
to  only  partial  imprints. 

Fossil  leaves  occur  as  dense  concentrations  of  flat-lying  and  stacked  leaf  material 
in  both  fine-grained  sandstone  and  sandy  siltstone.  The  leaves  are  preserved  as  iron- 
stained  imprints  and  are  largely  fragmentary.  The  leaves  were  collected  from  thin  ( 1 5 
cm  thick)  sandstone  and  siltstone  strata  interbedded  with  fossiliferous  sandstones  con- 
taining typical  "lower"  San  Diego  Formation  marine  invertebrate  taxa  (e.g.,  Patino- 
pecten  healeyi  and  Lucinoma  annulatd).  The  occurrence  of  terrestrial  plant  material  in 
this  marine  setting  points  to  offshore  transport,  no  doubt  first  under  fluvial  and  then 
under  marine  conditions.  The  concentration  of  leaf  material  in  thin  stratigraphic  ho- 
rizons suggests  marine  transport  by  storm-generated  debris  flows.  Otherwise,  it  seems 
doubtful  that  they  would  be  concentrated  in  a  local  area.  The  cones  probably  were 
transported  in  a  similar  manner.  The  cone  fragments  are  the  result  of  partial  preser- 
vation in  the  concretions,  the  remaining  portion  of  the  cone  was  not  preserved  in  the 
soft  compacted  sandstones  that  enclose  the  concretions. 

Not  described  in  this  report,  but  occurring  within  the  Pliocene  section  at  Chula 
Vista,  were  a  number  of  severely  shipworm-bored,  calcareously  cemented  logs.  Some 
of  these  logs  were  up  to  3  m  in  length  and  had  broken  branch  stems.  They  all  were 
lying  parallel  to  the  bedding  upon  locally  pebbly  fossiliferous  sandstone. 


281 

Table  1.     Systematic  list  of  floral  species  from  the  San  Diego  Formation,  Chula  Vista,  California. 

Pinaceae  Salicaceae 
Pinus  diegemis  n.  sp.  Populus  alexanderi  Dorf 

Finns  jejfreyoides  n.  sp.  Salix  wildcatensis  Axelrod 

Pinus  pieperiDod  T^^g^c^^^ 

Arecaceae  Quercus  lakevillensis  Dorf 

Gen.  et  sp.  indet.  (Palm)  Lauraceae 

Juncaceae  Persea  coalingensis  (Dorf)  Axelrod 

Gen.  et  sp.  indet.  (Reed)  Platanaceae 

Platanus  paucidentata  Dorf 

Composition 

The  small  paleobotanical  collection  from  Chula  Vista  includes  ten  species  distrib- 
uted among  three  conifers,  two  monocotyledons,  and  five  dicotyledons.  Two  of  the 
pines  are  new.  The  monocotyledons  are  represented  by  specimens  too  incomplete  to 
refer  to  species. 

The  fossil  taxa  are  represented  by  only  a  few  specimens  each,  and  most  are  frag- 
mentary, reflecting  their  transport  into  the  offshore.  Fifteen  specimens  are  referred  to 
Persea.  Of  the  remainder,  Pinus  diegensis  is  represented  by  5  specimens,  Populus  4, 
Platanus,  Salix,  Pinus  Jeffrey oides  3  each,  and  Pinus  pieperi,  palm,  Quercus  and  reed 
by  one  each. 

All  of  the  fossils  are  similar  to  living  taxa  and  have  been  assigned  to  fossil  species 
following  nomenclatural  convention  (Axelrod,  1980Z?:205).  As  judged  from  their  re- 
lationships, the  flora  is  composed  of  8  trees,  one  shrub  and  one  herbaceous  perennial. 

An  understanding  of  the  ecology  of  extant  species  closely  allied  to  the  fossil  plants 
in  the  Chula  Vista  flora  provides  a  basis  for  reconstructing  the  paleovegetation  and 
climate  of  the  region.  These  allied  modem  species  can  be  separated  into  four  ecologic 
and  geographic  groups. 

First,  several  of  the  allied  living  species  are  widely  distributed  in  central  and 
southern  California.  These  include  sycamore  {Platanus),  cottonwood  {Populus)  and 
willow  {Salix).  Today,  these  trees  inhabit  streambanks  in  the  coastal  area  from  near 
San  Francisco  Bay  southward,  reaching  also  into  the  interior.  The  oak  {Quercus  agri- 
folia)  forms  dense  woodlands  in  moister,  more  equable  sites  near  the  coast  and  con- 
tributes to  open  oak-grass  communities  at  low  to  moderate  elevations  in  more  inland 
areas  of  the  outer  Coast  Ranges.  Quercus  is  also  found  frequently  along  stream  margins 
where  it  is  associated  with  cottonwood,  sycamore,  and  willow.  The  fossil  cottonwood 
{Populus  alexanderi)  is  similar  to  P.  trichocarpa  var.  trichocarpa  that  has  roundish- 
ovate  leaves  and  lives  near  the  coast  in  mild  equable  climate  from  the  San  Francisco 
Bay  region  southward.  It  differs  from  the  black  cottonwood  that  occurs  in  the  mountains 
of  California  and  northward,  which  has  larger,  ellipitic-ovate  leaves  and  is  best  referred 
to  P.  hastata  Dode. 

Second,  digger  pine  {Pinus  sabiniana)  characterizes  the  pine-oak  woodland-grass 
vegetation  of  the  inner  Coast  Ranges  and  lower  slopes  of  the  Sierra  Nevada.  It  reaches 
the  margin  of  southern  California  in  Santa  Ynez  Valley  north  of  Santa  Barbara  and  on 
Liebre  Mountain  several  miles  southeast  of  Tejon  Pass.  It  only  approaches  the 
coast  in  the  Santa  Lucia  Mountains  near  Gorda  where  it  seems  to  be  a  relict  of  the 
Xerothermic  period.  This  also  appears  to  account  for  its  disjunct  occurrence  in  Santa 
Ynez  Valley.  In  the  south  Coast  Ranges  it  inhabits  flats  adjacent  to  rivers,  notably  the 
Salinas,  Santa  Maria,  Santa  Ynez,  and  Sisquoc,  which  at  times  of  flood  probably 
transport  its  large  cones  to  the  coast.  Among  its  associates  in  riparian  sites  are  species 
of  cottonwood,  willow,  sycamore,  and  live  oak  similar  to  taxa  in  the  Chula  Vista  flora. 

Third,  Pinus  jeffreyi,  allied  to  the  fossil  P.  jeffreyoides  new  species,  now  occurs  in 


282 


Figure  3.     Guadalupe  Island  pine  {Pinus  radiata  var.  binata)  and  palm  {Brahea  edulis)  on  Guadalupe  Island 
have  allied  laxa  in  the  Chula  Vista  fossil  flora.  (Photo  by  Reid  Moran.) 


the  mountains  of  eastern  San  Diego  County.  The  stand  nearest  to  Chula  Vista  is  55 
km  northeast  at  Pine  Valley,  at  an  elevation  of  1050  m.  There  it  is  associated  with  live 
oak  (Quercus  agrifolia).  Streams  in  the  area  also  support  species  of  willow,  cottonwood 
and  sycamore  similar  to  those  in  the  fossil  flora.  The  bordering  slopes  are  covered  with 
dense  chaparral  composed  of  Adenostoma,  Arctostaphylos,  Ceanothus,  Cercocarpus, 
Quercus,  and  others.  It  is  to  be  noted  that  P.  jejfreyi  replaces  P.  ponderosa  as  the  lowest 
montane  conifer  forest  species  in  the  southern  Peninsular  Ranges.  Here  it  is  a  distinct 
ecotype  as  compared  with  P.  jejfreyi  in  the  Sierra  Nevada  which  is  a  member  of  the 
red  fir-white  pine-mountain  hemlock  forest  that  is  snowbound  all  winter.  Only  an 
occasional  light  snow  occurs  at  its  lower  elevations  in  San  Diego  County  and  in  the 
Sierra  Juarez  and  Sierra  San  Pedro  Martir,  northern  Baja  California,  where  it  also  meets 
oak  woodland  and  chaparral  vegetation,  generally  at  levels  near  1400-1500  m  (see 
Nelson  1922). 

Fourth,  three  Chula  Vista  fossil  species  are  allied  to  living  taxa  in  western  Mexico. 
Avocado  (Persea),  which  was  abundant  in  California  into  the  close  of  the  Tertiary 
(Sonoma,  Wildcat,  Turlock  Lake,  Broken  Hill,  Coalinga  floras),  now  occurs  from  south- 
em  Sonora  southward  in  the  Sierra  Madre  Occidental,  and  other  species  are  in  the 
eastern  United  States,  chiefly  along  the  Atlantic  seaboard.  In  southern  Sonora  and 
Sinaloa,  Persea  is  a  frequent  member  of  pine-oak  woodland  vegetation,  associated  with 
sycamore,  willow,  palm  and  other  riparian  taxa  in  valley  bottoms.  It  occupies  sheltered 
sites  at  1000-1200  m  under  mild,  equable  climate  (Gentry  1942,  1946). 

Cones  of  Pinus  diegensis  new  species  are  similar  to  those  of  P.  radiata  var.  binata 
of  Guadalupe  Island,  situated  350  km  off' the  central  coast  of  Baja  California  at  29°N 
latitude.  This  pine  is  confined  chiefly  to  the  middle-upper  slopes  of  the  island,  generally 
at  altitudes  above  800  m  where  it  is  in  a  persistent  fog  belt  during  summer.  The 
occurrence  of  P.  diegensis  on  the  mainland  as  recently  as  3  m.y.  ago  suggests  that  the 
living  P.  radiata  var.  binata  did  not  originate  in  insular  isolation  but  probably  was 
confined  to  Guadalupe  Island  as  drier  climate  spread  over  the  region. 

A  fragmentary  specimen  of  palm  frond  from  Chula  Vista  shows  relationship  to 


283 


^st^'-'-r 


Figure  4.     Present  environment  in  the  area  of  the  Chula  Vista  fossil  flora. 


Sabal  and  also  to  Brahea  (^Erythea).  Sabal  is  a  frequent  member  of  the  pine-oak 
woodland  in  the  Sierra  Madre  Occidental  of  Sonora  where  it  occurs  along  streams  with 
avocado,  willow,  sycamore  and  other  riparian  inhabitants.  Of  considerable  interest  is 
the  presence  of  Brahea  edulis  H.  Wendl.  ex.  S.  Wats,  on  Guadalupe  Island  where  it 
occurs  in  deep,  moist  canyons  with  Pinus  radiata  var.  binata  (Howell  1941Z);  Libby  et 
al.  1968),  as  seen  in  Fig.  3. 

Vegetation 

The  community  relations  of  modem  species  most  similar  to  the  fossil  plants  suggest 
that  during  the  Late  Pliocene  this  portion  of  the  coastal  strip  was  covered  with  dense 
woodland  vegetation  in  contrast  to  the  open  coastal  sage  of  today  (Fig.  4).  The  abun- 
dance of  avocado  leaves  in  the  collection  implies  that  Persea  dominated  stream  valleys 
and  sheltered  slopes  near  the  shore.  On  moister  slopes  it  was  associated  with  fossil 
Guadalupe  pine  and  probably  formed  a  dense  forest.  Live  oak  was  regularly  associated 
with  the  pine  and  avocado.  Along  stream  borders  these  trees  occurred  with  sycamore, 
Cottonwood,  willow,  and  palm.  Toward  the  interior,  fossil  Guadalupe  pine  {P.  diegensis) 
and  live  oak  probably  contributed  to  a  more  open  woodland-grass  community  that 
gradually  merged  with  a  fossil  digger  pine  association  that  included  avocado  and  palm 
in  sheltered  sites,  as  well  as  riparian  taxa.  Fossil  digger  pine  probably  attained  optimum 
development  on  exposed,  hotter  and  drier  south-facing  slopes  provided  by  granitic  and 
metamorphic  basement  rocks  farther  inland.  At  higher,  cooler  and  moister  levels  farther 
east  were  stands  of  fossil  Jeffrey  pine  that  reached  down  to  a  lower  altitude  than  at 
present  because  precipitation  was  higher. 


Climate 

Judging  from  conditions  under  which  allied  living  taxa  occur,  annual  precipitation 
at  sea  level  during  the  Late  Pliocene  probably  was  near  58-63  cm  as  compared  with 


284 


20 


^    '  -<^  I I I I I I I I I    r 


^-! I I L 


'?C/ 


Figure  5.  Estimated  paleotemperature  for  the  Chula  Vista  coastal  strip  (A),  the  area  of  fossil  digger  pine 
(B),  and  the  fossil  Jeffrey  pine  forest  (C).  The  difference  in  mean  annual  temperature  between  Chula  Vista 
today  and  the  inferred  paleotemperature  (circled  area.  A)  is  approximately  0.5°C.  The  radiating  lines  of 
warmth  (W)  indicate  growing  seasons  defined  by  the  number  of  days  (d)  with  mean  temperature  warmer 
than  the  specified  temperature  (i.e.,  W  14°C  has  183  days  with  mean  temperature  warmer  than  14°C).  The 
arcs  M  70  and  M  60  represent  a  measure  of  equability  which  decreases  in  all  directions  from  T  14°C  and  A 
0°C.  For  further  data  see  Bailey  (1960,  1964). 


22  cm  at  Chula  Vista  today.  Precipitation  increased  to  about  70  cm  in  hills  to  the  east 
at  the  lower  margin  of  fossil  Jeffrey  pine  forest.  The  remaining  taxa  in  the  fossil  flora 
could  readily  survive  under  these  rainfall  totals.  Fossil  avocado,  palm  and  Guadalupe 
pine  (var.  binatd)  indicate  summer  rainfall.  This  resulted  from  a  warmer  sea-surface 
temperature  than  that  of  today,  as  shown  by  the  marine  macro-invertebrate  fauna  of 
the  San  Diego  Formation.  Hertlein  and  Grant  (1954)  estimated  on  the  basis  of  the 
molluscan  taxa  that  winter  sea  surface  temperature  during  the  Late  Pliocene  was  similar 
to  that  now  near  Cedros  Island  (ca.  17-18°C),  or  fully  3-4°C  higher  than  that  in  the 
San  Diego  area  today  (U.S.  Navy  1956-58;  U.S.  Weather  Bureau  1938).  With  warmer 
water,  hurricanes  from  the  eastern  Pacific  may  have  moved  farther  up  the  coast  than 
they  now  do.  In  this  regard,  the  summer  of  1 983  in  southern  California  was  characterized 
by  warmer-than-usual  water  offshore  and  increased  warm  season  rainfall  onshore. 

Because  of  the  mixture  of  taxa  from  both  coastal  and  inland  sites,  a  single  estimate 
cannot  represent  the  thermal  conditions  for  the  entire  fossil  flora.  As  for  the  coastal 
strip,  Guadalupe  pine  occurs  at  altitudes  generally  above  800  m.  To  judge  from  its 
occurrence  in  a  summer  fog  belt,  mean  annual  temperature  is  approximately  16°C  as 
inferred  from  sea  surface  temperature  (U.S.D.A.,  Climatic  data);  the  persistent  fog  belt 
implies  a  low  range  of  temperature,  probably  not  more  than  8-1 0°C.  Because  the  marine 
invertebrate  fauna  of  the  "lower"  San  Diego  Formation  indicates  that  sea  surface 
temperature  was  warmer  than  at  present,  temperature  along  the  coastal  plain  was  also 
warmer  than  at  present,  probably  with  a  mean  temperature  near  1 6°C.  This  is  suggested 
also  by  the  abundance  of  avocado  (Persea)  specimens.  Since  avocado  does  not  now 


285 


produce  fruit  well  in  areas  much  north  of  Santa  Barbara  (Fig.  5),  temperatures  near 
there  probably  represent  an  extreme  minimum  for  the  fossil  flora.  Digger  pine  occurs 
today  in  Santa  Ynez  Valley  (Cachuma  Lake),  not  far  from  avocado  groves,  where  mean 
temperature  is  about  16°C  and  the  range  is  10-1 1°C.  Farther  north  in  the  Coast  Ranges, 
as  at  Santa  Margarita  (alt.  300  m),  mean  annual  temperature  is  14.5°C  and  the  annual 
range  is  16°C.  The  palm,  Guadalupe  pine,  and  avocado  suggest  that  fossil  digger  pine 
lived  under  conditions  milder  than  those  now  at  Santa  Margarita,  and  probably  were 
like  those  in  the  Cachuma  Lake  area.  As  for  the  interior,  Jeffrey  pine  now  occurs  in 
Pine  Valley  where  mean  annual  temperature,  as  estimated  from  that  at  Cuyamaca,  16 
km  north  and  350  m  higher,  is  near  12.5°C  and  the  mean  monthly  range  is  about  17°C. 
These  estimates  suggest  that  during  the  Late  Pliocene  the  temperature  along  the  coast 
was  about  0.5-1.0°C  warmer  than  that  presently  at  Chula  Vista. 

In  view  of  the  greater  range  of  mean  monthly  temperature  in  the  interior  where 
fossil  Jeffrey  pine  lived,  mean  annual  temperature  there  probably  was  about  2-2. 5°C 
lower  than  on  the  coast  (Fig.  5).  This  suggests  a  depression  of  regional  climate  of  about 
300-400  m,  and  hence  a  lower  altitude  for  taxa  representing  upland  vegetation  as 
compared  with  their  present  occurrence.  Coupled  with  higher  rainfall,  under  the  pos- 
tulated paleotemperature  fossil  Jeffrey  pine  may  have  reached  down  to  near  600  m  as 
compared  with  1050  m  in  Pine  Valley  today.  Fossil  Jeffrey  pine  would  therefore  have 
lived  closer  to  the  marine  embayment,  in  sites  from  which  its  cones  would  more  likely 
be  transported  seaward,  especially  during  ffoods  resulting  from  hurricanes  that  pre- 
sumably were  of  more  frequent  occurrence.  As  suggested  in  Fig.  5,  equability  on  the 
coast  was  near  M  70  as  compared  with  M  67  at  Chula  Vista  today.  Frost  was  absent 
on  the  coast  where  the  growing  season  probably  had  a  mean  daily  temperature  warmer 
than  15.2°C  on  218  days  of  the  year.  Moderate  light  frosts  might  well  be  expected  over 
the  interior  where  fossil  Jeffrey  pine  lived.  The  estimated  paleotemperatures  are  shown 
by  the  circled  areas  A,  B,  and  C,  respectively,  for  the  immediate  coast,  the  central 
digger  pine  woodland,  and  the  lower  margin  of  Jeffrey  pine  forest  (Fig.  5). 

Regional  Comparisons 
Pliocene  Floras 

The  coastal  oak-avocado-palm-pine  woodland,  the  interior  digger  pine-oak  wood- 
land and  the  upland  Jeffrey  pine  forest  of  the  Pliocene  Chula  Vista  ffora  differ  consid- 
erably from  contemporaneous  vegetation  in  central  and  northern  California  (Fig.  1). 

The  ffora  from  the  Sonoma  Formation  at  Neer's  Hill,  Santa  Rosa  (Axelrod  1944c) 
represents  a  coast  conifer  forest  with  redwood  (Sequoia),  lowland  fir  (Abies  cf  grandis), 
Douglas  fir  (Pseudotsuga  cf  menziesii),  coast  hemlock  (Tsuga  cf  heterophylla)  and 
winged  seeds  of  weeping  spruce  (Picea  cf  brewehand).  Associated  with  the  conifers, 
and  also  forming  a  dense,  broadleaved  sclerophyll  forest  on  warmer  slopes,  were  species 
of  oak  (Quercus  cf  chrysolepis),  chinquapin  (Castanopsis  [Chrysolepis]  chrysophylla), 
California  laurel  (Umbellularid),  and  tanoak  (Lithocarpus).  Riparian  species  included 
sycamore,  cottonwood,  avocado,  and  willow.  A  few  Tertiary  relicts  are  recorded,  no- 
tably species  of  Ilex,  Persea,  Trapa  and  Ulmus.  It  is  estimated  that  the  area  received 
fully  89-100  cm  of  rainfall,  some  in  summer,  and  that  temperatures  were  cool  though 
frost  was  absent. 

Representatives  of  the  Sonoma  ffora  apparently  extended  farther  north,  as  suggested 
by  a  small  flora  from  the  upper  part  of  the  Wildcat  Group  near  Garberville  (Axelrod 
1944c:  187,  Dorf  1930).  This  fossil  flora  is  composed  chiefly  of  riparian  taxa  and  is 
characterized  by  an  absence  of  typical  forest  species.  The  flora  apparently  accumulated 
on  a  broad  floodplain  some  distance  from  forests  that  occupied  slopes  bordering  the 
lowland  marine  basin.  The  only  forest  representatives  are  logs  of  Sequoia  and  Pseu- 
dotsuga and  a  leaf  of  Ulmus.  Members  of  floodplain  vegetation  included  alders  (Alnus 
cf.  rhombifolia,  rubra),  avocado  (Persea  cf  borbonia),  sycamore  (Platanus  cf  racemosa), 
black  cottonwood  (Populus  cf  hastata),  willow  (Salix  cf  lasiolepis)  and  California 
laurel  ( Umbellularia).  Some  of  these  appear  to  represent  Miocene  relicts,  notably  the 


286 


Table  2.     Comparison  of  Chula  Vista  flora  with  related  living  taxa. 


Fossil  species 


Allied  living  species 


Pinus  diegensis 
Pinus  jeffreyoides 
Pinus  pieperi 
Platanus  paucidentata 
Populus  alexanderi 
Persea  coalingensis 

Quercus  lakevillemis 
Arecaceae  sp. 

Shrub 

Salix  wildcatensis 
Herbaceous  perennial 

Juncaceae  sp. 


P.  radiata  var.  binata  Englemann 

P.  Jeffrey i  Murray  ^ 

P.  sabiniana  Douglas 

P.  racemosa  Nuttall 

P.  trichocarpa  Torrey  &  Gray 

P.  podadenia  Blake;  P.  borbonia 

(Linne)  Sprengel 
Q.  agrifolia  Nee 
Braheal,  Sabal? 


S.  lasiolepis  Bentham 


Juncusl 


Persea,  Populus,  Ulmus  and  possibly  the  Platanus  which  is  represented  by  an  incomplete 
specimen.  Their  persistence  here  in  the  late  Tertiary  is  understandable  in  view  of  their 
coastal  position  where  there  was  high  rainfall  and  low  evaporation.  It  is  evident  that 
the  Garberville  flora  shows  little  relation  to  the  Chula  Vista  flora  which  is  expectable 
in  view  of  its  position  fully  1025  km  southeast. 

The  Napa  flora  (Axelrod  1950a),  situated  45  km  southeast  of  the  Sonoma  flora  at 
Santa  Rosa,  represents  a  pine  (Pinus  cf.  ponderosa)-Doug\as  fir  (Pseudotsuga)  forest 
living  near  sea  level.  Forest  associates  included  chinquapin  (Castanopsis  [Chrysolepis] 
cf.  chrysophylla,  C.  sempervirens),  Oregon  grape  (Mahonia  cf.  nervosa),  ocean  spray 
(Holodiscus  cf.  discolor),  goldcup  oak  (Quercus  chrysolepis),  interior  liveoak  (Q.  cf 
wislizenii),  and  California  laurel  (Umbellularia  cf.  californica).  Sequoia  is  rare,  with 
only  three  small  twigs  represented  in  the  sample  of  over  700  specimens.  Warmer  slopes 
were  covered  with  broadleaved  sclerophyll  vegetation  composed  of  Castanopsis,  Quer- 
cus, Umbellularia,  and  probably  Persea.  An  open  oak  woodland-grass  of  coast  liveoak 
(Quercus  cf.  agrifolia),  valley  oak  (Q.  lobata),  and  interior  liveoak  (Q.  wislizenii)  oc- 
cupied drier  slopes.  The  warmer,  driest  sites  supported  a  chaparral  of  whitehom 
(Ceanothus  cf.  leucodermis),  mountain  mahogany  (Cerocarpus  cf  betuloides,  C.  cf. 
ledifolius),  and  toyon  (Heteromeles  cf.  arbutifolia),  though  these  shrubs  also  occurred 
in  the  oak  woodland.  Stream-  and  lake-border  sites  supported  avocado  (Persea),  syc- 
amore (Platanus),  willows  (Salix  cf.  laevigata,  S.  scouleriana)  and  California  laurel 
( Umbellularia).  The  assemblage  reflects  a  drier,  more  continental  climate  than  that  in 
the  Santa  Rosa  area  to  the  northwest.  The  only  relationship  with  the  Chula  Vista  flora 
is  seen  in  a  few  riparian  taxa  that  ranged  widely  and  through  several  vegetation  zones. 

The  Turlock  Lake  flora,  slightly  older  than  the  Chula  Vista,  accumulated  on  the 
lowest  floodplain  in  front  of  the  central  Sierra  Nevada.  It  is  preserved  in  the  upper  part 
of  the  Mehrten  Formation  35  km  east  of  Modesto  (Axelrod  1980^?).  The  lake  border 
assemblage  included  avocado  (Persea),  sycamore  (Platanus),  paloblanco  (Forestiera), 
willow  (Salix),  and  California  laurel  ( Umbellularia).  The  bordering  slopes  were  covered 
with  oak  woodland-grass  (Quercus  cf.  douglasii,  Q.  wislizenii)  and  scattered  shrubs, 
including  scrub  oaks  (Quercus  cf.  dumosa,  Q.  dunnii),  ceanothus  (Ceanothus  cf  sore- 
diatus),  cofleeberries  (Rhamnus  californica,  R.  ilicifolia),  and  poison  oak  (Toxicoden- 
dron) that  may  have  formed  local  brushy  patches  on  exposed  drier  slopes.  Cooler, 
moister  sites  supported  broadleaved  sclerophyll  taxa,  notably  madrone  (Arbutus),  in- 
terior live  oak  (Q.  wislizenii),  and  California  laurel  (Umbellularia).  Reaching  down 
from  higher  levels  along  stream  valleys  were  members  of  a  moister  flora,  including 
pine  (Pinus  cf.  ponderosa)  and  smilax  (Smilax  cf.  californica).  Two  exotics  are  in  the 
flora,  an  aspen  (Populus  cf.  tremula)  and  a  cherry  (Prunus),  both  of  Asian  affinity. 
There  was  some  summer  rainfall  and  annual  precipitation  totalled  approximately  63 


287 


cm  as  compared  with  36  cm  today.  The  flora  shows  Httle  relationship  with  that  at 
Chula  Vista  apart  from  several  widely  distributed  riparian  species. 

The  small  Coalinga  flora  from  the  upper  San  Joaquin  Formation  10  km  south  of 
Coalinga  indicates  that  the  same  general  type  of  floodplain  vegetation  preserved  at 
Turlock  Lake  ranged  across  the  Central  Valley.  The  fossils  occur  in  sandstones  of 
fluviatile  origin  interbedded  with  marine  strata  containing  molluscs  that  represent  the 
Pecten  coalingensis  zone  (Dorf  1 930).  The  flora  has  abundant  leaves  of  avocado  (Persea) 
and  sycamore  {Plat  anus),  and  together  with  cotton  wood  (Populus)  and  hackberry  (Cel- 
tis)  are  indicative  of  riverbanks  and  moist  sites.  Adjacent  interfluves  were  covered  with 
oak  (Quercus  cf.  douglasii)  as  well  as  silk-tassel  bush  (Garrya  cf.  elliptica).  Further 
collecting  at  this  site  should  provide  a  better  representation  of  the  flora  and  vegetation 
of  this  area. 

An  indication  of  the  flora  that  occupied  the  region  slightly  earlier  (ca.  5  m.y.)  is 
provided  by  a  flora  from  Broken  Hill,  30  km  southeast,  at  the  south  end  of  North 
Dome,  Kettleman  Hills,  in  the  basal  part  of  the  San  Joaquin  Formation  (Axelrod  1 980^). 
It  also  provides  evidence  of  vegetation  in  the  area  prior  to  uplift  of  the  Coast  Ranges 
directly  west,  an  event  that  brought  a  drier,  semidesert  climate  to  this  area.  The  flora 
occurs  in  sandstones  of  fluviatile  origin  that  grade  upward  into  marine  beds.  Floodplain 
vegetation  included  alder  (Alnus),  sycamore  (Platanus),  cottonwood  (Populus),  poplar 
(Populus  euphmtica),  aspen  (Populus  cf.  tremula),  California  laurel  (Umbel- 
lularia)  as  well  as  several  willows  (Salix  cf.  exigua,  S.  lasiandra,  S.  laevigata,  S.  lasi- 
olepis),  and  soapberry  (Sapindus).  Avocado  (Persea)  is  abundant,  magnolia  is  present, 
and  both  contributed  to  the  floodplain  vegetation.  Well-drained  interfluves  were  cov- 
ered with  oak  woodland-grass  that  included  three  oaks  (Quercus  cf.  agrifolia,  Q.  doug- 
lasii, Q.  wislizenii).  There  is  also  evidence  of  broadleaved  sclerophyll  vegetation,  prob- 
ably on  higher  slopes  to  the  west.  These  included  tanoak  (Lithocarpus),  morheus  oak 
(Q.  morheus),  and  California  laurel  (Umbellularia),  as  well  as  avocado  and  magnolia. 
The  flora  has  several  exotics  in  addition  to  Magnolia  and  Persea,  notably  Populus  cf. 
tremula,  P.  cf.  euphratica,  Sapindus  and  Ulmus,  all  indicating  summer  rainfall.  Apart 
from  these,  the  flora  resembles  vegetation  of  the  lower  middle  slopes  of  the  Santa  Lucia 
Mountains  120  km  west.  Rainfall  is  estimated  to  have  been  near  75  cm  annually,  and 
climate  was  more  equable  than  that  now  in  this  semi-desert  region  where  precipitation 
totals  12  cm  yearly. 

Evidently  the  Pliocene  floras  of  central  and  northern  California  represent  vegetation 
quite  different  than  that  at  Chula  Vista.  Much  as  the  vegetation  of  these  areas  differs 
today,  the  differences  reflect  the  uniqueness  of  the  floristic  provinces  represented.  The 
principal  links  with  the  floras  to  the  north  were  riparian  taxa  that  ranged  widely  into 
very  different  vegetation  zones  and  which  have  a  broad  time  span.  The  transition  from 
the  northern  coast  conifer  forests  to  the  coastal  vegetation  of  the  Chula  Vista  area 
probably  corresponded  generally  with  the  axis  of  the  present  Transverse  Ranges. 

Plio- Pleistocene 

The  preceding  late  Neogene  floras  were  succeeded  by  those  that  represent  colder, 
moister  climates.  This  is  illustrated  by  the  Soboba  flora  from  interior  southern  Cali- 
fornia (Axelrod  1967a),  dated  by  the  Bautista  mammalian  fauna  at  1  m.y.  This  flora 
shows  that  mixed  conifer  forest  reached  down  fully  1000  m  below  its  present  level  in 
the  San  Jacinto  Mountains  which  now  tower  above  the  fossil  locality.  Included  as 
macrofossils  that  represent  montane  conifer  forest  are  Abies  concolor,  Calocedrus  de- 
currens,  Pinus ponderosa,  P.  lambertiana,  and  Populus  tremuloides.  Associates  included 
species  of  upper  woodland  and  chaparral  vegetation.  Precipitation  was  near  70  cm 
annually  as  compared  with  33  cm  in  the  area  today  and  mean  annual  temperature  was 
fully  5.5°C  lower  than  at  present  (Axelrod  1976:  fig.  5).  A  flora  of  comparable  age  is 
represented  in  the  Santa  Cruz  Mountains  near  Saratoga  (Dorf  1930).  It  records  forest 
taxa  (Libocedrus,  Pinus  cf.  lambertiana,  Pseudotsuga)  at  much  lower  altitudes  than 
occur  near  sea  level  in  this  area  today.  By  contrast,  the  fossil  site  supports  oak  woodland- 


288 


grass  and  broadleaved  sclerophyll  forest.  Redwood  forest  is  now  at  higher  levels,  and 
is  not  represented  in  the  fossil  flora. 

The  magnitude  of  the  climatic  shift  along  the  coastal  strip  in  the  Early  Pleistocene 
is  indicated  by  a  flora  from  the  marine  upper  Pico  Formation  (Early  Pleistocene,  1 
m.  y.),  situated  near  Seacliff'on  the  coast  west  of  Ventura  (Axelrod  1983).  Several  cones 
of  Douglas  fir  {Pseudotsuga  menziesii)  are  represented,  a  species  that  now  has  scattered 
populations  in  the  Santa  Lucia  Mountains  325  km  north  but  has  its  principal  southern 
area  in  the  Santa  Cruz  Mountains  450  km  north.  Associates  in  the  coastal  strip  were 
three  pines  that  are  of  coastal  occurrence  today,  Monterey  pine  (Pinus  radiata).  Island 
pine  {P.  remorata),  and  Stanton  pine  {Pinus  muricata  var.  stantonii).  The  Seacliff' 
assemblage  suggests  that  temperature  along  the  coastal  strip  was  approximately  13.5°C 
as  compared  with  15°C  today  and  that  rainfall  totalled  76  cm  in  contrast  to  38  cm  at 
present. 

Development  of  Modern  Forest  Geography 

The  conifer  forests  in  the  mountains  of  southern  California  and  Baja  California 
Norte  are  now  perched  as  discontinuous  stands  on  isolated  mountain  ranges  at  altitudes 
generally  above  1 500  m.  At  lower  altitudes,  semidesert,  chaparral  and  woodland  vege- 
tation thrive  under  warmer,  drier  climates.  The  discontinuous  montane  forests  have 
similar  dominants,  notably  Abies  concolor,  Calocedrus  decurrens,  Pinus  jeffreyi,  P. 
ponderosa,  P.  lambertiana,  P.  murrayana,  Populus  hastata  {""trichocarpd")  and  P.  tre- 
muloides  (see  Griffin  and  Critchfield  1972,  Munz  1935,  Nelson  1922,  Wiggins  1980). 
In  addition,  many  forbs,  herbaceous  perennials,  and  shrubs  link  these  montane  forests. 

The  now-isolated  forest  taxa  once  had  a  more  continuous  distribution  in  the  Late 
Pliocene-Early  Pleistocene  when  climate  was  considerably  wetter  and  cooler  than  at 
present,  as  demonstrated  by  the  Soboba  flora  from  San  Jacinto  Valley  in  interior 
southern  California  (Axelrod  1967<3)  and  by  the  limited  Chula  Vista  flora. 

In  the  Plio-Pleistocene  transition,  mountains  were  still  relatively  low,  probably 
scarcely  half  their  present  altitudes  which  now  reach  up  to  3000-3400  m.  As  colder, 
wetter  climates  spread  southward,  conifer  forests  shifted  from  the  Sierra  Nevada  south- 
ward into  the  Tehachapi-Mount  Pinos-Liebre  Mountain  region  and  thence  through 
the  San  Gabriel-San  Bernardino-San  Jacinto  Mountains  and  into  the  Sierra  San  Pedro 
Martir  in  Baja  California  Norte  (see  photos  in  Nelson  1922).  As  the  fault-block  moun- 
tains were  elevated  later  to  their  present  heights,  the  forests  were  stranded  as  disjunct 
stands  isolated  by  drier,  warmer  climates  occupied  by  woodland,  chaparral,  sage,  and 
semi-desert  vegetation.  As  the  forests  were  restricted  to  higher  altitudes,  some  taxa 
disappeared  locally,  thus  accounting  for  the  present  disjunct  occurrences  in  the  moun- 
tains of  southern  California  (see  maps  in  Griffin  and  Critchfield  1972,  Little  1971,  also 
discussion  in  Munz  1935).  Other  disjunct  distributions  of  Sierran  montane  taxa  in  the 
mountains  of  southern  California  are  exemplified  by  Acer  glabrum,  Cornus  nuttallii, 
Euonymus  occidentalis,  Phyllodoce  breweri,  Rhododendron  occidentale,  Prunus  emar- 
ginata,  and  many  forbs,  herbaceous  perennials  and  shrubs  listed  by  Munz  (1935). 
Greater  disjunctions  from  the  southern  Sierra  Nevada  are  seen  in  the  distribution  of 
Pinus  murrayana  (San  Gabriel,  San  Bernardino,  San  Jacinto  mountains,  and  San  Pedro 
Martir)  and  Populus  tremuloides  (San  Bernardino  Mountains  and  San  Pedro  Martir). 

Evolution  and  Biogeography  of  Pinus  radiata 

The  Chula  Vista  flora  provides  additional  evidence  regarding  the  probable  evo- 
lutionary history  of  the  five  living  populations  of  Pinus  radiata.  Based  on  their  cones 
these  populations  can  be  separated  into  two  groups.  The  populations  of  P.  radiata  on 
Cedros  Island  (var.  cedrosensis)  and  on  Guadalupe  Island  (var.  binata)  represent  one 
group  characterized  by  small  symmetrical  to  slightly  asymmetrical  cones  without  prom- 
inent apophyses  except  for  the  extreme  variation  seen  in  var.  binata.  By  contrast,  the 
other  group  represented  by  the  California  populations  at  Monterey,  Aiio  Nuevo  and 
Cambria  possess  larger,  regularly  asymmetrical  cones  with  large,  rounded  apophyses. 


289 


Table  3.     Trees  and  shrubs  in  coastal  southern  CaHfomia  with  close  relatives  in  equable  montane  areas  of 
Mexico. 

California  Mexico 

Arbutus  menziesii  A.  xalapensis 

Ceanothus  arboreus  C.  caeruleus 

Cercocarpus  traskiae  C.  mojadensis 

Comarostaphylis  diversifoHa  C.  spp.  ( 1 2  or  more) 

Myrica  californica  M.  mexicana 

Pinus  remorata  P.  oocarpa 

Pinus  torreyana  P.  oxacana 

Prunus  lyonii  P.  prionophylla 

Vaccinium  ovatum  V.  confertum 


There  is  considerable  similarity  between  Pinus  radiata  var.  cedrosensis  and  P. 
remorata  Mason  which  has  large  populations  in  Pine  Canyon  west  of  Lompoc,  on  Santa 
Cruz  and  Santa  Rosa  islands,  and  in  Baja  California  near  the  coast  southwest  of  San 
Vicente.  Cones  of  Cedros  Island  pine  differ  from  those  off.  remorata  in  having  some- 
what more  prominent  apophyses  and  slightly  thicker  cone  scales.  These  differences 
seem  to  reflect  the  different  environments  to  which  these  pines  are  adapted,  with  the 
cooler,  foggier,  moister  climate  in  the  north  favoring  the  persistence  of  a  more  "prim- 
itive type,"  as  exemplified  by  P.  remorata.  Cones  of  P.  remorata  are  similar  to  those 
of  P.  oocarpa,  though  in  its  northern  areas  P.  oocarpa  cones  approach  those  of  P. 
radiata  var.  cedrosensis  in  having  more  prominent  apophyses  (see  U.C.  Herbarium) 
than  in  areas  to  the  south. 

The  geologic  ages  of  these  pines  are  not  presently  known,  but  to  judge  from  their 
relation  to  Pinus  oocarpa  and  their  present  disjunct  distribution  (see  Martinez  1948), 
they  have  probably  been  in  existence  since  the  Middle  Miocene.  Up  to  500  km  of  right- 
lateral  strike-slip  movement  since  the  Middle  Miocene  along  the  San  Andreas  fault 
system  and  related  rifting  in  the  Gulf  of  California  is  probably  in  large  part  responsible 
for  the  present  biogeography  (Gastil  and  Jensky  1973).  The  fact  that  Baja  California 
was  once  part  of  the  Mexican  mainland  is  supported  by  floristic  evidence  that  provides 
additional  data  with  respect  to  derivation  of  Monterey  pines  from  ancient  members  of 
Pinus  subsect.  Oocarpeae.  Geologic  evidence  indicates  that  a  chain  of  low  coastal 
mountains  was  once  situated  west  of  the  San  Andreas  rift.  These  hills  were  probably 
covered  with  taxa  that  now  have  closely  allied  species  in  southern  California  and  in 
the  mountains  of  Mexico,  chiefly  in  areas  of  equable  climate  (Table  3). 

Today,  these  allied  taxa  are  now  isolated  by  the  broad  stretch  of  the  Sonoran  Desert 
and  adjacent  Thorn  Forest  vegetation.  Links  are  also  provided  by  a  number  of  taxa 
now  in  Mexico  that  have  allied  fossil  species  in  the  Miocene  Puente  and  Modelo  floras 
of  coastal  southern  California  (Axelrod  1977:162),  distributed  in  Clethra,  Magnolia, 
Nectandra,  Persea,  Quercus,  Sabal  and  others.  They  clearly  indicate  that  a  humid, 
equable  Miocene  route  connected  these  areas.  That  there  was  a  near-coastal  route  is 
indicated  also  by  the  Mint  Canyon  (Axelrod  1979:25,  32)  and  Tehachapi  floras  (Axelrod 
1939).  They  demonstrate  that  by  19  m.y.  ago— and  certainly  earlier— interior  southern 
California-Arizona  was  too  semiarid  and  hot  for  the  taxa  noted  above.  Hence,  allies 
of  the  mesic  species  now  in  coastal  California  or  in  Mexico  shifted  into  coastal  southern 
California  via  a  near-coastal,  not  an  interior  route.  It  was  this  route  that  brought  pines 
of  subsect.  Oocarpeae  to  California. 

At  that  time,  it  is  inferred  that  the  ancestor  of  Pinus  radiata  var.  cedrosensis  was 
in  the  mountains,  living  under  a  mild,  warm  temperate  climate.  Insularity  developed 
from  opening  of  the  Gulf  of  California  (since  5  m.y.),  and  submergence  of  the  outer 
continental  shelf  There  the  pine  has  persisted  on  the  higher  summits  of  Cedros  Island 
where  it  is  sheltered  by  a  regular  summer  fog  belt  at  altitudes  above  600  m.  The  smaller 
cones  of  the  Cedros  and  Guadalupe  Island  pines  seemingly  reflect  their  closer  relation 
to  P.  oocarpa  which  was  not  far  away  in  the  mountains  of  Sonora-Sinaloa,  to  which 


290 


Baja  California  was  joined  in  the  Miocene.  Further,  their  small  size  may  reflect  the 
summer  rain  regime  to  which  they  are  and  were  adapted. 

Pinus  diegensis,  closely  allied  to  P.  radiata  var.  binata,  may  have  been  derived 
from  var.  cedrosensis  by  adaptation  to  somewhat  greater  aridity  over  the  northern  part 
of  the  earlier  distribution  of  the  immediate  ancestor  of  var.  cedrosensis.  This  may 
account  for  the  greater  variation  in  symmetry  and  apophyses  development  in  P.  radiata 
var.  binata.  That  P.  diegensis  (cf.  binata)  may  represent  the  ancestral  form  that  gave 
rise  to  the  three  California  populations  of  Pinus  radiata  is  suggested  by  its  Pliocene 
mainland  occurrence,  and  by  its  cone  variation  which  is  intermediate  between  var. 
cedrosensis  and  the  type  Monterey  population. 

Cones  of  the  three  California  populations  o^  Pinus  radiata,  which  are  larger,  more 
asymmetrical,  and  have  prominent  apophyses,  presumably  reflect  adaptation  to  a  pro- 
gressively more  extreme  mediterranean  climate  of  dry  summers.  In  the  suggested  evo- 
lutionary sequence— oocarpa-remorata-cedrosensis-binata-Monterey-A  Ho  Nuevo- 
Cambria—iX  is  noteworthy  that  Pinus  diegensis  approaches  the  variation  of  the  Mon- 
terey population.  This  lends  further,  though  still  tenuous,  support  to  its  probable  place 
in  the  evolution  of  Monterey  pines. 

The  fossil  samples  now  available  suggest  that  Pinus  radiata  populations  with  cones 
comparable  in  size,  symmetry,  and  apophyses  development  to  the  Guadalupe  and 
Monterey  populations  were  rather  widespread  in  coastal  California  well  into  the  Pleis- 
tocene. At  Carpinteria  cones  more  nearly  approaching  the  larger-sized  Aiio  Nuevo 
population  appear  only  late  in  the  Pleistocene  (Axelrod  1980^).  Cones  the  size  of  the 
present  Cambria  population  are  not  now  known  as  fossils.  They  may  be  the  most  recent 
of  the  group,  possibly  originating  in  post-glacial  time. 


Conclusion 

In  the  Late  Pliocene  (ca.  3  m.y.),  increased  precipitation  and  lower  temperatures 
in  California  enabled  floras  of  more  mesic,  cooler  requirements  to  replace  floras 
adapted  to  drier,  warmer  climate,  as  illustrated  by  the  Mulholland  (Axelrod  \944a), 
Oakdale  (Axelrod  1 944b),  Piru  Gorge  (Axelrod  1 9506^)  and  Mount  Eden  (Axelrod  1937, 
\950b)  floras.  The  more  mesic,  younger  floras,  like  the  Santa  Rosa,  Napa,  Upper 
Wildcat,  Turlock  and  (probably)  Coalinga,  still  have  a  few  exotic  taxa  allied  to  species 
in  summer  rain  areas,  reflecting  a  warmer  sea  surface  than  that  of  today.  Owing  to  its 
more  southerly  position,  the  influence  of  cooler  climate  at  Chula  Vista  was  not  so 
pronounced,  though  precipitation  was  about  double  that  of  today.  The  lowland  flora 
of  southern  California  then  represented  a  different  fforistic  province  than  that  to  the 
north,  one  adapted  to  a  warmer  climate,  much  as  comparable  thermal  differences 
separate  the  floras  of  northern  and  southern  California  today. 

The  succeeding  colder,  wetter  climate  of  the  latest  Pliocene  and  early  Pleistocene 
enabled  Douglas  fir  {Pseudotsuga  menziesii)  forest  to  shift  south  into  coastal  southern 
California,  fully  350  km  (or  somewhat  more)  south  of  its  present  scattered  coastal 
stations.  Over  the  interior,  lowered  temperature  and  higher  precipitation  enabled  the 
Sierran  mixed  conifer  forest  to  range  southward  into  the  mountains  of  southern  Cal- 
ifornia and  Baja  California  Norte.  The  Sierran  mixed  conifer  forest  then  lived  fully 
1000  m  below  its  present  level  at  a  time  when  mountains  were  appreciably  lower.  The 
present  forests  were  stranded  as  isolated  patches  as  the  discontinuous  mountains  were 
uplifted  in  the  middle  and  later  Quaternary  and  drier,  hotter  climates  spread  over  the 
region.  These  later,  more  extreme  conditions  probably  account  for  the  present  disjunct 
occurrences  of  diverse  montane  taxa  in  the  mountains  of  southern  and  Baja  California, 
and  for  the  restriction  of  taxa  allied  to  those  in  the  Chula  Vista  flora  to  more  local 
areas. 

Evidence  suggests  that  Pinus  diegensis  (cf.  P.  radiata  var.  binata)  may  have  given 
rise  to  the  California  populations  of  Pinus  radiata  by  progressive  adaptation  to  in- 
creasing summer  drought.  Further,  P.  radiata  var.  binata  did  not  originate  in  insular 


291 


isolation,  but  like  many  other  insular  endemic  trees  and  shrubs,  was  confined  there  by 
unfavorable  land  climates  during  the  past. 

Systematics 

Family  Pinaceae 

Pinus  diegensis  new  species 

Figures  6A,  B,  7A,  B 

Description.— Cones  long-oval  to  elliptic  oval;  small  cone  (SDSNH  25135)  9  cm 
long  and  5  cm  broad,  medium  sized  cone  (SDSNH  25 165)  about  10-1 1  cm  long(estim.), 
and  about  6.5  cm  broad;  largest  cones  (SDSNH  25136,  25137)  somewhat  flattened, 
fragmentary.  Cone  scales  up  to  4.0-4.5  cm  long  and  2.0-2.3  cm  broad,  distally.  Tips 
of  cone  scales  broadly  flattened,  with  slightly  swollen,  convexly  rounded  apophyses 
well  inside  tip  of  scale. 

Types. -Holotype  SDSNH  no.  25135,  Paratypes  SDSNH  no.  251 10,  25136,  25137, 
25165. 

Discussion.  — Four  incomplete  cones  are  sufliciently  distinct  from  those  previously 
described  to  warrant  the  recognition  of  a  new  species.  Examination  of  the  large,  excellent 
collection  of  pine  cones  in  the  herbarium  of  the  U.S.  Forest  Service,  Institute  of  Ge- 
netics, Placerville,  California,  indicates  that  P.  diegensis  is  a  member  of  the  subsect. 
Oocarpeae  (Little  and  Critchfield  1 969).  The  fossil  cones  are  most  similar  to  the  variable 
cones  produced  by  Pinus  radiata  var.  binata  Engelmann  from  Guadalupe  Island.  The 
variation  present  in  the  small  fossil  collection  is  readily  duplicated  by  the  large  suites 
of  cones  that  have  been  recovered  from  the  island. 

Howell  (1941a)  discussed  cone  variation  in  the  Guadalupe  pine,  noting  that  asym- 
metrical cones  with  large  apophyses  on  the  outer  side  (like  fig.  6B)  were  similar  to  the 
type  which  comes  from  Monterey.  A  second  form  with  nearly  symmetrical  cones  (like 
fig.  6A)  and  with  scales  alike  on  all  sides  and  with  little  prominent  apophyses  devel- 
opment Howell  termed  forma  guadalupensis.  A  third  form,  intermediate  between  the 
latter  two  he  termed /orma  binata,  noting  that  it  corresponded  with  the  type  specimen 
collected  from  Guadalupe.  As  discussed  separately  in  the  section  on  Evolution,  the 
variable  binata  on  Guadalupe  Island  may  represent  a  population  close  to  that  which 
gave  rise  to  the  three  living  populations  in  central  California. 

In  southern  California,  fossil  pine  cones  allied  to  the  present  California  populations 
of  Pinus  radiata  have  been  recorded  previously  from  the  late  Tertiary  Mount  Eden 
(Axelrod  1937)  and  Lower  Pico  floras  (Dorf  1930).  It  is  also  known  from  Pleistocene 
floras  at  Carpinteria  (Chaney  and  Mason  1933),  near  Seacliff  on  the  coast  west  of 
Ventura  (Axelrod  1983),  and  at  Rancho  La  Brea  (Mason  1927).  In  coast-central  Cal- 
ifornia, it  is  known  from  Mussel  Rock  on  the  outer  coast  south  of  San  Francisco.  Here 
it  has  been  recovered  from  an  old  forest  soil  that  rests  on  Franciscan  diabase  and  is 
overlain  by  the  basal  beds  of  the  marine  Merced  Formation,  approximately  5-6  m.y. 
old  (Axelrod  1967Z>).  Pleistocene  records  in  central  California  are  near  Pt.  Sal  (Axelrod 
1961b),  Little  Sur  (Langenheim  and  Durham  1963),  Millerton  (Mason  1934,  Axelrod 
1980a),  and  Drakes  Bay  (Axelrod  1980a,  1983). 

The  present  record  is  the  most  southern  fossil  locality  now  known  for  species  of 
Monterey  pine. 

Pinus  jeffreyoides  new  species 
Figures  7C,  D 

Description.— Cone  large,  estimated  15-10  cm  long  and  fully  10  cm  broad;  cone 
scales  large,  up  to  4  cm  long  in  central  part  of  cone;  terminal  part  of  cone  scale  broadly 
triangular,  1.5-2.5  cm  wide  and  0.8-1.1  cm  thick  as  measured  below  the  umbo. 

Typ^'^.-Holotype  SDSNH  25166;  Paratypes  SDSNH  25138.  25167. 

Discussion.— This  is  a  member  of  subsect.  Ponderosae  Little  and  Critchfield  ( 1 969) 
and  is  well-matched  by  cones  of  the  living  Pinus  jeffreyi  Greville  and  Balfour.  Rela- 


292 


r"^-^. 


J  jtff^ 


Figure  6.     \,^—Pinus  diegensis  new  species,  A  — holotype,  SDSNH  25135,  B— paratype,  SDSNH  25165. 
All  specimens  x  ]  (latex  casts). 


tionship  with  P.  jeffreyi  is  apparent  in  the  morphology  of  the  distal  ends  of  the  cone 
scales,  which  are  much  thicker  than  those  of  the  allied  P.  ponderosa  Lawson. 

Pinus  pieperi  Dorf 
Figures  7B 

Pinus  pieperi  Dorf,  Carnegie  Inst.  Wash.  Publ.  412,  p.  69,  pi.  5,  figs.  7-10,  1930; 
Axelrod,  Carnegie  Inst.  Wash.  Publ.  476,  p.  156,  pi.  2,  figs.  2,  3,  1937. 

Referred  specimen. -S,X)SN\\  25168. 

Remarks.  — A.  single  fragment  of  a  cone  with  distinctive  cone  scales  that  have 


293 


Figure  7.  A,B—Pinus  diegensis  new  species.  A— paratype,  SDSNH  251 10,  B— paratype,  SDSNH  25137; 
CD- Pinns  jeffreyoides  new  species,  C-holotype,  SDSNH  25166,  D-paratype,  SDSNH  25138;  E-Pinus 
pieperi  Dorf,  SDSNH  25168.  All  specimens  x  1  (latex  casts). 


prominently  hooked,  large  apophyses  represents  this  species  which  is  allied  to  digger 
pine,  Pinus  sabiniana  Douglas  of  central  California.  The  specimen  measures  6.0  cm 
long  and  5.5  cm  wide. 

Pinus  pieperi  has  been  recorded  previously  in  southern  California  from  the  Plio- 
Pleistocene  rocks  north  of  Ventura  (Wiggins  1951).  in  the  lower  part  of  the  Pico  For- 
mation west  of  Ventura  (Dorf  1930).  in  the  upper  Pico  Formation  near  Seacliff  on  the 
coast  west  of  Ventura  (Axelrod  1 983).  as  well  as  in  the  Mount  Eden  flora  near  Beaumont 
(Axelrod  1937).  A  fragmentary  cone  scale  from  the  Anaverde  Formation  near  Palmdale 


294 


has  also  been  referred  to  Pinus  pieperi  (Axelrod  1950c).  Digger  pine  may  have  been 
eliminated  from  southern  California  as  drier  hotter  climate  spread  there  during  the 
Xerothermic. 

Family  Arecaceae 

Gen.  et  sp.  indet. 

Figure  8A 

Referred  specimen.— SUSNH  25 163. 

Remarks.— A  fragmentary  specimen  that  certainly  represents  a  palm  is  in  the 
collection.  It  is  7  cm  long  and  2.5  cm  broad,  the  blade  has  4  rays  each  with  a  prominent 
midvein  and  each  is  bordered  by  7-8  fine  parallel  veins  that  are  somewhat  less  than 
1  mm  apart. 

Comparison  with  several  genera  indicates  that  it  may  represent  a  species  of  Sabal 
or  Brahea  {=Erythea),  but  in  view  of  the  incomplete  nature  of  the  fossil  reference  to 
either  (or  any)  genus  seems  unjustified.  Both  of  the  noted  genera  extend  up  into  oak 
woodland  vegetation,  Sabal  in  northern  Sinaloa  and  adjacent  Sonora,  Brahea  in  the 
highlands  of  southernmost  Baja  California,  and  both  regions  with  ample  rainfall  in 
summer.  In  addition,  Brahea  is  associated  with  Monterey  pine  on  Guadalupe  Island. 

Family  Juncaceae 

Gen.  et.  sp.  indet. 

Figure  83 

Referred  specimen. -SDSNH  25164. 

Remarks.— A  single  slender  reed-like  leaf  impression  is  5  cm  long  and  7  mm  wide 
with  9-10  parallel  veins.  The  blade  is  without  a  midrib  and  clearly  represents  a  sedge- 
or  reed-like  plant  of  indeterminate  nature.  It  seems  comparable  to  some  leaves  of 
Cyperus,  and  especially  to  Juncus,  both  common  in  marshy  areas  and  on  banks  along 
slow,  meandering  rivers. 

Family  Salicaceae 

Populus  alexanderi  Dorf 

Figures  8E,  F 

Populus  alexanderi  Dorf,  Carnegie  Inst.  Wash.  Publ.  412,  p.  75,  pi.  6,  fig.  11  only, 
1930;  Axelrod,  Carnegie  Inst.  Wash.  Publ.  553,  p.  281,  pi.  48,  fig.  4,  1944;  Axelrod, 
Univ.  Cahf  Publ.  Geol.  Sci.  34,  p.  128,  pi.  19,  figs.  1-11,  1958. 

Referred  specimens. -^USHH  25159,  25160,  25161,  25162. 

Remarks.— This  fossil  species  represents  a  cottonwood  similar  to  the  coastal  Pop- 
ulus trichocarpa  Torrey  and  Gray,  the  type  of  which  comes  from  the  Santa  Clara  River 
near  Ventura.  This  species  differs  from  leaves  commonly  identified  as  P.  alexanderi 
and  P.  eotremuloides  Knowlton  in  its  ovate  shape  and  smaller  size  as  compared  with 
the  lanceolate-ovate  form  of  the  species  from  Miocene  floras  to  the  north  and  in  the 
north  coast  Pliocene  floras  as  well.  The  specimens  from  the  Verdi  flora  (Axelrod  1958, 
pi.  19)  illustrate  the  nature  of  the  species,  and  the  accompanying  plate  20  (loc.  cit.) 
shows  how  much  Populus  ""trichocarpa''^  from  the  mountains  differs  from  it;  this  mon- 
tane and  north  coastal  species  seems  to  represent  P.  hastata  Dode. 

P.  alexanderi  (restricted)  is  indicative  of  a  climate  with  mild  winters  as  compared 
with  P.  eotremuloides  which  inhabited  areas  with  colder  climate.  The  ovate-leaved  P. 
alexanderi  is  related  to  P.  emersoni  Condit  from  the  San  Pablo  flora  (Condit  1938; 
Lesquereux  1883,  pi.  55,  figs.  3,  5,  only),  though  the  latter  has  consistently  larger  leaves. 

Salix  wildcatensis  Axelrod 
Figures  8C,  D 

Salix  wildcatensis  Axelrod,  Carnegie  Inst.  Wash.  Publ.  553,  p.  132,  1944  (see  synon- 
ymy); Chaney,  Carnegie  Inst.  Wash.  Publ.  553,  p.  341,  pi.  58,  Fig.  2,  1944. 


295 


Figure  8.  A-Arecaceae  indet.,  SDSNH  25163;  B-Juncaceae  indet.,  SDSNH  25164;  C,D-Salix  wild- 
catensis  Axelrod,  C-SDSNH  25 1  53,  D-SDSNH  25 1 52;  E,F - Populus alexanderi  Dorf,  E-SDSNH  25 1 59, 
F-SDSNH  25160.  All  specimens  x  1. 


296 


Salix  coalingensis  Dorf  (in  part),  Carnegie  Inst.  Wash.  Publ.  476,  p.  170,  pi.  4,  fig.  8, 
1937. 

Referred  sped  mens. -SDS^H  25 1 52,  25 1 53,  25 1 54. 

Remarks.  — Three  fragmentary  specimens,  the  largest  7.0  cm  long  and  2.6  pm  broad, 
represent  this  willow.  It  is  allied  to  Salix  lasiolepis  Bentham,  a  common  shrub  to  small 
tree  that  is  widely  distributed  in  the  Coast  Ranges  and  lower  Sierra  Nevada,  reaching 
southward  into  the  coastal  slopes  of  northern  Baja  California.  It  is  also  disjunct  to 
southeastern  Arizona.  In  both  areas  it  is  a  common  member  of  riparian  vegetation  in 
oak  woodlands. 

Family  Fagaceae 

Quercus  lakevillensis  Dorf 

Figure  9C 

Quercus  lakevillensis  Dorf,  Carnegie  Inst.  Wash.  Publ.  412,  p.  82,  pi.  8,  figs.  4,  5,  1930; 
Axelrod,  Carnegie  Inst.  Wash.  Publ.  590,  p.  58,  pi.  3,  fig.  4,  1950;  Axelrod,  Univ. 
Calif.  Publ.  Geol.  Sci.  121,  p.  165,  pi.  18,  figs.  2,  3,  1980. 

Referred  specimen. -SI>S^H  25158. 

Remarks.— A  single  leaf  in  the  flora  is  similar  to  those  produced  by  the  common 
California  live  oak,  Quercus  agrifolia  Nee.  The  specimen  is  oval  in  outline,  has  broad 
sinuses  and  wavering,  irregular  secondaries  that  diverge  at  moderate  angles.  The  modem 
species  is  common  in  the  Coast  Ranges  from  Sonoma  County  southward  into  Baja 
California.  Near  the  coast,  where  it  is  subject  to  regular  summer  fog,  it  forms  pure 
dense  woodlands.  Elsewhere  it  is  a  member  of  diverse  communities,  including  redwood 
forest,  closed  cone  pine  forest,  broadleaved  sclerophyll  woodland  and  occurs  also  in 
the  coastal  sectors  of  digger  pine  woodland. 

Family  Lauraceae 

Persea  coalingensis  (Dorf)  Axelrod 

Figures  9A,  B 

Persea  coalingensis  (Dorf)  Axelrod,  Carnegie  Inst.  Wash.  Publ.  553,  p.  132,  1944  (see 
synonymy  and  discussion);  Axelrod,  Univ.  CaHf  Publ.  121,  p.  112,  pi.  12,  fig. 
4;  p.  167,  pi.  19,  fig.  7;  pi.  20,  figs.  2-4,  1980. 

Referred  specimens. -^US>N¥[  25149,  25150,  25151. 

Remarks.— The  long-elliptic  leaves  of  avocado  are  the  commonest  leaf  fossil  in 
the  Chula  Vista  flora.  Most  were  broken  during  transport  into  the  marine  basin  though 
sufficient  details  of  venation  are  present  to  permit  their  certain  reference  to  this  late 
Tertiary  species.  Two  nearly  complete  specimens  tentatively  grouped  under  Persea  may 
represent  Magnolia  but  the  finer  details  of  venation  which  would  make  their  identifi- 
cation certain  are  not  preserved. 

Family  Platanaceae 

Platanus  paucidentata  Dorf 

Figures  lOD,  E 

Platanus  paucidentata  Dorf,  Carnegie  Inst.  Wash.  Publ.  412,  p.  94,  pi.  10,  figs.  4,  9; 
pi.  11,  fig.  1;  pi.  12,  fig.  1,  1930;  Axelrod,  Carnegie  Inst.  Wash.  Publ.  476,  p.  174, 
pi.  5,  figs.  4,  5,  1937;  Axelrod,  Univ.  Calif.  Publ.  Geol.  Sci.  121,  p.  113,  pi.  13, 
fig.  1;  p.  168,  pi.  20,  fig.  1,  pi.  21,  fig.  7,  1980. 

Referred  specimens. -S\)S\<^H  25155,  25156,  25157. 

Remarks.— The  typical  lobed  leaves  of  this  sycamore  are  in  the  flora.  The  large 
leaves  regularly  produced  by  the  species  were  broken  and  mangled  during  transport 
into  the  marine  basin.  However,  the  distinctive  primary  venation  and  the  lobed  nature 
of  the  leaves  permits  their  ready  identification. 


297 


Figure  9.  A,B-Persea  coalingensis  (DorO  Axelrod,  A-SDSNH  25150,  B-SDSNH  25151;  C-Quercus 
lakcvillcnsis  DorfSDSNH  25158;  D,E- Platanus paucidentataDorf ,  D-SDSNH  25155,  E-SDSNH  25157. 
All  specimens  x  1. 


298 


The  fossil  species  has  leaves  similar  to  those  of  Platanus  racemosa  Nuttall,  found 
along  stream  banks  and  floodplains  in  oak  woodland  vegetation  from  central  California 
southward  into  Baja  California.  An  allied  species.  P.  wrightii  Watson,  is  in  Arizona 
and  border  areas.  The  fossil  species,  P.  paucidentata,  is  common  in  the  Miocene  and 
Pliocene  floras  of  California.  * 

Acknowledgments 

The  cooperation  of  Watt  Industries,  Rancho  Santa  Fe,  California,  and  Financial 
Scene,  San  Diego,  California,  is  gratefully  acknowledged  for  permitting  and  supporting 
the  collection  of  fossils  on  their  Chula  Vista  property. 

In  addition  the  Planning  Department,  City  of  Chula  Vista  and  especially  Douglas 
Reid  of  that  office  are  to  be  commended  for  realizing  the  significance  of  paleontological 
resources  within  their  jurisdiction. 

Special  thanks  are  extended  to  Richard  A.  Cerutti  and  Bradford  O.  Riney  who 
collected  most  of  the  paleobotanical  material  discussed  herein. 

The  junior  author's  work  on  this  project  was  supported  in  part  by  grants  from  the 
Parker  Foundation,  the  Scripps  Foundation,  and  the  J.  F.  Sefton  Foundation.  The 
additional  support  of  Joseph  and  Joanne  Parker  of  Comado,  California,  is  also  gratefully 
acknowledged. 

The  senior  author  worked  on  the  project  during  tenure  of  National  Science  Foun- 
dation Grant  DEB  80-25525,  which  is  acknowledged  with  thanks. 

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I  IU9    ^^  ^  TRANSACTIONS 

lob"^    .^W  OF  THE  SAN  DIEGO 

SOCIETY  OF 
NATURAL  HISTORY 


Volume  20  Number  16  pp.  301-312       20  November  1984 


Relationships  within  Eumalacostracan  Crustacea  ^p 


Frederick  R.  Schram 

San  Diego  Natural  History  Museum,  P.O.  Box  1390,  San  Diego,  CA  92112  USA 


"^ 


Abstract.  A  cladistic  analysis  was  performed  on  20  constituent  higher  taxa  within  the  Eumala- 
costraca  based  on  3 1  characters  of  external  anatomy.  Variants  of  the  most  parsimonious  scheme  are 
presented,  and  the  effects  of  tolerating  different  levels  of  uncertainty  are  evaluated.  It  is  concluded  that: 
1)  while  the  basic  outline  of  Caiman's  (1904)  taxonomy  of  Eumalacostraca  might  be  utilized,  the 
arrangement  within  peracarids  postulated  by  Siewing  (1956)  cannot  be  maintained;  2)  the  Baupldne 
approach  of  Schram  (1981)  has  some  merit  and  some  of  the  controversial  higher  taxonomic  groupings 
of  eumalacostracan  "orders"  originally  indicated  by  that  method  are  vindicated;  3)  the  idea  that  the 
carapace  is  a  derived  feature  within  eumalacostracans,  advanced  by  Dahl  (1983),  can  be  maintained 
only  if  a  high  level  of  homoplasy  is  tolerated;  4)  the  concept  of  a  taxon  Mysidacea  seems  best  abandoned. 


Introduction 

The  basic  modem  classification  of  eumalacostracan  crustaceans  was  outlined  by 
Caiman  (1904,  1909)  with  little  reference  at  that  time  to  what  the  details  of  phyletic 
relationships  between  and  within  groups  might  have  been.  However,  it  was  Siewing 
(195 1,  1956)  who  presented  a  phylogenetic  tree  for  eumalacostracans  widely  subscribed 
to  by  subsequent  authorities  (e.g..  Fryer  1964,  Hessler  1969). 

Recently,  however,  the  Calman/Siewing  scheme  for  Eumalacostraca  sensu  sthcto 
has  been  questioned.  Schram  (1981)  recognized  basic  structural  plans  within  the  Eu- 
malacostraca, but  the  methodology  he  employed  was  limited  by  the  number  of  char- 
acters that  could  be  handled  essentially  by  pencil  and  paper.  However,  the  method  was 
helpful  in  three  respects.  First,  it  illustrated  a  variable  range  of  possible  dendrograms. 
Each  variant  dendrogram  was  constrained  by  which  characters  received  initial  emphasis 
and,  thus,  demonstrated  the  basic  range  of  uncertainty  that  must  be  implicit  in  any 
phylogenetic  analysis.  Second,  the  analysis  suggested  certain  "supraordinal"  relation- 
ships which  were  a  bit  unexpected,  especially  within  the  peracarid  groups.  For  example, 
isopods  and  amphipods  were  united;  and  cumaceans,  tanaids,  and  spelaeogriphaceans 
were  allied  to  each  other  with  some  suggestion  of  more  distant  possible  links  of  these 
to  thermosbaenaceans.  Third,  the  method  also  produced  a  number  of  "paper"  Baupldne 
which  were  not  occupied  or  had  yet  to  be  discovered.  Implicit  in  these  hypothetical 
morphotypes  was  the  idea  that  if  the  method  had  any  merit  at  all,  some  of  those 
"empty"  Baupldne  might  eventually  be  found. 

Watling  (1981,  1983)  questioned  the  unity  of  the  superorder  Peracarida  as  a  natural 
taxon.  He  produced  two  different  cladograms  for  the  peracarids.  His  stated  purpose  (in 
Schram,  1983:347)  was  to  search  for  ".  .  .  Baupldne  that  include  the  fine  structure  .  .  .  ," 
and  he  believed  that  ".  .  .  the  first  step  in  the  analysis  is  to  look  at  all  these  structures 
for  pattern  .  .  .  ."  In  this  respect  Watling  (1983)  performed  a  useful  function  by  focusing 
attention  on  characters  that  had  largely  been  overlooked  by  previous  workers  such  as 
mandible  function,  maxillipede  form,  and  patterns  of  arterial  circulation. 

Dahl  (1983)  formally  proposed  a  concept  that  had  been  implicit  in  several  of  his 
earlier  papers,  viz.,  that  the  lack  of  a  carapace  is  a  primitive  feature,  that  the  evolution 
of  the  carapace  had  occurred  independently  several  times,  and  that  Caiman's  caridoid 


302 


facies  was  a  set  of  convergent  phenomena.  Dahl  presented  some  interesting  observations 
on  comparative  carapace  development  related  to  these  ideas.  Watling  (especially  1983) 
acknowledged  his  indebtedness  to  Dahl's  concept  of  the  carapace  as  a  derived  feature. 

Finally,  Hessler  (1983)  produced  a  "defense"  of  the  caridoid  facies  in  which  he 
attempted  cladistic  analysis  of  the  Siewing  scheme  for  peracarids  in  a  mpre  formal 
manner  than  had  ever  been  presented  before.  Hessler's  study,  however,  produced  a 
scheme  in  which  the  Siewing  arrangement  of  taxa  could  be  retained  only  by  tolerating 
a  great  deal  of  convergence  (10  of  his  23  characters  are  convergent  in  whole  or  part  to 
one  or  more  of  the  others). 

Thus,  several  items  bear  on  the  problem  of  eumalacostracan  relationships  and 
demand  some  sort  of  a  resolution.  First,  is  the  need  to  assess  relationships  among 
eumalacostracans  by  the  use  of  as  many  characters  as  possible,  and  use  these  characters 
across-the-board  for  all  taxa,  fosssil  and  Recent.  Second,  a  test  is  demanded  both  for 
Dahl's  concept  of  the  carapace  as  a  derived  feature,  as  well  as  some  of  the  "strange" 
higher  groupings  suggested  by  Schram  (1981).  And  third,  it  is  necessary  to  arrive  at  a 
scheme  which  will  group  the  taxa  in  question  strictly  by  their  shared  derived  character 
states  with  the  fewest  number  of  convergences  possible. 

Method 

One  way  to  analyze  large  numbers  of  characters  and  taxa  so  as  to  achieve  the  most 
parsimonious  arrangement,  based  solely  on  shared  derived  characters,  is  to  use  one  of 
the  various  versions  available  for  the  Wagner  78  program.  For  this  analysis,  I  decided 
to  "break  up"  certain  large  and  diverse  traditional  eumalacostracan  taxa  and  treat  their 
components  as  separate  units  to  test  both  the  viability  of  such  taxa  and  the  "reason- 
ableness" of  the  characters  used.  To  this  end  the  suborders  of  Mysidacea  (Mysida, 
Lophogastrida,  and  Pygocephalomorpha)  and  of  Decapoda  as  outlined  by  Burkenroad 
(1981)  (Dendrobranchiata,  Eukyphida,  Euzygida,  and  Reptantia)  were  evaluated  as 
separate  entities.  The  choice  of  taxa  for  the  decapods  was  somewhat  arbitrary  since, 
for  example,  Felgenhauer  and  Abele  (1983)  break  the  Eukyphida  into  two  groups 
coequal  with  the  others,  Procarididea  and  Caridea. 

The  3 1  characters  used  for  this  analysis  were  selected  by  repeated  trial  and  error 
(as  is  standard  in  any  computer-generated  cladistic  treatment  of  such  data),  rejecting 
potentially  useful  characters  which  had  low  consistency  indexes  (i.e.,  high  homoplasy 
values).  The  ultimate  aim  of  these  initial  assessments  of  potentially  useful  characters 
was  to  yield  the  most  parsimonious  and  congruent  cladogram  possible.  Only  characters 
that  could  be  assessed  for  all  groups  relatively  unambiguously  were  used.  For  example, 
I  did  not  use  the  lacinia  mobilis  because  I  do  not  feel  its  homology  has  been  dem- 
onstrated. As  has  been  shown  recently  (Dahl  and  Hessler,  1982),  this  character  is  not 
only  present  in  several  groups,  but  is  developed  differently  in  these  taxa.  How  can  one 
compare  the  massive  laciniae  of  lophogastrids  with  the  rather  delicate  ones  in  other 
peracarids?  Or,  how  are  larval  laciniae  to  be  judged  in  comparison  to  those  of  adults? 
More  needs  to  be  known  about  the  development  and  functional  morphology  of  laciniae 
before  they  can  be  adequately  assessed  in  a  phylogenetic  analysis.  Other  characters 
were  not  used  because,  while  they  serve  to  characterize  specific  taxa,  they  are  quite 
homoplastic  and  are  known  to  occur  convergently  in  widely  separated  groups.  For 
example,  the  use  of  the  presence  of  second  or  third  maxillipedes  was  avoided  in  the 
final  analysis  since  it  only  served  to  confirm  groupings  achieved  more  effectively  by 
singularly  derived  features.  The  characters  eventually  settled  upon  are  given  in  Table 
1,  the  numbers  indicated  corresponding  to  those  used  in  the  cladograms. 

The  program  was  run  using  several  different  outgroups,  Hoplocarida,  leptostracan 
Phyllocarida,  and  a  hypothetical  ancestor  arbitrarily  designated  primitive  for  all  31 
characters.  No  differences  in  any  of  the  resultant  eumalacostracan  cladograms  were 
noted.  Among  other  parameters,  the  program  also  calculated  total  lengths  of  trees  (i.e., 
the  total  number  of  incidences  of  derived  characters  in  the  cladogram)  and  the  total 
homoplasy  value  (i.e.,  a  measure  of  the  total  array  of  convergences  and  character 


303 


Table  1 .     Opposing  list  of  character  states  used  in  the  analysis  of  relationships  within  Eumalacostraca. 
Numbers  correspond  to  those  used  in  cladograms. 


Primitive 


Derived 


1 .  Non-caridoid  musculature 

2.  No  zoeal  larvae 

3.  Carapace  not  fused  to  all  thoracomeres 

4.  No  petasma 

5.  First  thoracomere  free  of  head 

6.  Maxillipede  with  epipodite 

7.  No  brood  pouch  formed  by  first  pleopod 

8.  No  scaphognathite 

9.  First  thoracopod  unmodified 

10.  Maxillipede  endopod  robust 

1 1 .  Eggs  not  brooded  on  pleopods 

12.  No  caridean  lobe 

13.  Biramous  thoracopods 

14.  All  pleopods  present 

15.  First  thoracopod  unmodified 

16.  Pereiopodal  epipodite  gills 

17.  First  thoracopod  unmodified 

18.  No  marsupium 

19.  Thoracic  endopods  non-filtratory 

20.  No  male  cones 

2 1 .  Thoracic  coxae  unmodified 

22.  Eyes  stalked  or  lobed 

23.  One  pair  of  uropods 

24.  Pleopods  non-respiratory 

25.  Carapace  not  short 

26.  Eggs  not  brooded  under  carapace 

27.  Maxillipedal  epipodite  if  present  simple 

28.  Rostrum  simple 

29.  Thoracic  exopods  non-respiratory 

30.  Maxillipedal  epipodite  as  a  single  segment 


31.  Carapace 

Character  reversal  used  in  analysis  portrayed  in  Figure  3 

3 1 .  No  carapace  carapace 


caridoid  musculature 

zoeae 

carapace  fused  to  all  thoracomeres 

petasma 

first  thoracomere  fused  to  head 

maxillipede  without  epipodite 

brood  pouch  between  first  pleopod  and 
venter  of  thorax 

scaphognathite  on  maxilla 

maxillipedes  with  lamellate  protopod, 
coxal/basal  endites  directed  mediad 

maxillipede  endopod  flagelliform 

eggs  brooded  on  pleopods 

caridean  lobe  on  maxillipede 

uniramous  thoracopods 

pleopods  lost  or  reduced 

maxillipedes  with  tendency  to  form 
gnathobasic  endites,  endopod  pediform 

no  pereiopodal  epipodite  gills 

maxillipedes  with  basal  endites  lobate 
and  directed  distad 

oostegite  marsupium 

thoracic  endopods  filtratory 

male  cones 

thoracic  coxal  plates 

eyes  sessile 

more  than  one  pair  of  uropods 

pleopods  respiratory 

carapace  short 

eggs  brooded  under  carapace 

epipodite  specialized  as  cup-  or  spoon- 
like respiratory  organ 

pseudorostrum  and  maxillipedal  siphons 

thoracic  exopods  respiratory 

epipodite  with  tendency  to  form  as  2-3 
segments 

carapace  absent 


reversals  in  the  cladogram).  These  factors  proved  useful  in  qualitatively  comparing 
different  cladograms. 


Results 

The  computer  program  generated  several  variant  cladograms.  That  variant  which 
was  most  parsimonious  and  yielded  the  fewest  number  of  convergences  and  character 
reversals  is  given  in  Figure  1.  In  the  series  of  cladograms  summarized  in  Figures  1-3, 
previous  outgroup  analysis  indicated  that  the  presence  of  a  carapace  should  be  treated 
as  primitive  because  it  is  present  in  all  hoplocaridans  and  phyllocaridans.  As  can  be 
seen,  the  program  produced  (Fig.  1)  an  unresolved  polychotomy  with  four  branches  at 
the  base  of  the  Eumalacostraca:  eucarids,  belotelsonids,  syncarids,  and  waterstonellid/ 
peracarids.  A  variant  of  this  scheme  (Fig.  2)  yields  an  unresolved  polychotomy  of  five 
branches.  Although  the  latter  cladogram  has  the  same  number  of  convergences  as  the 
former,  it  is  somewhat  shorter  than  that  of  Figure  1 .  A  convergence  in  the  secondary 
reevolution  of  pereiopodal  epipodite  gills  between  Mysida  and  Amphipoda  is  traded 
off  for  a  convergence  in  the  primary  loss  of  pereiopodal  epipodite  gills  in  Watersto- 


304 


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Figure  1 .  Cladistic  relationships  of  component  taxa  of  the  Eumalacostraca,  the  presence  of  a  carapace 
considered  primitive.  This  is  the  shortest  cladogram  with  the  lowest  homoplasy  value,  the  base  of  the 
cladogram  with  an  unresolved  quadrochotomy  between  eucarids,  belotelsonids,  syncarids,  and  waterstonel- 
lids/peracarids.  D  primitive,  ■  derived. 


nellidea  and  the  "peracarid"  line  above  Mysida.  Wagner  78  is  designed  to  produce  the 
best  resolved  cladogram  possible  from  the  data  given  and,  thus,  the  preferred  version 
is  that  seen  in  Fig.  1 .  If  on  the  other  hand  we  wish  to  tolerate  a  slightly  greater  degreee 
of  uncertainty  (Schram,  1983),  then  we  may  choose  the  variant  of  Fig.  2  in  which 
peracarids  can  be  recognized  as  a  distinct  lineage.  The  relationships  indicated  in  Figure 
1 ,  however,  are  not  without  considerable  biological  interest.  The  thrust  of  the  early 
evolution  of  the  waterstonellid/peracarid  line  was  towards  increasing  specialization  of 
thoracopods.  First  the  primitive  respiratory  epipodites  were  lost,  then  oostegites  and 
maxillipedes  were  evolved,  and  finally  some  further  specializations  occurred  in  specific 
lineages  such  as  filtratory  endopods  in  mysidans  (Attramadal,  pers.  comm.),  and  further 
maxillipedal  and  ambulatory  modifications  in  pygocephalomorphs  (Schram,  1974). 


305 


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Figure  2.  Cladistic  relationships  of  component  taxa  of  the  Eumalacostraca,  the  presence  of  a  carapace 
considered  primitive.  A  variant  cladogram  from  that  of  Fig.  1  exhibiting  ( 1 )  unresolved  quintichotomy 
at  the  base  that  allows  a  separation  of  waterstonellids  and  peracarids  (which  would  shorten  the  tree,  not 
involve  any  change  in  the  number  of  convergences  over  that  of  Fig.  1 ,  but  would  inject  a  higher  level  of 
uncertainty  into  the  cladogram)  and  (2)  an  association  of  pygocephalomorphs  as  a  sister  group  of  the  brachy- 
caridans  (which  would  not  involve  a  lengthening  of  the  cladogram  but  would  inject  one  extra  character 
reversal  over  that  seen  in  Fig.  1).  D  primitive,  ■  derived. 


Several  interesting  points  emerge  from  these  analyses.  Many  of  the  more-or-less 
controversial  higher  taxa  (Cohorts  and  Orders)  of  Schram  (1981),  emerge,  viz.,  Hemica- 
ridea  (Cumacea,  Tanaidacea,  and  Spelaeogriphacea),  Brachycarida  (Hemicaridea  and 
Thermosbaenacea),  Eucarida  (Euphausiacea,  Amphionidacea,  and  Decapoda),  and 
Acaridea  (Isopoda  and  Amphipoda).  The  latter  also  seems  to  bear  some  relationship 
to  a  yet  unnamed  new  order  being  proposed  by  T.  Bowman,  R.  Hessler,  and  H.  Sanders 
which,  interestingly,  seems  to  fill  one  of  the  "unoccupied"  5aw/7/a>2£' of  Schram  (1981). 
On  the  other  hand,  some  taxa  derived  from  Schram  (1981)  do  not  seem  viable:  e.g., 


306 


.&       -^ 


1^ 


Figure  3.  Cladogram  based  on  the  same  character  data  as  that  used  in  cladograms  of  Figure  1 ,  but  analyzing 
only  living  groups  and  excluding  the  4  extinct  taxa,  Palaeocaridacea,  Belotelsonidea,  Waterstonellidea,  and 
Pygocephalomorpha.  D  primitive,  ■  derived. 


Arthrostraca  in  the  sense  of  Haeckel  (1896),  Giesbrecht  (1913),  or  Grobben  (1919) 
which  unites  all  carapaceless  syncarid  and  acaridean  forms;  or  Mysoida  (Belotelsonidea, 
Mysidacea,  and  Waterstonellidea),  which  seems  invalid  as  a  cladistic  or  taxonomic 
unit. 

I  decided  to  test  the  effect  on  the  overall  scheme  of  relationships  when  the  fossil 
taxa  were  excluded  from  consideration  (Fig.  3).  Little  change  was  noted  except  to  ally 


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307 


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^ 


Figure  4.  Cladogram  with  all  taxa  and  character  data  the  same  as  in  Fig.  1,  except  the  scoring  of  character 
31  is  reversed  and  the  presence  of  a  carapace  is  treated  as  a  derived  feature,  as  favored  by  Dahl  (1983).  D 
primitive,  ■  derived. 


syncarids  and  eucarids  as  sister  groups.  Some  slight  modifications  occurred  in  the 
arrangement  of  higher  eucarids,  but  otherwise  the  basic  relationships  of  the  taxa  of 
Fig.  1  are  preserved.  The  total  homoplasy  value  (a  measure  of  the  amount  of  con- 
vergence) is  somewhat  higher  (388  vs.  372)  in  the  non-fossil  scheme  than  in  that  which 
includes  the  extinct  groups,  although  the  total  length  of  both  trees  is  not  that  much 
different,  44  without  fossils  as  opposed  to  46  with  extinct  groups  included. 


308 


I  also  tested  the  contention  of  Dahl  (1983)  that  the  carapace  is  a  derived  feature, 
the  lack  of  a  carapace  being  viewed  as  primitive  (Fig.  4).  This  test  resulted  in  a  somewhat 
longer  cladogram  than  those  in  Figure  1  (47  vs.  46),  but  one  which  has  a  dramatically 
higher  total  homoplasy  value  (510  vs.  372).  Similar  results  were  obtained  when  the 
data  based  on  Dahl's  concept  were  run  without  extinct  taxa.  It  would  seem, ♦therefore, 
that  the  suggestion  by  Dahl  that  the  carapaceless  state  is  the  primitive  one  for  eumalacos- 
tracans  engenders  a  more  complicated  and  less  parsimonious  array  of  relationships. 
Note,  however,  that  the  relationships  within  peracarids  persist,  including  that  of  break- 
ing apart  Mysidacea. 

Discussion 
Characters  difficult  to  use 

Certain  characters  were  deliberately  not  used  here  though  they  have  found  wide- 
spread employment  in  the  taxonomy  and  phylogenetic  discussions  of  Eumalacostraca 
by  various  authors. 

In  eucarids,  although  the  structure  of  maxillipedes  was  used  (lamellate  appendages 
with  endites  directed  mediad),  the  number  of  them  was  not  (three  maxillipedes  and 
thus  the  name  decapod).  The  anatomical  and  functional  state  of  thoracopods  in  higher 
eucarids  is  actually  more  varied  than  one  would  be  led  to  believe  from  the  automatic 
connotation  engendered  by  the  term  "decapod."  In  several  instances,  e.g.,  many  Den- 
drobranchiata,  the  so-called  third  maxillipede  is  actually  more  "pediform"  in  structure 
and  function  than  "maxillipediform."  Also,  certain  "pereiopods"  actually  have  little 
locomotory  function  but  are  utilized  in  food  acquisition  and  processing  as  well  as 
defense.  For  example,  in  euzygids  (=stenopodids)  the  fourth  and  fifth  thoracopods  are 
chelipedes  and  directed  anteriad  towards  the  mouth  field  resulting  in  a  hexapodous 
condition  instead  of  a  decapodous  one  in  these  animals.  In  astacideans  the  characteristic 
great  chelipedes  of  the  fourth  thoracopods  serve  in  food  procurement  and  defense, 
making  the  animal  functionally  octopodous.  So  while  there  are  good  maxillary  and 
maxillipedal  features  which  can  serve  to  delineate  a  taxon  Decapoda,  ironically  true 
decapody  is  not  a  particularly  good  character  to  assist  in  such  delineation. 

Another  feature  taken  for  granted  in  discussion  of  eumalacostracan  evolution  is 
the  fusion  of  the  first  thoracomere  to  the  cephalon.  Bathynellacea,  of  course,  do  not 
have  this  fusion.  The  Carboniferous  genera  Belotelson  and  Waterstonella  apparently 
had  free  first  thoracomeres  as  well,  as  they  lacked  maxillipedes  altogether.  Hence,  it  is 
imperative  to  resolve  whether  or  not  all  living  forms  with  a  carapace  do  or  do  not  fuse 
the  first  thoracomere  to  the  head.  For  example,  euphausiaceans  lack  a  maxillipede, 
have  the  carapace  fused  to  the  thoracomeres,  but  have  all  thoracomeres  associated 
together  separate  from  the  maxillary  segment.  Examination  of  mysidaceans  revealed 
a  variable  state  of  affairs.  Lophogastridans,  with  their  well-developed  maxillipedes, 
closely  associate  the  first  thoracomere  with  the  cephalon  and  separate  it  from  the  second 
and  following  thoracomeres.  However,  in  the  mysidan  Neomysis  americana  there  is  a 
separation  of  the  maxillary  from  the  thoracic  segments,  with  all  eight  sets  of  thoracopods 
closely  associated  and  separated  by  a  distinct  skeletal  bar  from  the  more  anterior 
mouthparts.  So  in  mysidans  the  first  thoracomere  is  clearly  not  fused  to  the  cephalon, 
although  there  is  a  tendency  to  develop  maxillipedes.  This  feature  serves  to  break  apart 
the  taxon  Mysidacea,  making  Mysida  a  sister  group  to  all  other  peracarids. 

I  also  excluded  three  characters  which  have  been  asserted  as  distinctly  peracaridan, 
including  the  lacinia  mobilis  mentioned  above.  The  presence  of  a  manca  stage  is 
frequently  cited  as  a  characteristic  of  peracarids.  Generally  workers  used  this  feature 
as  if  they  were  dealing  with  a  manca  larva.  Mancas,  however,  are  not  to  be  equated 
with  the  zoea,  cypris,  or  other  larval  types  which  have  considerable  cladistic  merit  (see 
for  example  character  2).  A  "manca"  is  a  stage  of  development  which  can  have  various 
forms  of  expression  (Newman  1983).  Amphipods  are  generally  said  to  lack  a  manca, 
yet  some  hyperiids  are  freed  from  the  female  in  a  virtual  manca  state  (Laval  1980). 
Some  adults  express  a  permanent  manca  condition,  e.g.,  the  genus  Thermosbaena. 


309 


Manca  stages  also  occur  outside  the  peracarids,  e.g.,  bathynellaceans  hatch  in  an  extreme 
"mancoid"  condition  lacking  several  of  the  posterior  thoracopods  and  in  the  adults  of 
some  forms  the  last  thoracopod  can  be  missing  or  greatly  atrophied.  The  presence  or 
absence  of  a  manca  may  be  better  understood  in  terms  of  constraints  placed  on  de- 
velopment by  egg  size  (e.g.,  Steele  and  Steele  1975).  Characters  of  marsupial  and 
maxillipedal  form  alone  can  be  used  to  delineate  peracarids  more  securely;  and  while 
the  presence  of  a  manca  stage  may  assist  in  this  delineation,  it  is  not  as  unambiguous 
as  one  would  suppose. 

The  same  observation  can  be  made  of  the  monocondylic  coxa/basis  articulation 
recently  noted  by  Hessler  (1982).  This  character  might  appear  to  be  a  useful  congruent 
feature  towards  establishing  a  concept  of  Peracarida.  However,  it  has  a  variety  of 
expression  difficult  at  this  time  to  evaluate.  For  example,  the  monocondyle  variously 
arises  from  positions  that  are  either  lateral  (tanaids),  purely  posterior  (Spelaeoghphus), 
or  postero-lateral  (all  other  peracarids).  Nor  do  all  thoracopods  have  this  joint.  In 
tanaids  the  third  through  fifth  limbs  have  a  dicondylic  joint  whereas  only  the  sixth 
through  eighth  have  the  distinct  monocondyle.  Completely  aberrant  condyle,  muscle, 
and/or  joint  arrangements  are  seen  in  amphipods  and  mysidaceans.  These  latter  two 
groups  also  display  different  degrees  of  expression  of  these  features  throughout  the 
whole  thoracopodal  limb  series.  Although  coxal/basal  structure  seems  to  second  per- 
acarid  monophyly,  problems  with  variety  of  expression  and  assessment  of  polarities 
between  these  variations  preclude  its  use  here. 

Characters  rejected  for  use 

Two  suites  of  traditional  characters  were  completely  rejected,  viz.,  those  of  gut 
structure  and  embryo  flexion  which  have  played  so  prominent  a  role  in  the  work  of 
Siewing,  and  which  resulted  in  the  diametric  separation  of  isopods  and  amphipods. 
The  more  that  is  discovered  about  gut  morphology,  the  more  it  seems  that  the  digestive 
system  is  too  plastic  to  yield  any  useful  data  for  phylogenetic  analysis.  Kunze  (1981, 
and  personal  communication)  has  noted  that  the  anatomy  of  the  stomach  of  isopods 
is  closely  tied  to  feeding  habits.  Ide  (1892)  and  Naylor  (1955)  provided  details  of  gut 
structure  in  Idotea  identical  to  that  supposedly  characteristic  of  amphipods,  including 
an  anteriorly  directed  mid-dorsal  caecum  in  /.  tricuspidata.  Carol  Diebel  {pers. 
comm.)  is  finding  that  stomach  structure  among  hyperiid  amphipods  is  so  diverse  as 
to  be  uncharacterizable  because  of  adaptations  to  particular  feeding  strategies. 

The  other  character  rejected  here,  but  given  great  weight  by  Siewing,  is  whether 
the  developing  embryo  is  flexed  ventrally  or  dorsally  within  the  egg  membranes.  First, 
few  studies  within  and  between  groups  of  peracarids  have  been  performed  to  determine 
the  distribution  of  these  states.  Second,  one  of  these  flexures  must  be  primitive  and 
the  other  advanced.  As  such,  only  one  of  them  can  be  used  to  characterize  one  of  the 
groups  which  possesses  it,  but  they  are  not  both  derived  characters.  It  might  appear 
that  the  dorsal  flexure  is  derived,  but  insufficient  data  exist  from  within  and  without 
peracarids  in  order  to  assess  polarity.  And  third,  flexure  in  embryos  seems  better 
understood  in  terms  of  the  mechanics  of  a  particular  developmental  sequence  rather 
than  in  terms  of  phyletic  trends.  Note  that  in  forms  with  a  ventral  flexure,  there  is 
typically  a  very  distinct  egg-nauplius  stage  in  early  development,  the  development  of 
the  teloblasts  lags  behind  that  of  the  primary  part  of  the  head.  The  development  of  a 
caudal  papilla  and  a  caudal  furrow  which  lead  to  ventral  flexure  is  thus  possibly  related 
to  the  rapid  development  of  the  naupliar  region.  In  contrast,  in  animals  with  a  dorsal 
flexure,  the  appearance  of  the  naupliar  anlagen  lags.  In  such  forms  the  teloblasts  not 
only  appear  early  in  the  sequence  of  events  around  the  blastopore,  they  initiate  their 
divisions  early  such  that  the  naupliar  and  anterior  metanaupliar  somites  appear  virtually 
simultaneously.  It  would  appear  that  because  of  the  slower  head  development  the 
proliferation  of  body  somites  is  allowed  to  occur  along  the  entire  ventral  and  posterior 
surfaces  of  the  egg  without  the  appearance  of  a  caudal  furrow  or  papilla  to  produce 
ventral  flexure.  Clearly  the  "phylogenetic  power"  of  the  apparent  differences  of  flexure 
between  isopods  and  amphipods  has  been  somewhat  overextended. 


310 


Variant  cladograms 

Two  variants  in  the  cladograms  were  produced  by  the  program  frequently  enough 
to  require  some  comment  here.  One  is  a  variation  in  the  higher  decapods  seen  in  Figures 
1  and  3.  In  one  (Fig.  1),  somewhat  more  parsimonious,  eukyphids  are  placed  as  a  sister 
group  to  euzygids  and  reptants.  In  the  other  (Fig.  3),  reptants  are  a  sister  group  of 
euzygids  and  eukyphids.  The  former  is  a  more  traditional  arrangement,  but  the  latter 
is  all  the  more  startling  in  light  of  the  pregnant  comment  of  Felgenhauer  and  Abele 
(1983)  that  it  was  their  belief  that  the  origins  of  the  so-called  "natant"  groups  of 
decapods  ".  .  .  are  to  be  found  among  those  groups  traditionally  considered  reptants." 

Indeed  the  entire  issue  of  relationships  within  the  decapods  is  under  intense  study 
right  now.  Burkenroad  (1981),  using  branchial  and  ontogenetic  characters  not  employed 
in  this  analysis,  essentially  obtained  an  arrangement  of  taxa  like  that  seen  in  Figure  3. 
However,  Felgenhauer  {personal  communication)  is  examining  various  features  of  ex- 
ternal and  internal  anatomy  of  natant  forms  in  an  attempt  to  arrive  at  an  assessment 
of  cladistic  relationships  within  decapods.  For  these  reasons,  it  may  be  wise  to  avoid 
use  of  terms  like  Decapoda  and/or  Pleocyemata  for  the  time  being,  and  rather  treat 
the  taxa  within  Eucarida  as  one  long  transition  series. 

Another  notable  variant  is  seen  in  the  higher  peracarids  between  Figs.  1  and  2. 
The  scheme  in  Fig.  2  is  slightly  less  parsimonious,  but  if  one  can  tolerate  the  ad- 
ditional character  reversal  it  entails,  then  the  arrangement  is  a  sequence  of  events  which 
is  of  considerable  biological  interest.  The  isopod/amphipod  line  seems  to  represent  one 
in  which  the  thrust  of  the  radiation  is  toward  varied  exploitation  of  food  resources 
because  of  the  great  plasticity  in  gut  structures.  The  brachycaridan  line,  especially  when 
the  pygocephalomorphs  are  associated  with  it,  seems  to  be  a  line  which  represents 
exploitation  of  reproductive  strategies.  Both  pygocephalomorphs  and  tanaids  have 
cones  on  the  males.  The  supposed  seminal  receptacles  mentioned  by  Brooks  (1962)  on 
pygocephalomorphs  bear  little  actual  resemblance  to  such  structures.  These  structures 
are  more  likely  large  genital  cones  on  the  eighth  thoracic  stemites  of  males.  The 
brachycaridan  line  is  generally  characterized  by  respiratory  specializations  of  the  max- 
illipedes  and  thoracopods.  It  is  also  a  transition  series  in  which  carapace,  pleopods, 
and  the  oostegite  brood  pouch  are  reduced  or  lost,  culminating  in  the  condition  seen 
in  the  thermosbaenaceans.  Insofar  as  the  component  taxa  are  currently  understood, 
this  line  also  exploits  reproductive  and  unusual  sexual  strategies  that  maximize  the 
number  of  offspring  from  any  one  generation  (e.g.,  see  Sieg  1983,  for  tanaidaceans,  or 
Corey  1981,  for  cumaceans). 

Conclusions 

Several  conclusions  can  be  drawn  concerning  the  analyses  made  here: 

1)  At  least  in  part,  the  taxonomic  scheme  for  the  Eumalacostraca  suggested  by  the 
identification  of  Baupldne  within  the  group  (Schram  1981)  is  supported,  especially  in 
regards  to  peracarid  types.  The  idea  of  a  taxon  Arthrostraca  is  not  favored,  but  the 
reassociation  of  isopods  with  amphipods  in  the  sense  of  the  old  taxon  Edriophthalma, 
and  the  linking  of  short  carapace  forms,  does  have  some  merit. 

2)  If  some  degree  of  uncertainty  is  accepted,  then  the  relationships  within  Peracarida 
postulated  by  Slewing  (1951,  1956)  can  be  subscribed  to,  but  only  if  considerable 
multiple  convergences  can  be  tolerated  within  a  distinctly  unparsimonious  scheme. 

3)  The  concept  of  the  carapace  as  a  derived  feature  in  the  sense  of  Dahl  (1983)  is 
acceptable  only  by  tolerating  a  great  many  more  convergences  than  occur  when  the 
presence  of  a  carapace  is  viewed  as  primitive. 

4)  The  concept  of  a  formal  taxon  Mysidacea  seems  best  abandoned.  The  three 
subtaxa  traditionally  placed  within  it  (Lophogastrida,  Mysida,  and  Pygocephalomor- 
pha)  are  distinct  from  each  other  regardless  of  whether  the  presence  of  a  carapace  is 
considered  primitive  or  derived. 

What  taxonomy  of  Eumalacostraca  should  be  derived  from  all  this?  The  eucarids 
are  destined  for  some  kind  of  realignment,  especially  of  the  higher  taxa.  The  phylo- 


311 


genetic  integrity  of  the  brachycaridans  is  stable  enough,  whether  or  not  pygocephalo- 
morphs  are  closely  associated  with  them.  The  resolution  of  relationships  within  the 
edriophthalman  branch  must  await  the  description  and  evaluation  of  the  new  order  of 
Bowman,  Hessler,  and  Sanders,  as  well  as  a  reevaluation  of  relationships  within  isopods 
and  amphipods  using  a  careful  analysis  of  character  states  in  all  subgroups  therein.  In 
regards  to  the  latter,  we  may  resurrect  the  old  taxon  Laemodipoda,  wherein  caprellids 
and  cyamids  are  separated  as  sister  groups  off  by  themselves.  Such  a  study  is  currently 
under  way. 

It  is  my  intent  here  to  point  out  two  things.  First,  there  is  merit  in  carefully  reflecting 
on  what  are  the  constituent  structural  plans  expressed  within  any  particular  taxon,  alert 
to  the  fact  that  any  particular  Bauplan  may  or  may  not  be  developed,  or  may  or  may 
not  be  the  basis  of  an  extensive  radiation.  Second,  regardless  of  the  ongoing  philo- 
sophical and  in  large  part  tautological  debate  on  taxonomic  theory,  we  must  make 
some  organized  careful  evaluations  of  characters  and  what  their  condition  and  polarity 
might  be  throughout  all  members  of  a  group.  These  are  problems  which  have  been  all 
too  often  neglected  in  the  history  of  crustacean  studies,  but  are  not  unique  to  the  study 
of  these  arthropods. 

Acknowledgments 

Special  thanks  must  be  extended  to  Dr.  Richard  Brusca,  San  Diego  Natural  History 
Museum  and  Allan  Hancock  Foundation,  for  his  valuable  collaboration  in  the  ongoing 
evaluation  of  character  states  in  malacostracans.  Without  his  expertise  and  encour- 
agement this  study  would  not  have  been  undertaken.  Mr.  Ernest  Iverson,  Allan  Hancock 
Foundation,  ran  the  various  series  of  data  sets  with  the  Wagner  78  program  using  the 
computer  facilities  of  the  University  of  Southern  California.  Graphic  work  was  done 
by  Susan  Mc Williams,  San  Diego  Museum  of  Natural  History.  Valuable  criticism  of 
the  manuscript  has  been  offered  by  R.  Brusca,  G.  Brusca,  P.  Delaney,  M.  Grygier,  R. 
Hessler,  and  W.  Newman. 

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Sieg,  J.    1983.    Evolution  of  Tanaidacea.  Crusta- 
cean Issues  1:229-256. 


Slewing,  R.  1951.  Besteht  ein  engere  Verwand- 
schaft  zwischen  Isopoden  und  Amphipoden. 
Zoologisches  Anzieger  147:166-180. 

.    1956.    Untersuchungen  zur  Morphologic 

der  Malacostraca.  Zoologisches  Jahrbuch,  Ab- 
teilung  Anatomie  751:39-176. 

Steele,  D.  H.,  and  V.  J.  Steele.  1975.  *Egg  size  and 
duration  of  embryonic  development  in  Crus- 
tacea. Internationale  Revue  der  gesamten  Hy- 
drobiologie  60:71 1-715. 

Watling,  L.  1981.  An  alternative  phylogeny  of 
peracarid  crustaceans.  Journal  of  Crustacean 
Biology  1:201-210. 

.  1983.  Peracaridan  disunity  and  its  bear- 
ing on  eumalacostracan  phylogeny  with  a  re- 
definition of  eumalacostracan  superorders. 
Crustacean  Issues  1:213-228. 


■  ■■/. 


,_  TRANSACTIONS 

"^«»4p/^^  OF  THE  SAN  DIEGO 

JUn  society  of 

0  1955  natural  history 


Volume  20  Number  17  pp.  313-336        30  January  1985 


History  and  status  of  the  avifauna  of  Isla  Guadalupe,  Mexico 

Joseph  R.  Jehl,  Jr. 

Hubbs-Sea  World  Research  Institute.  1700  South  Shores  Road,  San  Diego,  California  92109,  USA 

William  T.  Everett 

San  Diego  Natural  History  Museum,  P.O.  Box  1390,  San  Diego.  California  92112,  USA 

Abstract.  Since  1954,  renewed  interest  in  Isla  Guadalupe,  stimulated  by  the  research  of  the  late 
C.  L.  Hubbs,  has  resulted  in  much  new  information  on  the  avifauna.  In  this  paper  we  review  the 
status  of  the  birdlife  through  1982,  provide  an  historical  review  of  the  research,  including  information 
on  the  timing  and  extent  of  the  many  expeditions,  and  provide  a  bibliography. 

Resumen.  Desde  1 954,  interes  renovado  en  la  Isla  Guadalupe,  promovido  por  las  investigaciones 
del  Dr.  C.  L.  Hubbs,  ha  resultado  en  mucha  nueva  informacion  sobre  la  avifauna.  En  esta  presentacion 
detallamos  lo  conocido  sobre  la  historia  natural  de  los  aves  hasta  1982,  damos  un  resumen  historico 
sobre  las  investigaciones,  incluyendo  informacion  sobre  el  estacionamiento  y  duracion  de  las  expedi- 
ciones  y  ofrecemos  una  bibliografia. 


Introduction 

Because  of  its  unique  plant  and  animal  life,  Isla  Guadalupe,  Mexico,  has  fascinated 
biologists  since  the  time  of  its  scientific  "discovery"  in  1875.  This  rugged  and  remote 
volcanic  island,  220  miles  south  of  the  Mexico-United  States  boundary  and  160  miles 
west  of  the  peninsula  of  Baja  California,  was  the  home  of  ten  endemic  species  or 
subspecies  of  birds  (an  eleventh  has  recently  been  proposed).  But  the  history  of  the 
birdlife  "is  a  sad  one  of  reduction  and  extermination  through  destruction  of  habitat  by 
feral  goats,  predation  by  introduced  house  cats,  and  regrettably,  some  excess  of  zeal  by 
collectors"  (Howell  and  Cade  1954:  see  also  Huey  1924,  1925),  and  many  of  the 
endemics  are  gone. 

Much  has  been  written  about  the  avifauna,  and  the  history  of  some  of  the  extinct 
forms  has  been  well  documented.  Ridgway  (1876),  Bryant  (1887(2),  Thayer  and  Bangs 
(1908)  and  Hanna  (1925)  provided  comprehensive  reviews  of  the  birdlife  and  these 
were  made  current  by  Howell  and  Cade  in  1954.  Since  then,  sufficient  new  information 
has  been  obtained  to  prompt  a  further  compilation. 

For  a  description  of  the  island  and  its  general  ecological  settings  and  geology  see 
Howell  and  Cade  (1954),  Lewis  (1971),  and  Johnson  (1953).  A  good  summary  is 
provided  by  Lindsay  (1966:2);  who  wrote:  "Guadalupe  Island  is  about  22  miles  long 
from  north  to  south,  and  four  to  six  miles  wide  over  most  of  its  length.  The  highest 
part  is  at  the  north  end,  where  magnificent  sheer  cliffs  tower  over  the  sea  or  narrow 
beaches  of  cobbles  and  sand.  The  central  part  is  a  plateau  sloping  toward  the  south 
end,  but  the  whole  island  is  very  rugged.  Most  of  the  base  rock  is  red  lava,  and  several 
of  the  lesser  peaks  are  cinder  cones.  Recent  research  has  shown  that  the  oldest  lava 
flows  occurred  about  7,000,000  years  ago." 

Most  of  the  island  is  devoid  of  vegetation.  A  remnant  of  a  formerly  large  cypress 


314 


forest  (Cupressus  giiadahipensis)  about  Vh  miles  long,  is  located  in  the  north  central 
part  of  the  island.  The  endemic  fan  palm  {Erythea  edulis)  is  fairly  common  on  the 
north  slope  of  the  island,  and  atop  the  main  ridge  at  the  north  end  there  are  stands  of 
Island  Oaks  {Quercus  tomentella)  and  Guadalupe  Island  Pines  {Pinus  radiata  var. 
binata).  The  highest  peak.  Mount  Augusta,  rises  more  than  4200  feet  abore  the  sea. 
A  map  of  the  island,  with  names  of  major  localities,  is  given  in  Figure  1. 

History  of  Ornithological  Research 

The  Hungarian  explorer  Johan  Xantus  de  Vesey  is  generally  acknowledged  as 
having  been  the  first  naturalist  to  visit  the  island.  He  was  becalmed  there  on  1 7  March 
1859,  while  en  route  between  San  Francisco  and  Cabo  San  Lucas  (Madden  1949).  In 
his  journals  Xantus  described  several  species  of  birds  whose  identity  might  be  inferred. 
Yet,  his  accounts  of  the  island  itself  are  at  such  variance  with  those  of  an  earlier  French 
expedition  (duPetit-Thouars  1956)  that  they  undermine  the  credibility  of  his  reports, 
and  we  have  given  them  no  attention. 

It  was  the  work  of  the  botanist  Edward  Palmer  that  drew  first  attention  to  the 
island.  Palmer,  collecting  plant  specimens  for  the  U.S.  National  Museum,  arrived  in 
1875,  intending  to  remain  for  six  weeks  (Blake  1961).  Instead  he  found  himself  stranded 
for  four  months  (Table  1),  which  afforded  him  ample  time  to  collect  eight  of  the  nine 
endemic  taxa  of  landbirds,  all  of  which  were  quickly  described  by  Robert  Ridgway 
(1876). 

Palmer's  visit  came  just  in  time,  for  the  island  was  already  undergoing  a  series  of 
rapid  and  irreversible  changes.  Goats  had  been  introduced  as  a  source  of  meat  by  sealers 
or  mariners,  perhaps  as  early  as  the  18th  century,  and  more  were  introduced  for  a 
commercial  wool-producing  enterprise  in  the  ISlOs  (San  Diego  Union,  15  March  1873; 
Anon.  1874).  They  numbered  in  the  tens  of  thousands  by  the  time  of  Palmer's  visit, 
caused  the  elimination  of  much  of  the  island's  vegetation  (already  noted  by  the  French 
in  1838),  and  forced  much  of  the  birdlife  to  be  concentrated  in  the  few  wooded  areas. 

Spurred  by  Palmer's  discoveries,  Walter  E.  Bryant  spent  a  short  time  on  the  island 
in  January  1885,  then  returned  in  December  with  the  intention  of  spending  six  weeks 
there.  Three  and  a  half  months  elapsed  before  his  ship  returned.  Palmer's  work  had 


Table  1.     Chronology  of  ornithological  research  at  Guadalupe  Island,  1875-1953. 


Observers 


Jan  Feb  Mar  Apr  May  Jun  Jul  Aug  Sep  Oct  Nov  Dec 


References 


Palmer 

1875 

Bryant 

1885 

Bryant          1885 

-1886 

Anthony,  Streator 

1892 

Gaylord,  Anthony 

1896 

Anthony  et  al. 

1897 

Thobum 

1897 

Beck 

1900 

Hartert,  Rothschild     ? 

Brown  et  al. 

1906 

Townsend 

1911 

Beck 

1912 

Anonymous 

1913 

Anthony  et  al. 

1922 

Huey 

1923 

Hanna  et  al. 

1925 

Swarth 

1932 

Walker 

1938 

Vanderbilts 

1941 

Howell  and  Cade  1953 


Blake  1961 
Bryant  1887a 
Bryant  1887a 
Anthony  1901,  1925 

Gaylord  1897 

Davidson  1928,  Kaeding 

1905 
Thobum  1899 
Abbott  1933 
Howell  and  Cade  1955 
Thayer  and  Bangs  1 908 
Townsend  1911,  1923 
Abbott  1933 
Kimball  1922 

Anthony  1925,  Hanna  1925 
Huey  1924 
McLellan  1925 
Swarth  1933 
Huey  1954 

Bond  and  deSchauensee  1 944 
Howell  and  Cade  1954 


315 


PALMS 


ROCA  o 

ELEFANTE 


STEAMBOAT 

ROCK  -o 


PINES    AND    OAKS 

NORTHEAST    ANCHORAGE 

FORMER    CYPRESS    GROVE 


MT.    AUGUSTA    -    4257" 

TWIN    CANYONS 

MAIN    CYPRESS    GROVE-    3420 


RED    CINDER    CONE 


29  00N 


CAMPO  TEPEYAC 
WEST  ANCHORAGE 


ABANDONED  LOBSTER  CAMP 


ISLOTE  NEGRO 


ISLOTE  ADENTRO 
(EL  TORO) 


—  OLD  SEALING  STATION 
WEATHER  STATION 


MELPOMENE  COVE 


00  — 


ISLOTE  AFUERA 
(I.  ZAPATO) 


Figure  1.     Isla  Guadalupe,  showing  major  localities  and  forested  areas  mentioned  in  this  report. 


been  so  thorough  that  Bryant  was  unable  to  discover  any  additional  endemic  landbirds, 
but  he  did  amass  a  number  of  unusual  observations  (Bryant  1 887a).  More  importantly, 
he  discovered  the  Guadalupe  Petrel  {Oceanodroma  macrodactyla),  the  first  endemic 
seabird  to  be  recognized  (Bryant  1887/?). 

In  the  last  decade  of  the  19th  century,  several  expeditions  made  brief  stops  at 
Guadalupe.  A.  W.  Anthony  made  several  trips  and  was  the  first  to  report  the  destruction 
of  biota  by  feral  goats  (Anthony  1901).  Within  a  few  years  many  of  the  endemic  birds 
were  extinct.  The  last  report  of  a  Guadalupe  Caracara  {Polyhorus  plancus  lutosus)  was 
made  by  Rollo  Beck,  who  collected  nine  in  December  1900.  An  expedition  from  the 


316 


Table  2.     Expeditions  to  Guadalupe  Island  by  C.  L.  Hubbs. 


Year 

Jan  Feb 

Mar 

Apr 

May 

Jun 

Jul 

Aug 

Sep 

Oct  Nov  Dec 

Total  days 

1946 

■ 

2 

1950 

1 

*  8 

1954 

■ 

8 

1955 

■ 

7 

1956 

II 

5 

1957 

■ 

■ 

■       ■ 

26 

1958 

■ 

9 

1960 

1 

6 

1963 

I 

9 

1964 

I 

6 

1965 

■ 

7 

1966 

■ 

8 

1967 

■ 

9 

1969 

■ 

5 

1970 

I 

■ 

12 

1971 

■ 

6 

Thayer  Museum,  including  W.  W.  Brown,  spent  two  months  on  the  island  in  the  spring 
of  1906.  After  scouring  the  island,  they  declared  the  Guadalupe  Wren  {Thryomanes 
bewickii  brevicauda)  and  Guadalupe  Towhee  (Pipilo  erythrophthalmus  consobhnus) 
extinct  (Thayer  and  Bangs  1908),  and  predicted  the  quick  demise  of  the  Guadalupe 
nicker  (Colaptes  auratus  rufipileus).  Yet,  despite  intense  predation  by  cats,  the  Gua- 
dalupe Petrel  persisted  "in  large  numbers."  Only  six  years  later,  however,  in  1912, 
Beck  collected  two  downy  chicks,  the  last  known  examples  (Davidson  1928). 

After  the  basic  collections  had  been  made,  and  the  extinction  of  the  endemics  had 
been  documented,  interest  in  the  island  lagged.  Some  ornithologists  continued  to  visit 
it  (often  en  route  to  more  exciting  localities)  in  the  1920s  and  1930s  (e.g.,  Hanna  1925), 
but  little  new  information  resulted. 

Commencing  in  the  1940s  and  continuing  through  the  early  1970s  studies  by  Carl 
L.  Hubbs  and  his  many  colleagues  renewed  interest  in  Guadalupe.  Between  1946  and 
1971,  Hubbs  made  21  expeditions  to  the  island  (Table  2).  Hubbs  discovered  that  small 
islets  at  the  southern  end  of  the  main  island  were  important  seabird  colonies.  In  addition 
to  initiating  population  studies  on  seabirds,  he  reported  the  apparently  distinctive 
summer  and  winter  populations  of  Leach's  Storm-Petrels  (Hubbs  1960)  that  have 
received  much  subsequent  attention  (Crossin  1974,  Ainley  1980,  1983,  Bourne  and 
Jehl  1982).  He  also  discovered  the  only  known  fossil  locality  on  the  island  (Pleistocene), 
which  has  yielded  the  remains  of  a  few  seabirds  (Hubbs  and  Jehl  1976). 

In  1953,  T.  R.  Howell  and  T.  J.  Cade  spent  five  days  on  the  island,  on  one  of 
Hubbs'  expeditions.  Their  findings  (Howell  and  Cade  1954,  1955)  resulted  in  the  first 
discussion  of  the  birdlife  in  several  decades.  Additional  work  by  the  Pacific  Ocean 
Biological  Survey  Program  (POBSP)  in  the  late  1960s  and  early  1970s  (DeLong  and 
Crossin  MS,  Brownell  MS,  Crossin  MS,  1 974)  revealed  much  about  the  nesting  seabirds. 

Since  the  late  1960s  we  have  each  had  occasion  to  visit  Guadalupe  several  times. 
Our  observations,  and  those  of  many  other  biologists  who  have  generously  contributed 
data  (Table  3),  and  the  voluminous  field  notes  of  the  late  Carl  L.  Hubbs  provide  the 
basis  for  this  report.  Note,  however,  that  as  with  earlier  studies  nearly  all  of  the  recent 
work  has  been  concentrated  in  late  winter  through  early  summer  and  that  the  fall 
season,  with  one  exception,  is  unrepresented.  We  have  attempted  to  make  the  report 
comprehensive  and  current,  and  to  provide  a  complete  ornithological  bibliography. 
This  compilation  was  stimulated  in  part  by  a  request,  for  planning  purposes,  from  the 
Mexican  government,  through  Dr.  Martin  Gonzales.  Because  prospects  of  additional 
surveys  in  1982  and  1983  failed  to  materialize,  we  are  making  the  report  available 
now.  We  acknowledge,  however,  that  much  remains  to  be  learned  before  our  knowledge 
of  the  present  avifauna  is  adequate. 


317 


Table  3.     Recent  ornithological  observations  on  Isla  Guadalupe,  Mexico. 


Year 


Month 


Observers 


1965 

27-31  Jan 

1967 

26.  27,  30  Apr, 
1^  May, 
21-24  Oct 

1968 

17-22  Apr, 
20-23  Jun, 
28-29  Jun 

1969 

21-23  Feb 

1970 

19-27  Jan, 
12-18  Apr 

1971 

1 5  Mar, 
17-23  May 

1972 

mid  Dec 

1973 

22  Nov-26  Dec 

1975 

12-13  Jul 

1977 

20  Jun-13  Jul 

1978 

9-20  Feb 
16  Apr 
17-18  Dec 

1979 

24  Mar,  18  Apr 

1980 

4-5  May 

1981 

1 9-26  Aug 

1982 

12-15  Feb, 
4-6  Mar 

K.  L.  Kenyon 

R.  L.  DeLong  and  R.  S.  Crossin 

R.  L.  Brownell,  R.  S.  Crossin,  R.  L.  DeLong 

J.  R.  Jehl,  Jr. 
J.  R.  Jehl  Jr. 

J.  R.  Jehl,  Jr. 

K.  Briggs 

E.  Mirsky 

J.  R.  Jehl,  Jr. 

M.  Pierson  and  M.  Riedman 

M.  Pierson,  E.  Chu  et  al. 
W.  T.  Everett 
K.  Briggs 

W.  T.  Everett 

W.  T.  Everett 

D.  Duncan 

R.  Condit 


Annotated  List  of  Species 

In  the  following  section  we  provide  synoptic  information  on  all  species  of  birds 
recorded  on  Guadalupe  Island  and  in  waters  immediately  adjacent  (within  approxi- 
mately 1 5  km)  of  the  island.  We  include  information  on  many  species  of  seabirds, 
which  were  not  considered  by  Howell  and  Cade  (1954).  Nomenclature  follows  the  AOU 
Check-list  (1983),  except  for  Pluvialis  fulva  (see  Connors  1983).  The  following  abbre- 
viations are  used:  AMNH,  American  Museum  of  Natural  History;  CAS,  California 
Academy  of  Sciences;  HSWRI,  Hubbs-Sea  World  Research  Institute;  LACMNH,  Los 
Angeles  County  Museum  of  Natural  History;  SDNHM,  San  Diego  Natural  History 
Museum;  UCLA,  University  of  California  Los  Angeles. 

Arctic  Loon  (Gavia  arcticd) 

Uncommon  winter  visitor.  Palmer  found  an  adult  washed  ashore  on  20  May  1875 
(Ridgway  1876).  Hubbs  collected  one  on  13  February  1957  and  saw  two  others  on  his 
expedition  (8-15  February  1957),  and  Mirsky  (MS)  observed  one  on  23  November 
1973. 

Pied-billed  Grebe  {Podilymhus  podiceps) 

Accidental.  Hubbs  collected  one  on  27  October  1957;  location  of  specimen  un- 
known. 


Eared  Grebe  {Podiceps  nigricollis) 

Considered  accidental  by  Howell  and  Cade  (1955)  on  the  basis  of  a  specimen 
reported  by  Rothschild  and  Hartert  (1902).  There  are  four  subsequent  reports.  Hubbs 


318 


Figure  2.     Guadalupe  Storm-Petrel  {Oceanodroma  macrodactyla).  Photograph  of  a  mount  in  the  Field 
Museum  of  Natural  History,  Chicago. 


collected  two  on  12  February  1957  (LACMNH)  and  reported  taking  another  on  30 
October  1957;  Mirsky  (MS)  reported  one  on  23-25  November  1973. 

Western  Grebe  (Aechmophorus  occidentalis) 

Accidental.  Two  reports  by  Hubbs  of  single  birds  on  12  November  1954  and  17 
December  1957. 


Short-tailed  Albatross  {Diomedea  albatrus) 

Probably  regular  in  the  area  until  the  late  19th  century.  Bryant  (1889)  reported 
five  between  Ensenada  and  Guadalupe  in  April  1886.  Now  very  rare  and  not  reported 
in  Baja  California  for  many  decades. 

Black-footed  Albatross  {Diomedea  nighpes) 

Uncommon  but  regular  near  the  island  through  most  of  the  year,  with  many  sight 
records  from  September  to  June. 

Laysan  Albatross  {Diomedea  immutabilis) 

This  species  is  a  rare  but  regular  wanderer  to  Baja  California.  R.  Wisner  observed 
a  single  bird  about  3  km  off  the  north  end  of  the  island  on  22  April  1958,  and  R. 
Pitman  saw  one  90  km  to  the  northeast  on  4  January  1980.  Anthony  (1898Z))  took  a 
specimen  between  San  Geronimo  and  Guadalupe  islands  in  March  1897. 


319 


Northern  Fulmar  {Fulmams  glacialis) 

Irregular  visitor  in  winter.  Mirsky  (MS)  reported  one  on  6-7  December  1973.  This 
species  reaches  the  latitude  of  central  Baja  California  in  flight  years. 

Cook's  Petrel  {Pterodroma  cookii) 

This  species  occurs  regularly  off' the  coast  of  Baja  California  (Jehl,  pers.  obs.).  Two 
were  seen  on  1 1  April  1961,  100  km  north  of  the  island  (Small  1961).  Pitman  saw  one 
along  with  another  unidentified  Pterodroma  within  15  km  of  Guadalupe  on  1 1  October 
1979. 

Pink-footed  Shearwater  (Puffinus  creatopus) 

According  to  Grinnell  (1928):  "First  definitely  recorded  by  Gaylord  ...  as  seen  by 
A.  W.  Anthony  near  Guadalupe  Island,  September  17,  1896."  While  this  record  may 
be  correct— the  species  is  a  common  migrant  nearer  shore— we  suspect  that  it  more 
likely  pertains  to  the  Black- vented  Shearwater,  which  breeds  locally. 

Sooty  Shearwater  {Puffinus  griseus) 

Probably  a  regular  visitor  from  April  to  October.  Thayer  and  Bangs  ( 1 908)  reported 
taking  two  specimens  "near"  the  island  in  June  1906,  and  Pitman  saw  one  on  4  January 
1979. 

Black-vented  Shearwater  (Puffinus  opisthomelas) 

This  shearwater  has  long  been  known  to  breed  at  Guadalupe.  Anthony  (1900) 
reported  it  to  be  "rather  common  ...  in  several  parts  of  the  island,  but  in  no  place 
was  there  any  large  colony.  Their  nests  were  all  inaccessible  owing  to  the  nature  of  the 
sites  selected,  whether  in  natural  holes  in  the  lava  or  under  large  boulders,  and  no  eggs 
were  secured.  A  night  was  spent  on  the  top  of  the  island  in  a  heavy  cypress  growth, 
about  4000  feet  above  the  sea.  Here  the  shearwaters  were  heard  all  night,  their  choking, 
gasping  notes  coming  from  all  sides  as  they  flew  through  this  grove." 

Hubbs  discovered  and  for  many  years  surveyed  the  colony  on  Islote  Negro,  which 
in  the  late  1960s  and  early  1970s  contained  100-150  pairs  (Jehl,  pers.  obs.,  Crossin 
MS,  Brownell  MS).  Crossin  visited  Islote  Afuera  on  22-23  June  1968  and  found  a 
colony  of  150+  pairs.  We  suspect  that  there  may  be  other  colonies  on  the  main  island 
because  in  the  early  evening  flocks  of  500-2500  shearwaters,  too  many  to  be  accom- 
modated by  known  nesting  areas,  stage  at  the  south  end  of  the  island. 

The  breeding  season  at  Islote  Negro  can  be  outlined  with  some  confidence.  By  late 
November  single  birds  begin  to  occupy  burrows.  By  early  January  pairs  are  commonly 
found  and  fresh  eggs  are  present  by  5  March  (Hubbs  field  notes).  The  peak  of  the  egg- 
laying  season  occurs  in  early  April,  when  most  burrows  contain  an  egg,  but  may  extend 
to  late  June  (Jehl,  pers.  obs.;  Crossin  MS).  Young  are  present  as  early  as  late  April  and 
in  late  June  most  burrows  have  a  chick.  There  are  no  data  for  later  in  the  fall.  Crossin 
reported  that  the  phenology  of  the  Islote  Afuera  colony  was  somewhat  later  than  at 
Islote  Negro. 

Leach's  Storm-Petrel  {Oceanodroma  leucorhod) 

There  has  been  much  uncertainty  about  the  historical  status  of  storm-petrels  at 
Guadalupe,  much  of  which  may  never  be  fully  resolved  because  the  early  literature  is 
inconsistent,  and  because  cats  have  eliminated  most,  if  not  all,  storm-petrel  colonies 
on  the  main  island,  whose  location  and  species  composition  cannot  be  fully  recon- 
structed. The  complicated  case  of  the  Leach's  Storm-Petrel  is  best  treated  chronolog- 
ically. 

In  the  winter  and  spring  of  1885-86,  W.  E.  Bryant  was  stranded  on  Guadalupe 
for  nearly  four  months.  Near  the  end  of  his  stay  he  discovered  storm-petrels  nesting 


320 


among  the  pines  and  cypress  trees  at  the  northern  end  of  the  island.  Bryant  (1887a) 
reported  these  as  Leach's  Storm-Petrels  {Oceanodroma  leucorhoa),  and  recognized  that 
they  had  a  more  deeply  forked  tail  and  a  longer  middle  toe  than  other  races.  Bryant 
(1887^)  then  named  these  as  a  new  race  {O.  I.  macrodactyla),  the  Guadalupe  Storm- 
Petrel,  which  the  AOU  (1889)  elevated  to  species  rank. 

In  1889,  C.  H.  Townsend  secured  a  single  small  storm-petrel,  which  had  the  sides 
of  the  rump  "whitish,"  at  sea  near  Socorro  Island,  1200  km  SE  of  Guadalupe.  He 
described  it  as  a  new  species  {O.  socorwensis),  guessing  that  it  nested  at  Socorro  Island 
(Townsend  1890). 

A.  W.  Anthony  visited  Guadalupe  in  May  1892  and  collected  three  nestling  Gua- 
dalupe Storm-Petrels,  but  did  not  mention  other  species.  In  1896,  Anthony  and  H.  A. 
Gaylord  visited  Guadalupe  in  mid-September.  Anthony  spent  a  night  atop  the  island 
but  reported  no  petrels  of  any  species  (Anthony  1898c).  However,  Gaylord  (1897) 
stated:  "Regarding  the  Petrels  which  breed  on  the  island,  the  hunters  told  us  that 
while  doing  some  stone  work  in  the  region  of  the  Petrel  colony,  they  had  found  two 
different  species.  They  described  the  Guadalupe  Petrel  and  an  entirely  black  one,  which 
together  with  a  wing  found  on  the  trail  to  the  cypress  grove  makes  it  appear  that  O. 
homochroa  is  an  inhabitant  of  the  island." 

In  late  March,  Anthony  collected  a  series  of  adult  macrodactyla,  along  with  a  few 
eggs.  That  summer  he  returned  and  collected  young,  noting  that  macrodactyla  leaves 
the  colony  by  10  June.  On  neither  trip  did  he  find  other  species  in  the  colony,  but  in 
June  he  collected  a  storm-petrel  with  enlarged  ova  at  sea  120  mi  north  of  Guadalupe, 
which  became  the  type  of  Kaeding's  Petrel  {Oceanodroma  kaedingi).  This  new  form 
was  distinguished  by  its  small  size  and  white  rump,  but  Anthony  (1898a)  noted  that 
there  was  much  variation  in  rump  color.  From  observations  at  sea,  Anthony  (1898a) 
surmised  that  the  range  of  kaedingi  extended  from  the  Revillagigedos  to  southern 
California  and  guessed  (1898c)  that  its  breeding  grounds  were  on  Guadalupe.  Also,  he 
reidentified  the  wing  ascribed  to  homochroa  in  1896  as  belonging  to  kaedingi  (Anthony 
1898c). 

H.  B.  Kaeding  (1905),  who  accompanied  the  1897  expedition,  was  the  first  to  allege 
a  common  breeding  area  for  kaedingi  and  macrodactyla.  He  wrote  that  "the  breeding 
grounds  of  kaedingi  are  as  yet  unknown,  but  it  is  probable  that  the  birds  occupy  the 
burrows  of  Guadalupe  Petrels  .  .  .  after  the  breeding  season  of  the  former  is  closed." 
In  late  May  1906,  W.  W.  Brown  and  H.  W.  Marsden  visited  the  breeding  grounds  of 
the  Guadalupe  Storm-Petrel  (Thayer  and  Bangs  1908);  they  found  O.  macrodactyla 
but  no  other  species. 

By  this  time  most  of  the  islands  off  Baja  California  had  been  surveyed  and  tax- 
onomists  began  to  re-evaluate  the  variation  in  the  Leach's  Storm-Petrel  group  (including 
Swinhoe's  Storm-Petrel,  O.  monorhis,  of  the  western  Pacific),  a  process  that  still  con- 
tinues (e.g.,  von  Berlepsch  1906,  Emerson  1906,  Godman  1907-1910,  Obserholser 
1917,Loomis  1918,  van  Rossem  1942,  Austin  1952,  Todd  1955,  Palmer  1962,  Crossin 
1974,  Ainley  1980,  1983,  Bourne  and  Jehl  1982,  Power  and  Ainley  MS).  The  work 
was  made  difficult  because  of  the  petrels'  discontinuous  breeding  range,  the  scarcity  of 
specimens  from  breeding  colonies,  the  great  variability  in  some  populations,  and  the 
lack  of  knowledge  about  the  location  of  breeding  grounds.  The  last  point  was  illustrated 
by  A.  C.  Bent  (1922),  who  reported  that  "Mr.  Anthony  wrote  me  that  on  Guadalupe 
Island  the  Guadalupe  Petrels  breed  early,  April  20  or  earlier,  and  that  after  they  are 
through  breeding  the  Kaeding  Petrels  use  the  same  burrows."  As  Anthony  was  the  most 
knowledgeable  ornithologist  regarding  the  petrels  of  the  region  no  one  questioned  his 
perpetuation  of  Kaeding's  (1905)  idea,  even  though  all  expeditions  to  the  nesting  grounds 
of  macrodactyla  failed  to  report  any  other  species  as  nesting,  and  despite  the  fact  that 
the  nesting  grounds  of  kaedingi  had  yet  to  be  discovered  (e.g.,  Oberholser  1917,  Grinnell 
1918). 

On  11-17  July  1 922  Anthony  (1925)  returned  to  Guadalupe  and  visited  the  nesting 
area  of  macrodactyla.  That  species  had  disappeared,  but  the  expedition  did  find  nu- 
merous bodies  of  kaedingi  that  had  been  killed  by  cats  or  were  impaled  on  cholla,  and 


321 


even  found  a  week-old  chick  (CAS  no.  25561).  Unfortunately,  no  data  were  published 
on  precise  localities.  Hanna  (1925)  stated  that  kaedingi  "lived  among  the  loose  rocks 
and  in  holes  in  the  cliffs,"  whereas  Anthony  (1925)  said  that  it  "evidently"  nested  in 
high  cliffs  at  the  north  end  of  the  island.  Regardless,  the  discovery  of  the  chick  (i) 
provided  the  first  proof  that  a  species  other  than  macrodactyla  nested  on  the  main 
island,  (ii)  established  the  island  as  a  breeding  location  for  kaedingi,  (iii)  is  in  accord 
with  recent  data  that  kaedingi  is  a  crevice  nester  {macrodactyla  nested  in  burrows  in 
forested  areas),  and  (iv)  shows  that  Anthony  had  no  evidence  of  burrow-sharing  between 
macrodactyla  and  kaedingi. 

In  1950,  Carl  L.  Hubbs  discovered  and  began  to  study  the  Leach's  Storm-Petrels 
nesting  on  small  islets  at  the  south  end  of  the  main  island.  Hubbs  (1960)  was  the  first 
to  recognize  that  there  were  evidently  two  populations,  a  larger  form  with  white  rump 
that  bred  in  winter  and  fiedged  young  by  April,  and  a  smaller  form  with  less  white  on 
the  rump  that  began  laying  in  June  and  fledged  young  in  October  or  November.  Surveys 
by  the  POBSP  in  June  1968  determined  the  summer  population  on  Islote  Negro  at 
4000  adults  and  on  Islote  Afuera  at  3000  adults  (Crossin  MS,  1974).  Jehl  also  found 
a  nearly-grown  chick  on  Gargoyle  Rock  in  April  1970  and  Huey  (1952)  reported  a 
chick  there  in  January  1950.  Huey  also  reported  that  Hendrickson  collected  "a  number 
of  Oceanodroma  petrels  from  rock  crevices  on  the  hillsides  of  Melpomene  Cove,  sit- 
uated on  the  southern  end  of  Guadalupe  Island."  We  presume  that  small  colonies 
remain  to  be  discovered  on  the  main  island  but  none  are  known  with  certainty.  Evidence 
of  the  persistence  of  one  or  more  colonies  there  is  provided  by  repeated  observations 
of  birds  landing  on  ships  at  the  north  end  of  Guadalupe  in  late  spring  and  summer 
(Jehl,  pers.  obs.,  Anthony  1925,  Huey  1930),  as  well  as  records  of  a  few  petrels  flying 
over  the  north  end  of  the  island  in  May  1971  (Jehl  1972)  and  many  near  the  cypress 
grove  in  August  1981  (R.  Moran,  D.  Duncan,  pers.  comm.). 

The  correct  nomenclature  of  the  Guadalupe  populations  has  been  debated.  Van 
Rossem  ( 1 942)  pointed  out  that  the  type  of  socorroensis  was  a  small  bird  of  the  Gua- 
dalupe population;  since  that  name  had  priority,  kaedingi  was  relegated  to  synonymy. 
When  Hubbs  (1960)  found  that  two  populations  were  present  on  Islote  Negro,  he 
referred  to  the  summer  breeders  as  socorroensis  and  the  winter  breeders  as  kaedingi. 
However,  the  situation  is  evidently  even  more  complicated,  because  the  summer  birds 
are  highly  variable.  Crossin  (1974)  reported  that  all  birds  nesting  at  Islote  Negro  were 
dark-rumped,  while  on  Islote  Afuera  over  90%  of  the  population  was  white-rumped. 
Recently,  Ainley  (1980)  named  the  winter  breeders  as  a  new  race,  cheimomnestes,  on 
grounds  unconvincing  to  Bourne  and  Jehl  (1982)  but  reaffirmed  by  Power  and  Ainlev 
(MS). 

In  summary.  Leach's  Storm-Petrels  breed  commonly  on  islets  at  the  southern  end 
of  Guadalupe  Island.  There  is  also  strong  presumptive  evidence  for  the  persistence  or 
re-establishment  of  colonies  near  the  north  end  and  center  of  the  main  island.  According 
to  Power  and  Ainley  (MS),  the  winter  and  summer  populations  on  Islote  Negro  are 
temporally  segregated  and  morphologically  distinct.  If  so,  their  findings  would  require 
that  the  two  populations  be  considered  as  distinct  species,  not  subspecies.  Yet,  because 
of  the  lack  of  field  work  in  the  fall,  neither  Bourne  and  Jehl  (1982)  nor  Crossin  (1974: 
1 74)  were  convinced  that  the  populations  are  fully  isolated  temporally.  Furthermore, 
because  there  are  differences  in  coloration  (at  least)  between  summer  populations  breed- 
ing contemporaneously  on  I.  Negro  and  1.  Afuera.  and  because  the  morphological 
characters  and  breeding  seasons  are  not  known  for  the  presumed  colonies  on  the  main 
island,  the  situation  remains  unclear.  This  is  one  of  the  most  complicated  cases  of 
differentiation  known  in  birds,  and  one  that  will  require  additional  study. 

Ashy  Storm-Petrel  {Oceanodroma  homochroa) 

A  wing  attributed  to  this  species  by  Gaylord  (1897)  was  reidentified  as  that  of  O. 
leucorhoa  by  Anthony  (1898r).  In  Gaylord's  original  report  (see  above)  the  presence 
of  an  "entirely  black"  storm-petrel  near  the  colony  of  O.  macrodactyla  on  the  main 


322 


island  was  alleged  by  "hunters."  Probably  these  represented  dark-rumped  examples  of 
the  summer  population  of  6>.  /.  socorroensis  or,  less  likely,  wanderers  of  O.  /.  chapmani 
from  the  San  Benito  Islands.  O.  homochroa  may  occur  off  Baja  California  in  winter, 
but  the  nearest  breeding  colony  is  on  Los  Coronados,  320  km  to  the  north,  and  may 
consist  of  only  2-3  pairs  (Jehl  pers.  obs.).  , 

Wedge-rumped  Storm-Petrel  {Oceanodroma  tethys) 

A  specimen  obtained  in  1950  was  said  to  have  been  collected  from  a  crevice  on 
the  side  of  Melpomene  Cove,  at  the  south  end  of  the  main  island,  which  also  included 
a  downy  young  of  6>.  leucorhoa  (Huey  1952).  However,  the  precise  locality  is  not  clear. 
Howell  and  Cade  (1954)  claimed  it  was  collected  on  Gargoyle  Rock  at  the  end  of  the 
island,  and  Hubbs  told  Jehl  the  same  thing.  Jehl  landed  there  in  April  1970  and  found 
a  single  chick  of  O.  leucorhoa.  There  are  no  subsequent  records  although  the  species 
occurs  regularly  to  southern  Baja  California  (Jehl  pers.  obs..  Pitman  pers.  comm.). 

Black  Storm-Petrel  {Oceanodroma  melania) 

Kaeding  (1905)  reported  this  species  near  Guadalupe,  but  the  observation  could 
have  pertained  to  any  of  several  others.  Anthony's  (1898c)  report,  which  implies  that 
the  species  nests  on  Guadalupe  Island,  actually  alludes  to  the  nesting  season  of  the 
species  elsewhere. 

Guadalupe  Storm-Petrel  {Oceanodroma  macrodactyla) 

This  endemic  and  unusual  storm-petrel  (Fig.  2),  originally  described  as  a  race  of 
O.  leucorhoa  (Bryant  1887^),  bred  in  soil  burrows  atop  the  main  island,  among  the 
pines  at  the  north  end  and  in  the  cypress  grove.  Its  status  and  history  have  most  recently 
been  reviewed  by  Jehl  (1972).  Both  Howell  and  Cade  (1954)  and  Greenway  (1967) 
incorrectly  cite  the  last  report  of  the  species,  apparently  overlooking  the  work  of  Da- 
vidson (1928),  who  corrected  earlier  errors  and  established  August  1912  as  the  last 
acceptable  record.  A  winter  breeder,  it  laid  by  early  March  (Kaeding  1905)  but  some- 
times as  late  as  June,  for  Beck  collected  three  chicks  on  3  August  1912  (specimens  in 
AMNH).  Nothing  is  known  about  the  ecology  or  distribution  of  this  species  at  sea. 
Thobum  (1899)  reported  it  as  abundant  about  his  ship  at  night  in  late  June  1897,  as 
it  lay  anchored  at  Guadalupe,  and  reported  collecting  several.  If  so,  these  would  rep- 
resent the  only  documented  records  of  macrodactyla  away  from  the  breeding  colonies, 
but  we  have  been  unable  to  locate  any  specimens  and  suspect  that  his  reports  refer  to 
O.  leucorhoa. 

The  Guadalupe  Storm-Petrel  was  considered  abundant  in  the  colony  as  late  as 
1906  (Thayer  and  Bangs  1908),  but  was  being  preyed  upon  heavily  by  domestic  cats. 
Davidson  (1928),  in  declaring  it  to  be  extinct,  based  her  conclusion  on  the  negative 
results  of  the  1922  expedition  by  the  Cahfomia  Academy  of  Sciences  (Hanna  1925), 
which  took  place  in  mid-summer,  after  the  main  breeding  season,  and  on  the  erroneous 
assumption  that  the  CAS  expedition  of  April  1925  (McLellan  1926)  had  carefully 
searched  the  breeding  grounds.  Jehl  (1972)  spent  several  nights  listening  for  petrels  in 
the  early  1970's  in  the  pine-oak  woodlands  atop  the  island,  but  did  not  visit  the  cypress 
grove,  where  the  species  also  had  nested.  No  thorough  survey  of  the  breeding  grounds 
has  been  made  at  the  appropriate  season  since  1906.  The  apparent  persistence  of  <9. 
leucorhoa  on  the  main  island  despite  predation  by  cats  allows  some  hope  that  macro- 
dactyla may  still  exist. 

As  noted  above  there  is  no  acceptable  evidence  for  the  often-repeated  contention 
that  Guadalupe  and  Leach's  storm-petrels  ever  bred  in  the  same  colonies.  All  evidence 
suggests  that  they  used  different  habitats:  macrodactyla  burrowed  in  soil,  leucorhoa 
nested  in  crevices.  Further,  the  breeding  season  of  macrodactyla  probably  overlapped 
that  of  the  winter  and  summer  forms  of  leucorhoa,  so  that  sequential  use  of  burrows 
would  have  been  impossible. 


323 


Red-billed  Tropicbird  {Phaethon  aethereus) 

Probably  regular  in  the  area  but  the  only  records  are  of  individual  birds  25  and 
37  km  north  of  the  island  on  1 1  October  1979,  and  another  slightly  farther  north  on 
7  January  1980  (Pitman). 

Red-tailed  Tropicbird  {Phaethon  rubricauda) 

Anthony  (1898Z^)  reported  collecting  a  specimen  close  to  Guadalupe  on  23  July 
1897. 

Brown  Pelican  {Pelecanus  occidentalis) 

There  are  two  records,  an  immature  individual  reported  by  Anthony  (1925),  and 
a  probable  juvenile  reported  on  4  July  1977  (Pierson  and  Riedman,  MS).  This  coastal 
species  rarely  wanders  to  deep  waters  beyond  the  continental  shelf. 

Double-crested  Cormorant  {Phalacwcorax  auritus) 

Probably  a  rare  or  accidental  visitor,  but  its  status  requires  verification.  One  was 
reported  by  Gaylord  (1897)  and  Huey  (1924)  casually  mentions  the  species  as  being 
present  in  1923.  Hubbs  reported  the  species  on  several  trips  and  on  11  June  1955 
described  a  cormorant  with  "bill  and  pouch  yellow"  that  would  seem  to  be  this  species. 

Brandt's  Cormorant  (Phalacwcorax  penicillatus) 

Resident.  Seen  regularly  in  small  numbers  along  the  entire  east  side  of  the  island 
but  commonest  near  the  southern  end,  where  a  few  pairs  breed  on  outer  islet  (McLellan 
1926,  Crossin  MS),  and  on  Islote  Zapato  (Hubbs  notes,  Jehl  pers.  obs.).  The  maximum 
single  count  is  20  at  I.  Zapato  on  4  May  1966.  We  suspect  that  the  entire  island 
population  does  not  exceed  30-40  individuals.  Specimens  in  SDNHM. 

Pelagic  Cormorant  {Phalacwcorax  pelagicus) 

J.  Sefton  reported  this  species  on  Hubbs'  expedition  of  27  January-3  February 
1950  (Hubbs  field  notes).  His  identifications  were  doubted  by  Hubbs,  and  by  us. 

Magnificent  Frigatebird  {Fregata  magnificens) 

Sightings  of  an  immature  on  1  and  4  July  1973  (Pierson  and  Riedman,  MS) 
probably  represent  the  same  individual. 

Great  Blue  Heron  {Ardea  herodias) 

Probably  a  rare  but  regular  winter  visitor.  Hubbs  saw  one  or  more  on  five  different 
trips  (maximum  three,  two  trips),  between  November  and  February.  Other  records  are: 
1,  September  1896  (Gaylord  1897);  2-3  in  summer  1922  (Anthony  1925);  and  1, 
midwinter  1965  (Kenyon  MS). 

White-fronted  Goose  {A  nser  albifrons) 

Bryant  (1887(3)  shot  one  on  14  January  1885,  but  it  fell  over  a  cliff' and  could  not 
be  recovered. 

Brant  {Branta  nigricans) 

Mirsky  (MS)  reported  a  sick  bird  at  Northeast  Anchorage  on  22-24  November 
1973. 


324 


Mallard  {Anas  platyrhynchos) 

Hubbs  saw  several  and  collected  single  males  on   13  and   17  December  1957 
(LACMNH). 

Northern  Pintail  {Anas  acuta) 

Pitman  saw  one  40  km  north  of  the  island  on  1 1  October  1979. 

Blue- winged  Teal  {Anas  discors) 

Hubbs  reported  collecting  an  adult  male  on  30  October  1957.  The  location  of  the 
specimen,  if  preserved,  is  unknown. 

Cinnamon  Teal  {Anas  cyanoptera) 

Jehl  saw  a  male  swimming  along  the  shore  of  the  main  island  on  21  January  1970. 

Lesser  Scaup  {Aythya  affinis) 

Hubbs'  field  notes  list  "a  female  or  immature  male"  several  kilometers  from  the 
island  on  22  November  1964. 

Red-breasted  Merganser  {Mergus  senator) 

This  species  probably  is  an  occasional  winter  visitor.  There  are  two  records:  28 
January  1950  (Hubbs),  and  13-17  December  1973  (Mirsky  MS). 

Osprey  {Pandion  haliaetus) 

Status  uncertain.  Ospreys  have  nested  on  many  islands  along  the  Baja  California 
peninsula  and  perhaps  formerly  bred  on  Guadalupe,  although  proof  is  lacking.  Speci- 
mens were  collected  1 1  July  1922  (Anthony  1925,  Hanna  1925)  and  on  25  July  1941 
(Bond  and  Meyer  de  Schauensee  1944).  Kenyon  (MS)  visited  Guadalupe  early  in  1965 
and  saw  no  Ospreys  but  reported  two  presumed  nests  near  the  north  end  of  the  island. 
There  are  no  other  reports  or  indications  of  the  species'  presence. 

Red-tailed  Hawk  {Buteo  jamaicensis) 

Formerly  resident  in  small  numbers.  Howell  and  Cade  (1954)  considered  it  "ap- 
parently resident  until  at  least  1932,"  but  none  of  the  early  explorers  were  able  to  find 
any  nests.  Palmer  {in  Bryant  1887a)  considered  it  as  common  as  the  caracara.  Thayer 
and  Bangs  (1908)  and  Anthony  (1925)  reported  that  three  or  four  could  be  seen  in  a 
day;  Hanna  (1925)  also  considered  it  common.  This  hawk  wanders  to  many  offshore 
islands  in  fall  migration  and  probably  reaches  Guadalupe  infrequently.  However,  we 
know  of  no  recent  reports  for  any  season,  and  it  is  not  resident  at  this  time. 

Crested  Caracara  {Polyborus  plancus  lutosus) 

Extinct;  formerly  resident  in  small  numbers.  The  detailed  history  of  this  endemic 
form  and  its  taxonomy  have  been  reviewed  by  Abbott  (1933)  and  Brown  and  Amadon 
(1968). 

American  Kestrel  {Falco  sparverius) 

Resident  in  small  numbers.  Bryant  (1887a)  stated  that  they  were  found  most  often 
in  the  central  and  higher  portions  of  the  islands.  Howell  and  Cade  (1954)  reported  a 
pair  with  young  on  a  cliff' overlooking  the  sea  at  Northeast  Anchorage;  other  birds  were 
in  the  area.  In  recent  years  the  species  has  been  seen  regularly  near  Northeast  Anchorage 
and  at  the  southern  end  of  the  island.  D.  A.  Duncan  {pers.  comm.)  visited  Guadalupe 
19-26  August  1981  and  reported  it  as  common  everywhere,  one  or  two  being  seen  at 


325 


most  localities.  The  Guadalupe  population  was  described  as  an  endemic  race  (gua- 
daliipensis)  by  Bond  (1943);  its  validity  was  accepted  by  the  AOU  Check-list  (1957) 
though  not  by  the  Mexican  Check-list  (Friedmann,  Griscom,  and  Moore  1950). 

Peregrine  Falcon  {Falco  peregrinus) 

This  large,  maritime  falcon  is  likely  to  have  occurred  regularly  during  migration, 
but  we  know  of  only  one  report,  a  single  bird  seen  on  19  September  1896  (Gaylord 
1897). 

Prairie  Falcon  {Falco  mexicanus) 

Br>'ant  (1889)  reported  that  the  species  was  seen  on  "two  or  three  occasions"  in 
1886,  but  we  suspect  that  these  sightings  pertain  to  the  Peregrine  Falcon. 

Pacific  Golden  Plover  (Pluvialis  fulva) 

Jehl  saw  a  flock  of  20,  two  km  north  of  the  settlement  at  the  south  end  on  22 
February  1969.  Mirsky  (MS)  reported  from  1  to  12  birds  along  the  shore  at  Northeast 
Anchorage  from  23  November-16  December  1973.  Presumably  all  records  of  golden 
plovers  pertain  to  this  species  {see  Connors  1983). 

Killdeer  {Charadrius  vociferus) 

Hubbs  reported  two  on  13  December  1957. 

Willet  {Catoptrophorus  semipalmatiis) 

The  only  report  is  a  single  bird  observed  between  10-14  February  1977  (E.  Chu). 

Wandering  Tattler  {Heteroscelus  incanus) 

This  is  a  regular  visitor  to  the  island  from  fall  through  spring;  there  is  one  summer 
record.  One  or  two,  often  more,  are  seen  on  most  trips. 

Ruddy  Turnstone  {Arenaria  interpres) 

Though  not  reported  by  Howell  and  Cade  (1954),  this  species  is  a  regular  visitor 
in  small  numbers.  There  are  specific  records  for  June,  November-January,  and  April. 
At  least  three  were  present  in  November  1964  (Hubbs).  Hubbs  also  collected  several 
specimens,  the  location  of  which  is  not  known. 

Black  Turnstone  {Arenaria  melanocephala) 

Uncommon  but  regular  in  migration  and  during  the  winter.  There  are  records  for 
October-February,  and  April.  The  maximum  count  is  seven  on  20-26  January  1970 
(Hubbs,  Jehl).  Specimen  LACMNH. 

Sanderling  {Calidris  alba) 

Two  were  seen  on  22  January  1970  (Jehl). 

Western  Sandpiper  {Calidris  mauri) 

One  was  photographed  at  Northeast  Anchorage  on  16  April  1978  (Everett). 

Short-billed  Dowitcher  {Limnodromus  griseus) 

Hubbs  collected  an  immature  that  landed  on  his  boat  about  2  km  off  the  south 
end  of  the  island  on  29  August  1956  (LACMNH). 


326 


Common  Snipe  {Gallinago  gallinagd) 

One  record,  atop  the  northern  end  of  the  island  on  8  June  1953  (Howell  and  Cade 
1954). 

Red  Phalarope  {Phalaropus  fulicaria) 

Regular  in  migration.  This  phalarope  is  seen  irregularly,  sometimes  in  fair  numbers, 
between  November  and  May;  it  has  also  been  reported  in  late  June  (Thayer  and  Bangs 
1908). 

Jaegers  {Stercorarius  spp.) 

Jaegers  certainly  occur  near  the  island  during  migration,  but  the  only  published 
record  seems  to  be  that  of  Gaylord  (1897),  who  reported  two  Long-tailed  Jaegers  {S. 
longicaudus)  on  17  September  1896.  Pitman  has  seen  several  jaegers  in  the  area  in 
January,  and  identified  a  Pomarine  {S.  pomahnus)  on  4  January  1980. 

Heermann's  Gull  {Larus  heermanni) 

Two  adults  were  photographed  by  S.  Leatherwood  in  January  1 973  (photo  HSWRI). 

Ring-billed  Gull  {Larus  delawarensis) 

Jehl  and  R.  DeLong  saw  one  immature  at  Northeast  Anchorage  on  22  January 
1970.  This  species  rarely  ventures  beyond  the  coastal  beaches.  Hubbs  reported  "a  few" 
on  28  January  1950,  but  his  identification  seems  questionable. 

California  Gull  {Larus  californicus) 

Though  not  recorded  by  Howell  and  Cade  (1954),  this  gull  is  a  regular,  sometimes 
common,  winter  visitor.  It  avoids  the  elephant  seal  beaches,  because  of  competition 
with  the  larger  gulls,  and  tends  to  occur  at  sea.  Twenty  in  February  1978  (Chu  et  al. 
MS)  is  the  largest  number  recorded  (but  see  Herring  Gull). 

Herring  Gull  {Larus  argentatus) 

Common  winter  visitor  from  November-April,  at  times  being  as  common  as  the 
Western  Gull.  Hubbs  reported  that  it  was  by  far  the  commonest  gull  in  January- 
February  1950,  and  counted  360  at  the  south  end  of  the  island  in  late  January  1960. 
However,  we  suspect  that  many  of  these  were  California  Gulls,  for  at  that  season  Herring 
Gulls  congregate  near  the  elephant  seal  rookeries. 

Thayer's  Gull  {Larus  thayeri) 

Uncommon  but  regular  winter  visitor.  There  are  several  records,  all  for  immature 
or  sub-adult  birds;  21  February  1969,  16  April  1970,  30  January  1971  (3)  and  15 
March  1971  (Devillers  et  al.  1971). 

Western  Gull  {Larus  occidentalis) 

This  species  is  resident  at  Guadalupe.  Hubbs  recognized  that  the  local  population 
differed  slightly  from  the  mainland  birds.  There  are  minor  differences  in  the  color  of 
the  fleshy  parts  (Howell  and  Cade  1 954)  and  also  in  the  pattern  of  the  primary  markings. 
Hubbs  (1960)  suggested  that  it  might  represent  an  endemic  race,  but  no  formal  analysis 
of  the  variation  has  been  attempted.  In  winter  the  local  population  is  probably  enhanced 
by  representatives  from  the  mainland,  as  both  Hubbs  and  Jehl  have  seen  many  birds 
with  pinkish  (rather  than  whitish)  legs  at  that  season. 

In  Jehl's  opinion  the  population  in  1969-71  consisted  of  only  30-40  pairs.  Crossin 
(MS)  reported  that  the  species  is  "rather  sparse,"  and  guessed  that  the  local  population 


327 


in  June  was  no  larger  than  200  birds.  Jehl  found  a  nest  with  three  eggs  near  the  old 
Lobster  Camp  on  21  May  1971,  and  Crossin  reported  another  on  Islote  Afuera  on  20- 
23  June  1968.  In  contrast  to  mainland  gulls,  the  Guadalupe  birds  nest  singly,  well  back 
from  the  shore,  and  there  is  no  evidence  of  colonies.  In  November-December  1973, 
Mirsky  reported  100  at  the  Northeast  Anchorage;  all  had  whitish  legs.  Chu  et  al.  (MS) 
counted  100-125  along  the  entire  eastern  shore  of  the  island  in  February  1978,  and 
noted  that  adults  outnumbered  juveniles  by  about  10:1.  Pierson  and  Riedman  (MS) 
reported  at  least  100  birds  during  a  circumnavigation  in  the  first  week  of  July  1977, 
most  of  which  were  attending  nests;  at  least  15  large  nestlings  were  seen. 

Bryant  (1887a)  was  told  that  gulls  nested  commonly  at  the  southern  end  of  the 
island,  ''where  they  were  not  so  frequently  molested  by  the  'Quelelis'"  (=Caracaras). 

Glaucous-winged  Gull  {Larus  glaucescens) 

Regular  winter  visitor,  most  frequently  reported  at  the  Northeast  Anchorage  in 
January-March,  when  they  and  other  gulls  feed  on  elephant  seal  remains  and  placentas. 
Up  to  25,  adults  and  immatures,  have  been  seen  at  that  time  (Kenyon,  MS).  There  are 
records  from  November-May,  the  latest  being  1  May  1967  (Hubbs). 

Black-legged  Kittiwake  (Rissa  tridactyla) 

The  Kittiwake  occurs  in  winter;  it  is  common  in  some  years,  absent  in  others. 
Flocks  of  up  to  100  were  seen  around  the  island  on  20-26  January  1970  by  Hubbs  and 
Jehl. 

Sabine's  Gull  {Xema  sabini) 

This  migrant  is  probably  uncommon  but  regular  in  spring  and  fall.  Hubbs  reported 
10  birds  5  km  east  of  the  island  on  26  April  1967.  An  additional  report,  on  27  January 
1950  (Hubbs)  almost  certainly  pertains  to  an  immature  kittiwake. 

Royal  Tern  {Sterna  maxima) 

Gaylord  (1897)  reported  one  near  the  island  on  17  September  1886. 

Arctic  Tern  {Sterna  paradisaea) 

This  species  certainly  occurs  regularly  off  the  coast  of  Baja  California,  but  there 
are  few  records.  Pitman  identified  one  near  the  island  on  1 1  October  1979  and  saw  a 
second  tern,  probably  of  the  same  species. 

Xantus  Murrelet  {Synthliboramphus  hypoleuca) 

This  small  alcid  breeds  on  at  least  two  of  the  small  islets  at  the  southern  end  of 
the  main  island;  the  nesting  grounds  were  discovered  by  Hubbs.  Crossin  (MS)  estimated 
the  Islote  Negro  population  at  800  birds  (300  non-breeding)  and  the  Islote  Afuera 
population  at  4000  birds  (1000  non-breeding)  in  June  1968.  In  1977  remains  of  nine 
birds  were  found  in  caves  along  cliffs  at  the  east  side  of  the  island  (Pierson  and  Riedman 
MS),  which  suggests  the  possibility  of  a  mainland  breeding  locale. 

The  species  occurs  near  the  islands  from  late  December  through  August,  and  many 
fly  aboard  ships  at  night.  The  birds  apparently  first  visit  the  nesting  grounds  in  February. 
Hubbs  found  none  on  the  islands  between  October  and  January  (five  trips  total)  but 
found  fresh  eggs  as  early  as  5  March.  The  peak  of  the  breeding  season  is  late  April- 
June.  Yet,  the  breeding  season  may  be  protracted,  as  Hubbs  found  fresh  eggs  as  late 
as  29-30  August.  Brownell  (MS)  reported  that  "adequate  nesting  grounds  on  the  small 
islotes  off  Guadalupe  are  almost  fully  utilized."  If  so,  nest  site  limitation  would  be  a 
strong  selective  agent  for  an  expanded  breeding  season,  as  has  apparently  occurred  in 
Leach's  Storm-Petrels. 


328 


Geographic  variation  in  the  species  has  been  discussed  by  Jehl  and  Bond  (1975); 
the  local  form  is  S.  h.  hypoleuca. 

Cassin's  Auklet  {Ptychoramphus  aleuticus) 

Although  many  ornithologists  have  noted  this  species  at  Guadalupe,  particularly 
near  the  southern  end  of  the  island  (e.g.,  Thayer  and  Bangs  1908),  it  remained  for 
Hubbs  to  discover  the  nesting  area  on  Islote  Negro.  The  species  is  not  known  to  nest 
on  Islote  Afuera  (Crossin  MS).  Brownell  (MS)  estimated  the  population  at  200  pairs 
in  April  1968,  a  figure  that  is  supported  by  Hubbs'  and  Jehl's  data. 

Hubbs  and  associates  banded  many  birds  on  I.  Negro.  In  1968  Brownell  banded 
56  and  recovered  two  that  had  been  banded  two  years  earlier,  one  as  an  adult  and  one 
as  a  downy  chick.  Other  banded  birds  were  recovered  in  April  1970  (Jehl  pers.  obs.), 
but  details  are  not  available. 

The  breeding  season  begins  in  January.  Hubbs  reported  nests  with  fresh  eggs  on 
30  January.  The  peak  in  laying  occurs  by  April  and  by  late  April  many  nests  may 
contain  young.  There  is  annual  variation  in  the  nesting  period.  For  example,  on  19 
April  1957  Hubbs  reported  numerous  burrows,  fresh  eggs,  eggs  with  embryos,  newly 
hatched  young,  and  well-developed  young;  on  23  April  1963,  27  nests  contained  only 
downy  young.  In  most  years  nesting  is  completed  by  late  June.  On  1 3  June  1955  Hubbs 
found  young  ready  to  fledge.  On  22  June  1968,  most  of  the  colony  had  completed 
nesting;  85  adults  and  40  chicks  were  present  (Crossin  MS). 

Rhinoceros  Auklet  {Cerorhinca  monocerata) 

Probably  an  irregular  winter  visitor.  There  are  records  for  1 9  April  1 925  (McLellan 
1926),  9  February  1957  (Hubbs,  specimen  LACMNH),  and  4  January  1980  (Pitman). 

Rock  Dove  (Columba  livid) 

According  to  Hubbs,  the  species  was  introduced  to  the  island  in  1956  by  residents 
of  the  settlement.  Kenyon  (MS)  reported  20  at  the  weather  station  in  1965.  In  1977 
flocks  of  up  to  six  were  recorded  at  Twin  Canyons  and  the  Lobster  Camp  (Pierson  and 
Riedman  MS).  Six  were  seen  in  the  village  (along  with  a  peafowl  [Pavo  cristatus]),  on 
5  May  1980  (Everett),  and  Duncan  reported  15-20  there  on  19-26  August  1981. 

White- winged  Dove  {Zenaida  asiaticd) 

A  specimen  of  Z.  a.  mearnsi  was  collected  on  10  June  1953  (Howell  and  Cade 
1954). 

Mourning  Dove  {Zenaida  macroura) 

This  species  was  considered  accidental  by  Howell  and  Cade  (1954),  perhaps  based 
on  the  report  of  Gaylord  (1897).  It  has  since  colonized  the  island.  Hubbs  made  the 
following  observations:  31  August  1956—1;  23  November  1964—1;  10-14  February 
1967  —  24  near  the  Lobster  Camp.  By  1970  the  species  was  widespread.  Jehl  found  a 
nest  with  two  young  near  the  village  on  14  April  and  found  a  pair,  almost  certainly 
with  a  nest,  at  the  Lobster  Camp  on  16  April.  Another  pair  was  present  on  Islote  Negro 
on  18  April.  In  November-December  1973,  Mirsky  (MS)  reported  a  few  at  Northeast 
Anchorage  and  30  or  more  near  springs.  In  August  1981,  D.  Duncan  {pers.  comm.) 
estimated  the  population  to  be  in  the  low  hundreds. 

Great  Horned  Owl  {Bubo  virginianus) 

The  presence  of  large  owls  has  not  been  verified.  Bryant  (1 887a)  reported  that  "Dr. 
Palmer's  assistant"  stated  that  a  large  owl  {Bubo)  was  present  on  the  island,  and  further 
noted  that  the  Mexican  inhabitants  reported  hearing  "hooting"  at  night.  They  said, 
however,  that  the  owl  was  very  rare.  Ridgway  (1876)  also  noted  that  "two  kinds  of 


329 


owls  were  seen"  by  the  Palmer  parly  but  that  no  specimens  were  taken.  In  1981,  the 
base  commander  told  D.  Duncan  of  large  owls  in  the  canyons  to  the  south  of  the 
airstrip. 

Burrowing  Owl  {Athene  cuniculaha) 

This  small  owl  is  widespread  and  common  on  the  main  island;  it  also  occurs  on 
Islote  Negro.  The  island  population  is  indistinguishable  from  the  mainland  form  {A. 
c.  hypugaea)  (Thayer  and  Bangs  1908). 

Vaux's  Swift  {Chaetura  vau.xi) 

One  was  seen  at  the  Sealer's  Camp  on  5  May  1980  (Everett). 

White-throated  Swift  {Aeronautes  saxatalis) 

"Regular  visitor,  at  least  formerly.  Unreported  since  1922"  (Howell  and  Cade 
1954). 

Anna's  Hummingbird  {Calypte  anna) 

According  to  Howell  and  Cade  ( 1 954)  this  hummingbird  was  evidently  uncommon 
to  rare  prior  to  1953  but  shortly  thereafter  became  established  in  the  Nicotiana  grove 
at  Northeast  Anchorage.  However,  Bryant  (1887a)  was  told  that  they  were  common 
in  palms  on  the  northwestern  slope  and  collected  one.  Howell  and  Cade  ( 1 954)  estimated 
the  population  at  1 5-20  birds  and  called  attention  to  the  different  song  of  the  local 
population,  a  difference  subsequently  established  by  Mirsky  (1976).  Mirsky  estimated 
the  population  at  approximately  100  individuals.  On  19  May  1971,  Jehl  found  a  nest 
with  two  eggs  in  a  low  shrub  near  the  top  of  the  island,  in  a  canyon  above  Barracks 
Beach. 

Allen's  Hummingbird  (Selasphorus  sasin) 

Power  (1972)  incorrectly  listed  this  species  as  breeding.  We  know  of  no  evidence 
for  its  occurrence. 

Belted  Kingfisher  (Ceryle  alcyori) 

Although  not  listed  by  Howell  and  Cade  (1954),  the  kingfisher  is  an  uncommon 
but  regular  winter  visitor.  Between  1957-1969,  Hubbs  had  eight  records  (nine  indi- 
viduals) between  25  October  and  20  April.  It  has  since  been  reported  almost  annually 
(many  observers). 

Northern  Flicker  {Colaptes  auratus  rufipileus) 

This  endemic  race  was  formerly  resident  in  the  forested  areas  atop  the  island  but 
is  now  probably  extinct.  Habitat  depletion  and  predation  by  cats  have  been  considered 
the  responsible  agents.  Apparently  it  was  fairly  common  and  as  late  as  1906,  when  last 
seen,  the  population  was  reported  as  "not  more  than  forty  individuals"  (Thayer  and 
Bangs  1908).  The  history  of  this  local  population  has  been  reviewed  by  Greenway 
(1967);  see  also  Grinnell  (1928). 

There  are  recent  reports  of  flickers  at  Guadalupe.  K.  Briggs  {pers.  comm.)  reported 
the  species  in  the  pine  forest  on  17-18  December  1972,  and  Mirsky  (MS)  saw  one  at 
Northeast  Anchorage  in  late  November-early  December  1973;  whether  these  are  fall 
migrants  from  the  mainland  or  remnants  of  the  endemic  population  is  unresolved. 

Least  Flycatcher  {Empidonax  minimus) 

Accidental.  A  specimen  of  this  eastern  species  was  taken  on  25  October  1962 
(Stager,  specimen  LACMNH). 


330 


Say's  Phoebe  (Sayornis  saya) 

The  only  report  is  of  eight  at  Northeast  Anchorage  in  November-December  1973 
(Mirsky  MS). 

* 
Northern  Rough-winged  Swallow  {Stelgidopteryx  serripennis) 

Single  birds  were  seen  on  23  November  1964  (Hubbs)  and  on  18  May  1971  (Jehl). 

Barn  Swallow  {Hirundo  rustled) 

Two  seen  on  19  May  1971  (Jehl)  were  presumed  to  be  migrants. 

Clark's  Nutcracker  {Nucifraga  columbiand) 

In  the  invasion  year  of  1972,  at  least  one  nutcracker  was  observed  in  the  pine 
forest  at  the  north  end  of  the  island  on  17-18  December  (K.  Briggs  pers.  comm.). 

Red -breasted  Nuthatch  {Sitta  canadensis) 

This  nuthatch  is  resident  in  small  numbers  in  the  pine  woods  at  the  north  end  of 
the  island.  In  1971  Jehl  found  five  pairs  there,  and  on  12-13  April  1970  he  observed 
two  pairs  feeding  young  and  found  an  additional  nest.  It  occurred  in  the  cypress  grove 
in  1953  (Howell  and  Cade  1954)  and  probably  still  does. 

Rock  Wren  {Salpinctes  obsoletus  guadalupensis) 

This  endemic  race  is  abundant  in  all  open  areas  of  the  island,  from  the  beach  to 
the  crest;  it  is  much  less  common  in  forested  areas.  In  1981,  in  one  open  area  on  top 
of  the  island,  D.  Duncan  counted  one  wren  per  50  m  in  a  20  m  wide  transect.  There 
are  no  current  estimates  of  numbers,  but  the  total  population  is  certainly  in  the  thou- 
sands. 

Bewick's  Wren  ( Thryomanes  bewickii  brevicauda) 

Extinct,  last  seen  in  1892  (Anthony  1901).  The  history  of  this  endemic  form  has 
been  reviewed  by  Grinnell  (1928)  and  Greenway  (1967).  It  resided  in  brushy  areas  and 
pines,  but  was  never  numerous.  Habitat  depletion  by  goats  and  predation  by  cats 
caused  its  demise. 

Ruby-crowned  Kinglet  {Regulus  calendula  obscurus) 

This  endemic  race  formerly  nested  in  the  cypress  grove  as  well  as  in  the  pine  forest, 
and  apparently  was  fairly  common.  Howell  and  Cade  (1954)  reported  five  singing  males 
in  the  cypress  grove  on  1 1  June  1953.  Mirsky  (MS)  reported  five  in  the  cypress  grove, 
two  in  the  pine-oak  grove,  and  one  in  the  Nicotiana  (presumably  near  the  beach)  in 
November-December  1973.  However,  birds  seen  in  winter  could  be  migrants  and  the 
current  status  of  the  endemic  population  requires  verification. 

Mountain  Bluebird  (Sialia  currucoides) 

Three  wintered  on  Guadalupe  in  1885-86;  one  was  collected  (Bryant  1887a). 

Townsend's  Solitaire  {Myadestes  townsendi) 

One  seen  on  22  March  1897  (Kaeding  1905)  is  the  only  record. 

Hermit  Thrush  (Cat hams  guttatus) 

Bryant  (1887a)  collected  three  in  the  cypress  woods  between  December  1885  and 
March  1886.  The  race  has  not  been  verified  (Miller  et  al.  1957). 


331 


American  Robin  {Turdus  migratorius) 

Bryant  (1887a)  saw  several  in  December-January  1886-87  in  the  cypress  grove. 
Mirksy  (MS)  saw  one  at  Northeast  Anchorage  on  5  December  1973. 

Varied  Thrush  {Ixoreus  naevius) 

One  was  observed  in  the  pine  forest  on  4  March  1886  (Bryant  1887^3). 

Northern  Mockingbird  {Mimus  polyglottos) 

Considered  accidental  by  Howell  and  Cade  (1954),  apparently  on  the  basis  of  a 
report  by  Bryant  (1887a),  who  saw  two  and  collected  one  on  16  March  1886.  One  was 
described  to  Jehl  on  22  February  1969. 

Sage  Thrasher  {Oreoscoptes  montanus) 

One  was  collected  on  7  January  1886  (Bryant  1887a). 

Water  Pipit  {Anthus  spinoletta) 

This  pipit  is  probably  rare  but  regular  in  migration.  Bryant  reported  a  flock  of  25 
on  2  February  1886  (Bryant  1887a). 

Cedar  Waxwing  {Bombycilla  cedwrum) 

Bryant  (1887a)  collected  one  in  the  winter  of  1885-86. 

Loggerhead  Shrike  {Lanius  ludovicianus) 

Bryant  (1887a)  saw  two  and  collected  a  female  that  had  fed  on  a  Ruby-crowned 
Kinglet  on  29  December  1885.  The  other  bird  was  heard  singing,  which  suggests  the 
possibility  of  a  mated  pair. 

European  Starling  {Sturnus  vulgaris) 

On  15  May  1971,  Jehl  saw  one  at  the  settlement  at  the  south  end  of  the  island, 
and  on  18  May  found  three  in  the  pine  forest  at  the  north  end  of  the  island. 

Yellow-rumped  (Audubon's)  Warbler  (Dendroica  coronatd) 

This  warbler  is  probably  a  regular  winter  visitor.  It  was  first  reported  by  Bryant 
(1887a)  and  has  been  seen  by  many  observers.  Mirsky  (MS)  reported  up  to  30  in 
November-December  1973  at  Northeast  Anchorage.  Everett  saw  an  example  of  the 
eastern  race  {D.  c.  coronatd)  on  18  April  1979. 

Townsend's  Warbler  {Dendroica  townsendi) 

Mirsky  (MS)  reported  three  in  the  pine-oak  woods  on  6  December  1975. 

Black-and-white  Warbler  {Mniotilta  varia) 

One  in  the  pine  forest,  19  May  1971  (Jehl). 

Ovenbird  {Seiurus  aurocapillus) 

A  specimen  of  5".  a.  aurocapillus  was  collected  on  9  June  1953  (Howell  and  Cade 
1954).  The  late  date  is  typical  for  eastern  vagrants  on  the  west  coast  in  spring. 

Common  Yellowthroat  {Geothlypis  trichas) 

One  was  collected  on  12  November  1938  (Huey  1954). 


332 


Wilson's  Warbler  ( Wilsonia  pusiUa) 

Probably  regular  in  migration  but  there  are  only  two  records;  18  May  1971  (Jehl) 
and  18  April  1979  (Everett). 

Summer  Tanager  (Piranga  rubra) 

A  specimen  of  the  eastern  race  (P.  r.  rubra)  was  collected  in  the  cypress  grove  on 
12  October  1913  (Kimball  1922). 

Rose-breasted  Grosbeak  {Pheucticus  ludovicanus) 

Two  records,  24  October  1962  (Stager,  specimen  LACMNH)  and  5  December 
1973  (Mirsky,  specimen  UCLA). 

Black-headed  Grosbeak  {Pheucticus  melanocephalus) 

The  wing  of  a  male  was  found  on  the  east  side  of  the  island  on  29  June  1977 
(Pierson  and  Riedman  MS). 

Guadalupe  Rufous-sided  Towhee  (Pipilo  erythrophthalmus  consobrinus) 

Extinct.  The  history  of  this  endemic  race  has  been  summarized  by  Grinnell  (1928) 
and  more  fully  by  Greenway  (1967).  It  was  known  to  occur  in  the  cypress  grove  and 
perhaps  elsewhere,  and  was  last  observed  in  1897.  Its  extinction  was  due  to  habitat 
depletion  by  goats  and  predation  by  cats. 

Chipping  Sparrow  (Spizella  passerina) 

Bryant  (1887<3)  collected  one  on  6  January  1886,  and  Mirsky  (MS)  reported  the 
species  in  the  Nicotiana  and  in  the  pine-oak  forest  in  November-December  1973. 

Fox  Sparrow  {Passer ella  iliaca) 

An  example  of  P.  /.  sinuousa  collected  on  16  February  1886  (Bryant  1887a),  seems 
to  represent  the  southernmost  record  for  the  species  on  the  Pacific  coast. 

Lincoln's  Sparrow  {Melospiza  lincolnii) 

Bryant  (1887a)  collected  individuals  on  5  and  1 9  February  1886,  and  Swarth  (1933) 
reported  a  specimen  taken  on  16  March  1932. 

White-throated  Sparrow  {Zonotrichia  albicollis) 

One  collected,  10  October  1913  (Kimball  1922). 

Golden-crowned  Sparrow  {Zonotrichia  atricapilla) 

Bryant  (1887a)  collected  two  on  16  February  and  one  on  4  March  1886,  in  the 
pines. 

White-crowned  Sparrow  {Zonotrichia  leucophrys) 

Probably  regular  in  migration  but  the  only  report  is  of  two  near  the  south  end  of 
the  island  on  14  April  1970  (Jehl). 

Guadalupe  Dark-eyed  Junco  {Junco  hyemalis  insularis) 

Knowledge  of  this  endemic  junco  was  fully  summarized  by  Howell  (1968);  addi- 
tional information,  including  variation  in  the  song,  was  provided  by  Mirsky  (1976). 
At  one  time  the  junco  was  one  of  the  most  abundant  birds  on  the  island  (Palmer,  in 
Ridgway  1876).  Today  it  is  uncommon  and  is  much  less  abundant  than  the  House 


333 


Finch  or  Rock  Wren.  It  may  be  found  scattered  along  the  northern  half  of  the  island 
wherever  there  is  vegetation.  It  often  feeds  on  the  ground,  in  litter  at  the  base  of  pine 
trees  but  also  in  the  oaks.  However,  it  seems  adaptable  and  now  occupies  stands  of 
Nicotiana  on  the  beach.  Breeding  occurs  from  late  January  (Bryant  1887a)  to  at  least 
late  April  (Howell  1 968).  On  1 7  May  1 97 1  Jehl  saw  young  juncos  that  were  independent 
of  the  parents.  The  taxonomic  relationships  of  this  junco  have  been  fully  discussed  by 
Miller  (1941),  who  argued  that  the  local  population  was  derived  from  migratory  ances- 
tors. "The  Guadalupe  junco  is  distinguished  principally  by  its  relatively  long  bill  and 
short  wing  and  tail  .  .  .  and  virtual  absence  of  sexual  dimorphism  in  color"  (Howell 
1968).  The  long  bill  is  used  to  extract  seeds  from  deep  in  pine  cones  (Jehl  pers.  obs.). 
Power  (1980)  also  discussed  the  morphology  of  this  species.  Bryant  (1887a)  collected 
a  migrant  of  one  of  the  mainland  races  {thurberil,  cf.  Miller  et  al.  1957)  on  6  January 
1886,  that  was  being  attacked  by  a  resident  junco. 

Western  Meadowlark  {Sturnella  neglecta) 

Bryant  (1887a)  reported  one  on  the  crest  of  the  island  on  22  March  1886. 

Brewer's  Blackbird  {Euphagus  cyanocephalus) 

A  female  was  seen  on  12  December  1973  (Mirsky  MS). 

Scott's  Oriole  {Icterus  parisorum) 

Mirsky  (MS)  reported  two  males  and  three  females  in  the  Nicotiana  at  Northeast 
Anchorage  from  23  November  to  3  December  1973. 

Guadalupe  House  Finch  {Carpodacus  mexicanus  amplus) 

This  endemic,  the  second  commonest  landbird,  may  occur  almost  anywhere,  in- 
cluding Islote  Negro,  but  is  most  common  near  vegetation  and  at  the  village.  The  entire 
population  may  exceed  1000.  Bryant  (1887a)  provided  information  on  nests  and  nest 
sites.  He  also  noted  that  the  finches  were  captured  and  eaten  by  locals.  The  evolution 
and  geographic  variation  of  this  race  have  been  reviewed  by  Power  (1979). 

Red  Crossbill  {Loxia  curvirostra) 

Howell  and  Cade  (1954)  reported  this  species  as  "formerly  resident;  no  definite 
breeding  record;  unreported  since  1 903."  Evidently  it  was  once  fairly  common  as  Bryant 
( 1 887a)  reported  about  20  in  the  pines  in  1886,  and  reported  collecting  nine  specimens, 
including  an  immature  in  February-March  1896.  According  to  K.  C.  Parkes,  six  birds 
collected  by  A.  W.  Anthony  on  20  September  1896  include  a  female  almost  molted 
out  of  juvenile  plumage  and  five  full-grown  juveniles.  The  species  was  also  reported 
as  being  "resident"  by  Gaylord  (1897),  though  he  did  not  observe  it.  Kaeding  (1905) 
reported  "a  few"  in  1897.  Grinnell  (1928)  reviewed  the  status  of  the  species  and 
examined  the  specimens,  which  he  attributed  to  L.  c.  bendirei.  A.  R.  Phillips,  however, 
now  refers  all  specimens  to  L.  c.  benti  {fide  K.  C.  Parkes). 

Goldfinch  {Carduelis  sp.) 

Townsend  (1916)  states  "the  Goldfinch  was  observed."  There  is  no  additional 
information. 

Discussion 

The  Guadalupe  avifauna  was  well-studied  in  the  late  1 9th  and  early  20th  centuries. 
Recent  studies  have  provided  new  information  on  seabirds;  yet,  much  remains  to  be 
learned.  The  known  colonies  are  diflftcult  to  reach  and  most  visits  to  them  have  been 
made  in  winter  or  spring.  Studies  during  other  seasons  are  needed  to  clarify  breeding 


334 


seasons.  Efforts  are  also  needed  to  locate  seabird  colonies  on  the  main  island,  especially 
in  light  of  recent  reports  of  storm-petrels  calling  there  at  night,  and  to  determine  the 
morphological  characters  of  any  such  populations.  It  is  not  inconceivable  that  the 
Guadalupe  Storm-Petrel  has  escaped  extinction. 

The  island's  rugged  topography  and  lack  of  fresh  water  have  inhibited  recent  work 
along  the  central  axis.  Surveys  in  forested  areas  during  the  breeding  season  are  needed 
to  determine  the  status  of  the  endemic  races  of  the  Ruby-crowned  Kinglet  and  Northern 
Flicker.  Both  species  are  common  on  the  mainland  and  highly  migratory,  so  sight 
records  are  not  proof  of  the  persistence  of  endemic  races.  Even  evidence  of  breeding 
may  be  equivocal,  as  secondary  invasions  by  these  species  could  have  taken  place. 
These  studies  will  be  difficult  and  will  require  capturing  or  collecting  some  birds. 

Faunas  of  oceanic  islands  are  not  constant.  New  species  arrive  regularly;  some 
become  established,  and  others  disappear.  The  factors  that  affect  successful  colonization 
or  promote  extinction  are  difficult  to  establish  (Jehl  and  Parkes  1983)  but  are  critical 
to  understanding  avian  distribution.  On  Guadalupe,  a  new  food  source  {Nicotiana 
glauca),  may  have  been  a  major  factor  in  allowing  Anna's  Hummingbirds  to  colonize 
(or  become  more  common?)  in  the  past  several  decades.  Mourning  Doves  have  also 
become  established,  though  the  reasons  why  are  unstudied. 

In  view  of  the  importance  of  island  faunas  to  current  theories  in  biogeography,  regular 
surveys  should  be  encouraged  (e.g.,  at  least  every  decade)  so  that  changes  can  be  detected 
as  they  are  occurring  or  shortly  afterward.  Such  data  will  be  especially  useful  from 
islands,  like  Guadalupe,  where  a  strong  historical  record  has  been  established. 

Acknowledgments 

We  are  indebted  to  Laura  C.  Hubbs  and  Elizabeth  N.  Shor  for  access  to  the  field 
notes  of  the  late  Carl  L.  Hubbs,  and  to  the  following  for  permitting  us  to  include  their 
unpublished  records:  M.  Bonnell,  K.  Briggs,  R.  Brownell,  E.  Chu,  R.  Condit,  R.  Crossin, 
R.  DeLong,  D.  Duncan,  K.  Kenyon,  B.  LeBoeuf,  E.  Mirsky,  M.  Pierson,  R.  Pitman, 
M.  Riedman,  B.  Tyler,  and  R.  Wisner. 

R.  DeLong,  R.  McConnaughey  and  R.  Moran  assisted  in  various  aspects  of  the 
field  research.  We  especially  acknowledge  the  kindness  of  the  late  Carl  L.  Hubbs,  who 
made  it  possible  for  Jehl  and  many  others  to  participate  in  research  at  Guadalupe. 

The  photograph  of  the  Guadalupe  Storm-Petrel  was  provided  by  the  Field  Museum 
of  Natural  History  through  the  courtesy  of  M.  Traylor.  K.  C.  Parkes,  G.  Pregill,  and 
D.  Steadman  made  helpful  comments  on  an  earlier  draft  of  this  paper. 

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