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HARVARD    UNIVERSITY 

Library  of  the 

Museum  of 

Comparative  Zoology 


LIBRARY 

DEC  2  01982 

nARVAKD 
UNIV^RJ^ITY 


VOLUME  23 
1981-1982 


TULANE  UNIVERSITY 
NEW  ORLEANS 


TULANE  STUDIES  IN  ZOOLOGY  AND  BOTANY,  a  publication  of  the  Biology 
Department  of  Tulane  University,  is  devoted  primarily  to  the  biology  of  the  waters  and 
adjacent  land  areas  of  the  Gulf  of  Mexico  and  the  Caribbean  Sea,  but  manuscripts  on 
areas  outside  this  geographic  area  will  be  considered.  Each  number  contains  an  indivi- 
dual monographic  study  or  several  minor  studies.  Normally  two  numbers  plus  an  index 
and  a  table  of  contents  are  issued  annually.  Preferred  citation  of  the  journal  is  Tulane 
Stud.  Zool.  and  Bot. 

INFORMATION  FOR  AUTHORS:  Manuscripts  submitted  for  publications  are  eval- 
uated by  the  editors  and  by  an  editorial  committee  selected  for  each  paper.  Contrib- 
utors need  not  be  members  of  the  Tulane  faculty.  Manuscripts  of  20  or  more  pages, 
double-spaced,  are  preferred.  We  recommend  conformance  with  the  principles  stated 
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Illustrations  should  be  proportioned  for  one  or  two  column  width  corresponding  to  our 
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Volume  23.  $8.50  domestic,  $9.50    foreign. 

Copies  of  Tulane  Studies  in  Zoology  and  Botany  sent  to  regular  recipients,  if  lost  in  the 
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COMMUNICATIONS:  Address  all  queries  and  orders  to:  Editor,  TSZ&B,  Depart- 
ment of  Biology,  Tulane  University,  New  Orleans,  Louisiana  701 18,  U.S.A. 
Harold  A.  Dundee,  Editor 


CONTENTS  OF  VOLUME  23 
Number  Page 

1.  BIOSYSTEMATICS  OF  THE  KINOSTERNON  HIRTIPES  SPECIES 
GROUP  (TESTUDINES:  KINOSTERNIDAE) 

JohnB.  Iverson        1 

LIFE  HISTORY  OF  ETHEOSTOMA  COOSAE  (PISCES:  PERCIDAE) 

IN  BARBAREE  CREEK,  ALABAMA  

Patrick  E.O'Neil      75 

THE   TAXONOMIC   RELATIONSHIP   BETWEEN  MALACLEMYS 
GRAY,    1844  AND  GRAPTEMYS  AGASSIZ,    1857  (TESTUDINES: 

EMYDIDAE) 

James  L.Dobie      85 

2.  CHANGES  IN  MELANIN  MIGRATION  INDUCED  BY  NORADREN- 
ERGIC AND  HISTAMINERGIC  AGENTS  IN  THE  FIDDLER  CRAB, 
UCA  PUGILATOR  

Mukund  M.  Hanuamante  and  Milton  Fingerman     103 

ADDITIONAL    TREMATODES    OF    MAMMALS    IN    LOUISIANA 
WITH    A    COMPILATION   OF   ALL   TREMATODES    REPORTED 

FROM  WILD  AND  DOMESTIC  ANIMALS  IN  THE  STATE 

Wesley  L.  Shoop  and  Kenneth  C.  Corkum     109 

COMPARATIVE     VISCERAL     TOPOGRAPHY     OF     THE     NEW 
WORLD  SNAKE  TRIBE  THAMNOPHIINI  (COLUBRIDAE,  NATRI- 

CINAE) 

Nita  J.  Rossman,  Douglas  A.  Rossman,  and  Nancy  K.  Keith     123 


TULANE  STUDIES  IN  ZOOLOGY  AND  BOTANY 

VOLUME  23 

INDEX  TO  SCIENTIFIC  NAMES 

(NEW  TAXONOMIC  ENTITIES  IN  BOLDFACE) 


JOL 


Ablates  baliodeira,  129,  137 
Acrochordidae,  129,  163 
Acrochordis,  137 

arafurae,  137,  163 

granulatus,  129,  137,  163 
javanicus,  129,  137,  163 
A  de  lop  his,  124 

Agamodistomum  marcianae,  112 
Agkistrodon  piscivorus,  129 
A haetulla  prasinus,  1 37 
Alaria  alahodes,  110-112,  115,  119,  120 

americana,  112-113 

canis,  112 

da  th  rata,  112 

marcianae.  111,  113, .115,  119-120 

minnesotae,  112 

mustelae,  110-111,  119 

pseudoclathrata,  112 
Ambloplites  rupestris,  11 
Amnicola,  118,  120 

Integra,  118 
longiqua,  114 
Amphiesma  vibakari,  164 
Amphimerus,  114 

caudalitestis,  113-114 

interrupt  us,  113-114 

minimus,  113-114 

/jeo  tropicalis,  113-114 

parciovatus,  113-114 

/?r/ce/,  113-114 

pseudofelinus,  113-114,  120 

speciosus.  111,  114-115,  119-120 
Ancistrodon  rhodostoma,  137 
Aniliidae,  129,  137 
Aromochelys  carinatus,  41 

Baschkirovitrema  incrassatum,  1 10-1 11,  117, 

119-120 
Boidae,  129,  137,  163 
Boiga,  163 
Bolyeria,  163 

Brachylaima,  virginiana,  110-111,  119 
Bungarus  candidus,  137 
fasciatus,  137 

Calamaria  multipunctata,  129,  137 
Calloselasma  rhodostoma,  137 
Cambarellus  puer,  118 
Campostoma  anomalum,  11 
Carneophallus  basodactylophallus,  110-111,  119 
Catonolus,  78 
f label  lore,  76 


DEC  20'    '^ 


species,  78 
Catostomus  commersoni,  116 
Causus  rhombeatus,  163-164 
Cerberus  rhynchops,  137 
Cercaria  marcianae,  1 1 2 
Chamaetortus,  163 
Chinosternum  hirtipes,  46 
Chrysemys,  88-90,  92-95,  97-99 
insculpta,  95 

p/cro,  8,  94-95,  97-98,  100 
Cinosternon  henrici,  41,  61 
hippocrepis,  22 

/7/W//7e5,  18,  23,  41,  44-45,  48,  65 
pensylvanicum,  21-22,  45-46,  48-49 
species,  46,  48 
Cinosternum  flavescens,  41,  61 
integrum,  15,  46,  49,  51 
sonoriense,  41,  49 
Claudius  angustatus,  23 
Clemmys,  88-89,  92-95 
gwr/a?fl,  90,  95,  98,  100 
insculpta,  90,  95,  98,  100 
marmorata,  72,  100 
muhlenbergi,  100 
Clonophis,  126-127,  134-135,  137,  139,  142, 
144-146,  155,  157 
kirtlandi,  125,  128-132,  134-136,  145-156,  159 
Codoma  ornatus,  1 
Coluber  melanurus,  129,  137 
oxycephalus,  163-164 
radiatus,  137 
Colubridae,  129,  137,  163-164 
Compsemys,  97 
Corallus,  163 
Cottus  carolinae,  11 
Crotaphopeltis  hotamboeia,  163 
Cryptocotyle,  116 
concava.  111,  114-116,  119-120 
echinata,  114 
Cylindrophis  rufus,  129,  137 

Deirochelys,  87-90,  92-95 

com.  98 

reticularia,  98,  100 
Dendrelaphis  pictus,  129,  137 
Dendrophis  pictus,  137 
Didelphis  virginiana,  109,  118 
Diplostomidae,  110 
Diplostomum,  112 

alaroides,  110,  112 

fosteri,  1 1 2 
Dipsadoboa,  163 


Distoma  concava,  114 
Dryophis  praslnus,  1 37 

Echinochasmus  schwartzi,  119 
Echinocirrus  metis,  116 
Echinostoma  revolutum,  117 
Echinostomatidae,  116 
Echmatemys,  98-99 

pusilla,  98 
Elaphe  flavilineata,  129,  137 

radio  t  us,  137 
Elapidae,  129,  137 
Elapoides  fuscus,  137,  164 
Emydoidea,  88-90,  92-95,  98 
bland ingi,  98,  100 
Enhydrina  schistosa,  137 
Enhydris  alternans,  129,  137 

enhydris,  137,  164 

plumbea,  129,  137,  164 
Enhydrodiploslomum,  1 12 

alarioides,  110 
Etheostoma,  76,  82 

acuticeps,  82 

barbouri,  81 

blennioides,  82 

coosae,  75-81 

rfi/ry/,  75-76,  79 

flabellare,  76 

fonticola,  82 

gracile,  82 

jordani,  11 

kennicotti,  81 

nigrum,  82 

proeliare,  81 

radiosum  cyanorum,  82 

simoterum,  75,  79 

species,  79 

squamiceps,  78,  80 

stigmaeum,  11 
Eunectes,  163 
Euparyph  'um,  116-117,  1 20 

beaveri,  117,  120 

A«e//5,  116-117 
Exiliboa  placata,  164 

Easciola  putori,  1 16 

trigonocephala,  1 1 6 
Felis  domesticus,  109,  113 
Fibricola  cratera,  110-111,  119 

/wc/rffl,  110-111,  119 
Fordonia  leucobalia,  129,  137 
Fundulus  stellifer,  11 

Gasterosteus  aculeatus,  1 1 6 
Gongylosoma  baliodeira,  129,  137 
Gonysoma  oxycephalus,  163-164 


Graptemys,  85-88,  90-102 

barbouri,  86,  91,  96,  98,  100 

cag/e/,  86,  91,  98,  100 

cordifera,  97 

ftavimaculata,  86-87,  91,  98,  100 

geographica,  89-91,  93,  98,  100 

inornata,  97 

nigrinoda,  86,  90-92,  96,  98,  100 

oculifera,  86,  91,  98,  100 

ouachitensis,  98,  100 

ouachitensis  ouachitensis,  91,  100 

ouachitensis  sabinensis,  86,  91,  100 

pseudogeographica,  86,  90-94,  98,  100 

pseudogeographica  kohni,  101 

psdudogeographica  pseudogeographica,  100 

pulchra,  86-87,  89,  91-94,  98,  100-101 

ver^a,  86,  91,  98,  100,  101 
Gyrauiis  parvus,  119 
Gyrosoma  singulare,  110-111,  119 

Hasstilesia  texensis,  110-111,  119-120 
Heterobilharzia  americana,  110-111,  1 1 9- 1 20 
Heterophyidae,  114 
Homolopsinae,  129,  137,  164 
Homalopsis  buccata,  129,  137 
Hydrophis  fasciatus,  129,  137 
Hypentelium  etowanum,  11 
Hypsirhina  alternans,  129,  137 
plumbea,  137,  164 

Ictalurus  natalis,  11 

species,  8 
Isonychia,  75 
Isthmiophora,  116,  120 

mefc.  111,  115-116,  119-120 

Kinosternidae,  1,9,  11,  13 

Kinosternon,  13,  15,  19,  21-22,  29-30,  35,  41,  43, 

46,  55 
acutum,  23,  35 
alamosae,  1,  2,  7,  21,  23,  41 
angustipons,  35 
/70«/-/,  35,  55 
cobanum,  23 
dunni,  35 
flavescens,  2,  4,  6,  7-11,  22,  40-41,  44,  46,  50, 

55,  60 
henrici,  40,  41,  61 
herrerai,  22,  55 
hertipes,  46,  50 
hirtipes,  1-4,  6,  8-24,  26-31,  34-35,  38-41,  44, 

46-56,  60,  64 
hirtipes  chapalense,  subsp.  novum,  46,  51,  54, 

57,  65 
hirtipes  chapalense  X  K.h.  murrayi,  65 
/?/W//7e5  group,  1-2,  4-7,  11,  15,  20,  22,  24-28, 

33-38,  42,  46,  54-55 


hirtipes  hirtipes,  37,  46,  48-52,  59,  63,  65 
hirtipes  magdalense,  subsp.  novum,  6,  33,  46, 

53-54,  58,  64 
hirtipes  megacephalum,  subsp.  novum,  6,  33, 

37,  46,  52,  54,  58,  64 
hirtipes  murrayi,  6,  21.  33,  34,  37,  39,  46,  48-52, 

59,62 
hirtipes  tarascense,  subsp.  novum,  33-34,  37,  46, 

52,  54,  58,  64 
integrum,  1-5,  7,  12-18,  21-23,  40-41,  44-45,  48, 

55,  64-65 
leucostomum,  21,  23 
leucostomum  group,  35 
murrayi,  21,  46,  50 
oaxacae,  1,  2 
oblongum,  44 
pennsilvanicum,  11,  45 
punctatum,  41 
scorpiodes,  1,  2,  8,  20-21 
scorpiodes  group,  1,  2,  5,  22-23,  29,  35 
sonorensis,  41-43,  46,  55,  61 
sonoriense,  1-8,  12-14,  19-20,  22-31,  33-35, 

37-38,  40-46,  54-55,  60,  63 
sonoriense  longifemorale,  subsp.  novum,  6, 

42-44,  54,  56,  62 
sonoriense,  sonoriense,  56,  60 
species,  48-49 
steindachneri,  55 
subrubrum,  1\-11,  31,  35,  41,  55 
subrubrum  group,  55 
subrubrum  hippocrepis,  11 
subrubrum  steindachneri,  31,  55 
subrubrum  subrubrum,  35 
subrubrum  triliratum,  23 

Lepomis  cyanellus,  11 

gulosus,  11 

macrochirus,  11 

megalotis,  11 
Leptodira  hotamboeia,  163 
Lichanura,  163 

L  ins  to  wiella  szidati,  110-111,  1 1 9- 1 20 
Loxocemus,  163 
Lutra  canadensis,  109-110 
Lynxrufus,  109,  112 

Malaclemys,  85-88,  90,  92-100 

terrapin,  86,  91-93,  95-98,  100 

terrapin  centrata,  87 

terrapin  littoralis,  87 

terrapin  macrospilota,  87 

terrapin  pileata,  87 

terrapin  terrapin,  70 
Maritreminoides  nettae,  110-111,  1 1 9- 1 20 
Microphallidae,  117 
Microphallus  opacus.  Ill,  115,  1 17-120 


ovatus,  117 
Micropterus  coosae,  11 

punctulatus,  11 
Moxostoma  duquesnei,  11 
Mustek  vison,  109-110,  114,  117 

Naja  tripudians,  137 
Natricinae,  124,  129,  137,  164 
Matrix  chrysarga,  129,  137 
erythrogaster  alta,  1 1 
subminiata,  129,  137 
trianguligera,  129,  137,  164 
vibakari,  164 
vittata,  129,  137 
Nerodia,  126-127,  134-137,  139,  141,  144-147, 
149-152,  155,  157 
cyclopion,  125,  128,  130-132,  135-136,  138, 

140-143,  145-156,  159 
erythrogaster,  125,  128-132,  134-136,  138, 

140-143,  145-156,  159 
fasciata,  125,  128,  130-132,  135-136,  138, 

140-143,  145-156,  159 
rhombifera,  125,  128-132,  135-136,  138, 

140-143,  145-156,  159 
sipedon,  125,  128,  130-132,  135-136,  138, 

140-143,  145-156,  159 
valida,  125,  128,  130-132,  134,  138,  140-143, 
145-156,  159 
Notocotylidae,  118 
Notocotylus  quinqueserialis,  118 

urbanensis,  118 
Notropis  asperifrons,  11 
callistius,  11 
lirus,  11 
ornatus,  1 
stilbius,  11 
trichroistius,  11 
venustus,  11 
xaeocephalus,  11 
Nyctanassa  violacea,  114 
Nudacotyle  novicia,  119 

Ondatra  zibethica,  109,  118 
Opisthorchidae,  113 
Ozotheca  hirtipes,  45 
odorata,  21,  45,  48 

Paragonimidae,  118 
Paragonimus  kellicotti.  111,  1 18-120 
Paramonostomum  pseudalveatum,  119 
P  ere  in  a  caprodes,  11 

nigrofasciata,  77,  81 
Phagicola  angrense,  1 1 9 

nana,  119 
Pharyngostomoides  procyonis,  110-111,  119 
Phenacobius  ca  tost  am  us,  11 


Philothalmus  semivariegalus,  1 63- 1 64 
Phopalias  macracanthus,  119 
Platylhyra  flavescens,  22 
Poecilia  reticulatua,  82 
Pomatiopsis  lapidaria,  1 1 8 
Procambarus  clarki,  118 
Procyon  lotor,  109,  112-113,  116-117 
Psamnophis  sibilans,  163 
Pseudemys,  85-90,  92-96,  98-99 

alabamensis,  101 

concinna,  87-88,  94,  101 

floridana,  88,  94,  101 

nelsoni,  101 

rubiventris,  87-88,  101 

5cr//?/a.  7,  10-11,  14,  70,  87-88,  101 

scrip ta  elegans,  18 

stejnegeri,  101 
P/v'a5  korros,  137 

mucosa,  137 

Quinqueserialis  hassali,  118 

quinqueserialis,  111,  115,  1 1 8- 1 20 

/?eg/«fl,  126-127,  135,  137,  139,  141,  144,  146,  147 
a//e/j/,  125,  128-132,  135-136,  138,  140,  145, 

146-151,  153-156,  159 
grahamii,  125,  128,  130-132,  134-136,  138, 

140-143,  145-156,  160 
rigida,  125,  128,  130-132,  134-136,  140-143, 

145-156,  160 
septemvittata,  125,  128,  130-132,  135-136,  138, 
140-141,  145-156,  160 
Rhabdophis  chrysarga,  129,  137 

subminiata,  129,  137 
Rhinoclemmys,  88-90,  98-99 
areola  ta,  101 
pulcherrima,  101 
sp.,  101 
Rhopalias  macracanthus,  110-111 

Sellacotyle  vitellosa,  110-111,  120 

Seminatrix,  126-127,  134-135,  139,  141-142,  144, 

155,  157 
pygaea,  126,  128-132,  134-136,  145-156,  160 
Semotilus  atromaculatus,  11 
Sinonatrix  trianguligera,  129,  137,  164 
Staurotypus  triporcatus,  23 
Sternotherus  odoratus,  9,  21,  46,  50 
Storeria,  126,  129,  134,  137,  139,  141,  144,  155, 

157,  159 
dekayi,  125,  127-128,  130-132,  134-136,  138, 

140-143,  145-156,  161 
occipitomaculata,  126-128,  130-132,  134-138, 

140-143,  145-156,  160 
Swanka  henricii,  41 


Sytvilagus  aquaticus,  119 

Terrapene,  87-90,  92-95 
Carolina,  101 
ornata,  101 
Testude  pensilvanica,  21-22 
Thalassophis  anomalus,  137 
Thamnophiini,  124,  137,  164 
Thamnophis,  126-127,  129,  134,  137,  139,  151-152, 

154-155 
angustirostris,  126 

brachystoma,  126,  128,  130-132,  145-156,  160 
butleri.  126,  128,  130-132,  145-156,  160 
chrysocephalus,  130-132,  135-136,  145-156,  160 
couchii,  125 
couchii  A,  128,  130-132,  135-136,  138,  140-143, 

145-156,  160 
couchii  B,  128,  130-132,  135-136,  138,  140-143, 

145-156,  160 
couchii  couchii,  126,  128,  138 
couchii  hydrophilus,  126,  128,  160 
cyrtopsis,  126,  128,  130-132,  135-136,  138, 

140-143,  145-156,  160 
elegans,  125,  135 
elegans  gxouv,  126,  130,  132,  139,  141-144, 

157-158 
elegans  A,  128,  130-132,  135-136,  138,  140-143, 

145-156 
elegans  B,  128,  130-132,  135-136,  138,  140-143, 

145-156 
elegans  terrestris,  126,  128,  160 
elegans  vagrans,  126,  128,  160 
eques,  125-126,  128,  130-132,  134-136,  138, 

140-143,  145-156 
eques  megalops,  125,  160 
eques  virgatenuis,  125,  160 
godmani,  126,  128,  130-132,  135-136,  145-156, 

160 
marcianus.  126,  128,  130-132,  135-138,  140-143, 

145-156,  160 
megalops,  126 
melanogaster,  126,  128-132,  134-138,  140-143, 

145-156,  160 
nigronuchalis,  126,  128-136,  138,  140-143, 

145-156,  160 
ordinoides,  126,  128,  130-132,  145-156,  160 
proximus,  126,  128-132,  134-139,  140-143, 

145-156,  160 
radix,  126,  128,  130-132,  134-156,  161 
radix  gxonp,  126,  130,  132,  139,  141-144,  157-158 
rufipunctatus,  126,  128-136,  145-156,  161 
sauritus,  126,  128-132,  134-138,  141-143, 

145-156,  161 
sauritus  gxoxx^p,  126,  130,  132,  139,  141-144,  153, 

157-158 


scalaris.  126,  128,  130-132,  135-137,  145-156,  161 
sirtalis  A,  128,  130-132,  135-136,  138,  140-143, 

145-156 
sirtalis  B,  128,  130-132,  135-136,  138,  140-143, 

145-156 
sirtalis  fitchi,  126,  128,  138,  161 
sirtalis  group,  126,  130,  132,  139,  141-144, 

157-158 
sirtalis  sirtalis,  126,  128,  161 

Tocotrema  concava,  114 

Thyrosternum  henrici,  41 
hirtipes,  45 
sonoriense,  41 

Trachemys,  88,  94-95 

Trachyboa  gularis,  163 

Trimeresurus  gramineus,  1 37 

Trionyx  spiniferus,  4,  10 

Tropidoclonion,  126-129,  134,  137,  139,  141-142, 
144,  155,  157 
lineatum,  125-126,  134-136,  161 
lineatum  A,  130-131,  133,  145-151,  153-156 
lineatum  B,  130-131,  133,  135-136,  138,  140-143, 
145-156 

Tropidophiidae,  164 

Tropidophis,  163 

Uca  pugilator,  103-106 
Ulocentra,  76,  79 

Viperidae,  163 

Virginia,  126-127,  134,  137,  139,  141,  144,  155,  157 
striatula,  126,  128,  130-131,  133-138,  140-143, 

145-156,  161 
valeriae,  126,  128,  130-131,  133-138,  140-143, 

145-156,  161 
valeriae  elegans,  128 
Vulpesfulva,  109 

Xenochrophis  vittata,  137 
Xenopeltis,  163 
unicolor,  137 

Zamenis  florulentus,  163 
rhodorachis,  163 


i(D®IL(D(g^ 


JAN  4     iQn9 


Volume23  .  Number  1 


$5.50 


K^ft}^,9n5?iCSo,  1981 
UN/IVK.cjsiTV 


BIOSYSTEMATICS  OF  THE  KINOSTERNON  HIRTIPES 
SPECIES  GROUP  (TESTUDINES:  KINOSTERNIDAE) 

JOHN  B.  IVERSON 


p.  1 


LIFE  HISTORY  OF  ETHEOSTOMA  COOSAE  (PISCES:  PERCIDAE) 
IN  BARBAREE  CREEK,  ALABAMA 

PATRICK  E.  O'NEIL 


p.  75 


THE  TAXONOMIC  RELATIONSHIP  BETWEEN  MALACLEMYS  GRAY,  1844 
AND  GRAPTEMYS  AGASSIZ,  1857  (TESTUDINES:  EMYDIDAE) 


JAMES  L.  DOBIE 


p.  85 


TULANL  UNIVERSITY 
NEW  ORLEANS 


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TULANE  STUDIES  IN  ZOOLOGY  AND  BOTANY 


Volume  23,  Number  1 


December  30,  1981 


BIOSYSTEMATICS  OF  THE  KINOSTERNON  HIRTIPES 
SPECIES  GROUP  (TESTUDINES:  KINOSTERNIDAE) 

JOHN  B.  IVERSON' 

Dept.  of  Biology 

Earl  ham  College 

Richmond,  Indiana  47374 


Abstract 

Geographic  variation  in  scute  and  shell  measure- 
ments (via  muhivariate  statistical  analysis),  body 
size,  head  scale  and  chin  barbel  morphology,  size  of 
first  neural  bone,  shell  carination,  and  head  size  and  pat- 
terns in  populations  of  the  Kinostemon  hiritpes  species 
group  were  analyzed.  The  results  support  the  retention 
of  allopatric  K.  sonoriense  and  K.  hirtipes  as  full  species 
in  the  group,  and  the  recognition  of  two  eillopatric  sub- 
species (one  new)  of  K.  sonoriense  and  six  subspecies 
(four  new  and  all  apparently  allopatric)  of  K.  hirtipes. 
The  description  of  each  taxon  includes  complete  synon- 
omies  and  ecological  and  reproductive  data.  Also  in- 
cluded are  a  key  to  adults  and  a  discussion  of  all  taxa. 

Introduction 

Prior  to  1970,  members  of  the  Kinoster- 
non  hiriipes  species  group  were  cited  more 
than  233  times  in  the  literature.  At  least  half 
of  those  citations  contained  errors  in  identi- 
fication, locality,  and/or  orthography.  Iver- 
son  (1976,  1978),  Conant  and  Berry  (1978), 
Iverson  and  Berry  (1979),  and  Berry  and 
Legler  (1980)  have  each  addressed  some  of 
the  problems  dealing  with  members  of  this 
group  in  the  American  southwest,  adjacent 
northwestern  Mexico,  and  northeastern 
Mexico.  Clearly  the  distribution,  identifica- 
tion, systematics,  and  phylogeny  of  the  tur- 


'Adjunct  Assistant  Curator  of  Herpetoiog)',  Florida 
State  Museum,  University  of  Florida,  Gainesville,  FL 
32611 


ties  of  the  Kinosternon  hirtipes  species 
group  are  poorly  understood.  The  purpose 
of  this  report,  as  part  of  a  continuing 
analysis  of  relationships  within  the  family 
Kinosternidae,  is  to  rectify  this  situation. 

My  objectives  here  are  1)  to  redefine  the 
members  of  this  group  taxonomically,  2)  to 
analyze  patterns  of  geographical  variation 
in  external  morphological  characters,  3)  to 
develop  a  phylogeny  of  these  members,  and 
4)  to  correct  and  bring  order  to  the  confus- 
ing and  erroneous  literature. 

Identification  Of  The 
Kinosternon  Hirtipes  Species  Group 

One  of  the  primary  obstacles  to  the  study 
of  Mexican  kinosternids  has  been  the  diffi- 
culty in  distinguishing  members  of  the  K. 
hirtipes  species  group  (K.  hirtipes  and  K. 
sonoriense)  from  those  of  the  A',  scorpioides 
group  (fide  Berry  1978;  including  K.  scor- 
pioides, K.  alamosae,  K.  oaxacaedSid^K.  in- 
tegrum), especially  where  the  groups  occur 
sympatrically.  Adult  males  of  the  hirtipes 
group  are  readily  distinguished  by  the  pres- 
ence of  a  patch  of  elevated  scales  on  the 
posterior  crus  and  thigh  of  each  hindleg 
(vinculae:  fide  H.  M.  Smith  and  R.  B. 
Smith  1980),  absent  in  turtles  of  the  scor- 
pioides group,  but  adult  females  of  the  hir- 
tipes group  lack  these  structures  and  are 
thus  often  difficult  to  identify.  An  elabora- 
tion of  the  differences  between  K.  integrum 


EDITORIAL  COMMITTEE  FOR  THIS  PAPER: 

Dr.  James  F.  Berry,  Assistant  Professor  of  Biology,  Elmhurst  College, 
Elmhurst,  Illinois  60126 

Dr.  Robert  G.  Webb,  Professor  of  Biological  Sciences,  University  of  Texas  at 
El  Paso,  El  Paso,  Texas  79999 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


and  members  of  the  K.  hirtipes  group  is 
therefore  justified  (see  also  Iverson  and 
Berry  1979),  especially  since  they  coexist  in 
several  Mexican  drainage  basins  (see  MA- 
TERIALS AND  METHODS).  Kinosternon 
scorpioides  is  not  sympatric  with  members 
of  the  K.  hirtipes  group. 

Identification  Of  The 
Kinosternon  Hirtipes  Species  Group 

One  of  the  primeuy  obstacles  to  the  study 
of  Mexican  kinostemids  has  been  the  diffi- 
culty in  distinguishing  members  of  the  K. 
hirtipes  species  group  (K.  hirtipes  and  K. 
sonoriense)  from  those  of  the  K.  scorpioides 
group  (fide  Berry  1978;  including  K.  scor- 
pioides, K.  alamosae,  K.  oaxacae  and  K.  in- 
tegrum), especially  where  the  groups  occur 
sympatrically.  Adult  males  of  the  hirtipes 
group  are  readily  distinguished  by  the  pres- 
ence of  a  patch  of  elevated  scales  on  the 
posterior  crus  and  thigh  of  each  hindleg 
(vinculae:  fide  H.  M.  Smith  and  R.  B. 
Smith  1980),  absent  in  turtles  of  the  scor- 
pioides group,  but  adult  females  of  the  hir- 
tipes group  lack  these  structures  and  are 
thus  often  difficult  to  identify.  An  elabora- 
tion of  the  differences  between  K.  integrum 
and  members  of  the  K.  hirtipes  group  is 
therefore  justified  (see  also  Iverson  and 
Berry  1979),  especially  since  they  coexist  in 
several  Mexican  drainage  basins  (see  MA- 
TERIALS AND  METHODS).  Kinosternon 
scorpioides  is  not  sympatric  with  members 
of  the  K.  hirtipes  group. 

The  primary  criteria  for  distinguishing 
adults  of  K.  integrum  and  the  K.  hirtipes 
group  appear  in  Table  1.  Juveniles  are 
much  more  difficult  to  distinguish  and 
remain  poorly  studied.  In  general,  small 
specimens  of  the  K  hirtipes  group  have 
smaller  plastra,  narrower  bridges,  and  more 
axillary-inguinal  scute  contact  than  K.  inte- 
grum (Fig.  1).  More  precise  discrimination 
of  small  turtles  must  await  additional  data. 

Members  of  the  K.  hirtipes  group  also 
differ  from  K.  integrum  ecologically.  The 
former  are  virtually  restricted  to  permanent 
water  habitats,  rarely  leaving  the  water 
except  to  nest;  migrating  behavior  is  unre- 
ported. Kinosternon  integrum  is  an  excel- 
lent colonizing  species  (fide  MacArthur  and 


Wilson,  1967).  It  is  extremely  vagile,  mi- 
grates considerable  distances  during  the 
rainy  season,  and  may  aestivate  under 
ground  as  K.  fiavescens  and  K.  alamosae 
do.  The  number  of  specimens  and  locality 
records  for  K.  integrum  in  museum  collec- 
tions (see  lists  in  H.  M.  Smith  and  R.  B. 
Smith,  1980)  reflects  the  more  frequent  oc- 
currence of  A",  integrum  than  K.  hirtipes  on 
roads.  Thus,  K.  integrum  may  be  found  in 
almost  any  temporary  pond  or  roadside 
pool,  habitats  where  K.  hirtipes  would  al- 
most never  occur. 

In  addition,  although  their  thermoregula- 
tory behavior  has  not  been  studied  in  detail, 
I  suspect  thermal  preference  and  tolerance 
levels  are  higher  in  K.  integrum  than  in  the 
K.  hirtipes  group.  This  is  reflected  in  the 
very  different  basking  habits  of  the  two 
forms.  I  have  observed  K.  integrum  basking 
at  many  Mexican  localities  in  Michoacdn, 
Jalisco,  Sinaloa,  and  Oaxaca,  but  K.  hir- 
tipes basking  only  once  at  2400  m  elevation 
in  Durango,  and  once  (adult  females  only) 
at  1800  m  in  Jalisco.  This  perhaps  reflects 
their  coastal  lowland  (integrum)  versus  high 
plateau  (hirtipes)  origins.  The  absence  of  A'. 
integrum  in  the  highest  (i.e.  coldest)  basins 
of  the  southern  Mexico  Plateau  (Pa'tzcuaro, 
San  Juanico,  and  Villa  Victoria;  see  later) 
may  be  related  to  thermal  requirements 
rather  than  to  historical  zoogeography. 
Likewise,  despite  its  vagility,  K.  integrum 
ranges  no  farther  northward  in  Sonora  than 
29 °N  latitude  in  the  Rio  Yaqui  basin.  Per- 
haps its  range  there  is  also  limited  by  tem- 
perature regime.  Further  study  may  show 
other  behavioral  and  ecological  differences 
between  these  two  groups  on  the  Mexican 
Plateau. 

Materials  And  Methods 

Specimens  and  field  work 

I  have  examined  nearly  all  specimens  of 
the  species  of  the  Kinosternon  hirtipes 
group  in  United  States  museums.  In  addi- 
tion, most  of  the  world's  museums  were 
canvassed  for  locality  data  of  other  speci- 
mens. All  available  type  specimens  were 
examined.  Each  locahty  was  pinpointed 
(and    its   elevation    determined)   on   the 


No.  1 


Kinosternon  Biosystematics 


Figure  1.  Plastral  comparison  of  juvenile  Kinoster- 
non hirtipes  (left;  UMBM  2403)  and  A:.  integrum 
(UMBM  2411),  both  from  3.2  km  SE  Ocotlan, 
Jalisco,  Mexico. 


1:500,000  sheets  of  "La  carta  general  de 
la  Repiiblica  Mexicana"  (published  by  the 
Ex-Comision  Intersecretarial  de  Mexico, 
D.  F.,  1958),  with  the  help  of  the  "Offi- 
cial Standard  (Geographic)  Names  of 
Mexico",  published  by  the  Office  of  Ge- 


ography of  the  U.S.  Dept.  of  Interior 
(1956).  These  localities  were  then  mapped 
on  Miller's  (1968)  drainage  map  of  Mex- 
ico (Figs.  3  and  4). 

Field  trips  to  sample  critical  areas  for 
K.  hirtipes  were  taken  in  May  1977  (12 
days;  10  localities  in  Durango  and 
Chihuahua),  June  1978  (11  days;  12 
localities  in  San  Luis  Potosi,  Guanajuato, 
Michoacan,  and  JaUsco),  June  1979  (part 
of  14  day  trip;  5  localities  in  Mexico 
State),  July  1980  (12  days;  7  localities  in 
Coahuila,  Chihuahua,  and  Durango),  and 
May  1981  (8  days;  10  localities  in  Jalisco, 
Mdxico  State,  Michoacan,  and  Puebla). 
Field  work  with  K.  sonoriense  in  Arizona 
was  also  undertaken  in  January  1971  (2 
days),  May  1974  (4  days),  January  1976  (4 
days),  and  July  1980  (1  day),  and  in  Chi- 
huahua in  August  1980  (1  day). 

All  known  specimens  and  localities  for 
members  of  the  K.  hirtipes  species  group 
are  in  the  SPECIMEN  LIST  and  plotted 
in  Figures  3  and  4;  museum  acronyms  fol- 
low Duellman,  Fritts,  and  Leviton  (1978) 
except  for  the  following: 


CAS-SU 

DMNH 

EAL 

ENMU 

FB 

FWMNH 

JBI 

JFB 

LTU 

MES 

MSU 

MU 

NMSU 

RSF 

SENCK 

SM 

SRSU 

TAI 

UAZ 

UF 

UG 

UMKC 

UNSM 

UOK 

USA 

USL 


California  Academy  of  Sciences  -  Stanford  University 

Collections 
Dallas  Museum  of  Natural  History 
Ernest  A.  Liner,  Houma,  Louisiana 
Eastern  New  Mexico  University 
Thomas  R.  VanDevender,  Tucson,  Arizona 
Fort  Worth  Museum  of  Natural  History 
John  B.  Iverson,  Richmond,  Indiana 
James  F.  Berry,  Elmhurst,  Illinois 
Louisiana  Tech  University 
Michael  E.  Seidel,  Huntington,  West  Virginia 
Michigan  State  University 
Midwestern  University,  Wichita  Falls,  Texas 
New  Mexico  State  University 
R.  S.  Funk,  Normal,  Illinois 
Senckenberg  Museum 
Strecker  Museum,  Baylor  University 
Sul  Ross  State  University 
Texas  A&I  University 
University  of  Arizona 

University  of  Florida,  Florida  State  Museum 
University  of  Georgia 
University  of  Missouri,  Kansas  City 
University  of  Nebraska  State  Museum 
University  of  Oklahoma 
University  of  South  Alabama 
University  of  Southwestern  Louisiana 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


This  study  is  based  on  the  examination 
of  at  least  1298  museum  specimens  of  the 
Kinosternon  hirtipes  species  group,  as 
well  as  other  specimens  collected  and  re- 
leased in  the  field.  Population  samples  of 
turtles  correspond  to  the  inhabited  drain- 
age basins,  which  are  listed  and  described 
below  from  approximately  north  to 
south.  The  reader  is  referred  to  Blasques 
L.  (1959)  and  Tamayo  (1962,  1964)  for 
more  general  descriptions  of  the  geogra- 
phy and  hydrography  of  the  drainage 
basins  in  Mexico. 

Bill  Williams  River  basin,  Arizona 
(WILL).  —  The  Bill  Williams  River  and 
its  major  tributaries,  the  Big  Sandy,  Bur- 
ro, and  Santa  Maria  rivers  drain  a  small 
area  in  west  central  Arizona  and  empty 
into  the  Colorado  River  at  Parker  Dam, 
about  90  km  below  the  Nevada  border. 
Kinosternon  sonoriense  is  the  only  fresh- 
water turtle  known  from  this  basin  (four 
localities;  800-1200  m).  Stebbins'  (1966) 
K.  flavescens  records  from  this  basin  were 
based  on  K.  sonoriense  (Iverson,  1978: 
477). 

Gila  and  Lower  Colorado  River  basins, 
Arizona,  California,  New  Mexico,  and 
Sonora  (GILA).  —  Most  of  Arizona 
south  of  the  Mogollon  Rim  and  a  portion 
of  west  central  New  Mexico  are  drained 
by  this  system.  Miller  (1961),  Ohmart,  et 
al.  (1975),  and  McNatt  (1978)  described 
changes  in  the  aquatic  habitats  along  the 
Colorado  and  Gila  rivers  and  their  tribu- 
taries over  the  past  100  years.  Kinoster- 
non sonoriense  occurs  throughout  the 
basin  (Iverson,  1976,  1978)  and  reaches  its 
maximum  known  elevation  (2042  m)  in 
the  Gila  River  in  western  New  Mexico 
(Niles,  1962;  Degenhardt  and  Christian- 
sen, 1974),  and  its  lowest  known  elevation 
(ca.  43  m)  near  Yuma,  Arizona. 

The  range  of  K.  sonoriense  in  the  Colo- 
rado River  is  poorly  known.  It  apparently 
once  occurred  there  at  least  upstream  to 
southeastern  Nevada  (LaRivers,  1942,  as 
K.  flavescens;  see  Iverson  1978:476). 
More  field  work  is  needed  along  the  Colo- 
rado River  between  Needles,  California, 
and  Yuma,  Arizona,  to  establish  the  pre- 


sent range  of  K.  sonoriense.  The  only 
other  freshwater  turtle  which  may  occur 
naturally  in  this  basin  is  K.  flavescens,  but 
I  have  elsewhere  (Iverson,  1978)  ques- 
tioned its  recent  occurrence  in  the  Gila. 
The  introduced  Trionyx  spinferus  does, 
however,  also  occur  in  the  Gila  and  Colo- 
rado rivers  from  southwestern  Utah  and 
western  New  Mexico  to  the  mouth  of  the 
Colorado  River  (Webb,  1973). 

Southwestern  New  Mexico  interior 
drainages  (SWNM).  —  K.  sonoriense 
occurs  in  the  permanent  water  basins  of 
the  eastern  and  western  slopes  of  the  Pe- 
loncillo  Mountains  of  Hidalgo  Co.  in 
southwestern  New  Mexico  and  adjacent 
Arizona  from  1150  m  to  1700  m  (Niles, 
1962,  and  Degenhardt  and  Christiansen, 
1974,  briefly  discussed  turtle  habitats  in 


f^^\ 


Figure  2.  Comparison  of  papillae  on  the  tail  of 
female  Kinosternon  hirtipes  (above;  FMNH 
71029;  137  mm  CL;  Guanajuato,  Taboado)  and 
A',  integrum  (FMNH  71031;  126  mm  CL;  same 
locality). 


No.  1 


Kinosternon  Biosystematics 


Table  1 .  Character  Comparison  of  adults  of  the  Kinosternon  hirtipes  species  group  and 
Kinosternon  integrum  (K.  scorpioides  species  group). 


Kinosternon  hirtipes 
group 


Kinosternon  integrum 


Elevated  scale  patches  on  hind  legs  of 
males 

Head  shield  in  adults  geographically 
variable,  from  reduced  crescent-shape 
to  large,  V-shape,  or  to  even  larger  tri- 
angle or  bell  shape  (latter  characteris- 
tic only  of  A",  sonoriense  and  Valley  of 
Mexico  K.  hirtipes);  posterior  margin 
of  shield  often  concave. 

Axillary  and  inguinal  scutes  nearly  al- 
ways in  broad  contact. 

Plastron  usually  yellow  or  greenish  yel- 
low, sometimes  darkly  stained. 

If  carination  present  on  posterior  cara- 
pace, then  only  one  medial  keel  usual- 
ly evident. 

Skin  very  papillose;  tail  with  numerous 
rows  of  large  papillae  (Fig.  2). 

Maximum  carapace  length,  185  mm; 
maximum  plastron  length,   160  mm. 

First  vertebral  scute  width  averages  24.5 
(range  20  to  3207o)  and  25.1%  (20  to 
30%)  of  carapace  length  in  male  A". 
hirtipes  and  K.  sonoriense,  respective- 
ly; 24.7  (18  to  31%)  and  26.1%  (20  to 
32%)  in  females,  respectively. 

Bridge  length  averages  20.1  (range  16  to 
24%)  and  21.4%  (18  to  25%)  of  cara- 
pace length  in  male  K.  hirtipes  and  K. 
sonoriense,  respectively;  23.6  (18  to 
29%)  and  24.8%  (22  to  28%)  in  fe- 
males, respectively. 

Bridge  length  averages  82.0%  (range  61 
to  120%)  and  85.3%  (62  to  115%)  of 
first  vertebral  scute  width  in  male  K. 
hirtipes  and  K.  sonoriense,  respective- 
ly; 95.5%  (64  to  133%)  and  95.0%  (70 
to  123%)  in  females,  respectively. 

Maximum  posterior  width  of  plastral 
forelobe  averages  43%  (range  36  to 
51%)  and  47%  (range  42  to  53%)  of 
carapace  length  in  male  K.  hirtipes 
and  K.  sonoriense,  respectively;  48% 
(42  to  54%)  and  49%  (44  to  54%)  in 
females,  respectively. 

Nuchal  and  first  neural  bones  occasion- 
ally (41%  in  K.  sonoriense;  10%  in  K. 
hirtipes)  in  contact. 


No  elevated  scale  patches  on  hind  legs  of 
males 

Adult  head  shield  large,  triangular  or 
bell  shaped,  with  posterior  margin 
convex;  shield  not  reduced  or  furcate 
behind 


Axillary  and  inguinal  scutes  usually  not 
in  contact;  if  touching,  contact  is  nar- 
row. 

Plastron  usually  yellow-orange,  almost 
never  darkly  stained. 

If  carination  present  on  posterior  cara- 
pace, then  three  keels  usually  evident. 

Skin  hardly  papillose;  tail  with  few,  very 
reduced  papillae  (Fig.  2). 

Maximum  carapace  length,  210 -(-  mm; 
maximum  plastron  length,  195-1-  mm. 

First  vertebral  scute  width  averages 
21.3%  (range  17  to  26%)  of  carapace 
length  in  males;  22.5%  (19  to  28%  in 
females). 


Bridge  length  averages  26.1%  (range  20 
to  28%)  of  carapace  length  in  males; 
26.8%  (20  to  30%)  in  females  (ex- 
cludes coastal  Jalisco  specimens). 


Bridge  length  averages  114%  (range  88 
to  151%)  of  first  vertebral  scute  width 
in  males;  115%  (91  to  158%)  in  fe- 
males (excludes  coastal  Jalisco  speci- 
mens). 

Maximum  posterior  width  of  plastral 
forelobe  averages  47%  (range  42  to 
54%)  of  carapace  length  in  males; 
53%  (45  to  57%)  in  females  (excludes 
coastal  Jalisco  specimens). 


Nuchal  and   first  neural  bones  not  in 
contact. 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


this  area).  Huntington  (1914:70)  reviewed 
the  historical  isolation  of  the  Animas 
Valley,  which  lies  to  the  east  of  Peloncil- 
los  and  receives  the  drainages  of  the  east- 
ern slopes  of  those  mountains.  Hubbs  and 
Miller  (1948)  examined  the  geography  of 
this  and  other  independent  drainage  bas- 
ins in  southwestern  New  Mexico. 

Rio  Sonoyta  (-  Sonoita)  basin,  Arizo- 
na and  Sonora  (SNTA).  —  The  Rfo 
Sonoyta  lies  along  the  northwestern 
boundary  of  the  state  of  Sonora,  Mexico. 
The  river  disappears  in  the  desert  sands 
near  the  eastern  border  of  the  Pinacate 
lava  flows.  The  physical  geography  of  the 
basin  is  reviewed  by  Ives  (1936).  K. 
sonoriense  is  found  in  the  more  perma- 
nent portions  of  the  basin  between  about 
350  and  450  m,  near  the  U.S.  border.  It  is 
abundant  at  Quitobaquito  Pond  in  Organ 


Figure  3.  Distribution  of  the  subspecies  of  Kino- 
sternon  sonoriense.  Dots  indicate  actual  records; 
hatching,  suggested  total  ranges.  The  range  of  K. 
sonoriense  longifemorale  is  marked  (A);  the  re- 
maining hatched  area  marks  the  range  of  the 
nominate  subspecies.  Question  mark  in  Nevada  is 
discussed  in  Iverson  (1978);  that  in  southeastern 
Chihuahua,  in  the  present  text.  Stippled  area  il- 
lustrates portion  of  the  allopatric  range  of 
Kinosternon  hirtipes  (see  Fig.  4). 


Pipe  Cactus  National  Monument,  Pima 
Co.,  Arizona  (Hulse,  1974;  Iverson,  field 
notes).  The  aquatic  habitat  at  Quitoba- 
quito was  described  by  Cole  and 
Whiteside  (1965).  Hubbs  and  Miller 
(1948:113)  discussed  the  historical 
geography  of  the  basin. 

Kinosternon  flavescens  is  the  only  other 
native  aquatic  or  semiaquatic  turtle 
known  from  the  Rio  Sonoyta  system  (H. 
M.  Smith  and  Hensley,  1957;  Iverson, 
1979),  but  Hulse  (1974:94)  reported  that 
Chrysemys  picta  dorsalis  has  been  intro- 
duced into  Quitobaquito  Spring. 


Figure  4.  Distribution  of  the  subspecies  of  Kinoster- 
non hirtipes.  Dots  indicate  actual  records;  hatch- 
ings, suggested  total  ranges.  Subspecies  ranges 
are  marked:  K.  hirtipes  murrayi  (A-E);  K.  h. 
chapalaense  (F);  K.  h.  chapalaense  x  A",  h. 
murrayi  (G);  A",  h.  magdalense  (H);  K.  h. 
tarascense  (I);  K.  h.  megacephalurn  (J),  and  A",  h. 
hirtipes  (K).  Problematical  localities  (San  Luis 
Potosi,  C;  Balsas,  D;  and  Puebla,  E)  are 
discussed  in  text.  Stippled  area  indicates  a  portion 
of  the  allopatric  range  of  K.  sonoriense  (see  Fig. 
3). 


No.  1 


Kinosternon  Biosystematics 


Rio  Magdalena,  Sonora  (MAGD).  — 
The  Rio  Concepcidn  (the  major  tributary 
of  the  Rio  Magdalena)  arises  in  the  hills 
near  Nogales,  Arizona,  flows  as  a  perma- 
nent stream  through  the  Magdalena  Val- 
ley, and  disappears  into  the  coastal  sands 
of  northwestern  Sonora  (Tamayo  1964: 
102).  Kinosternon  sonoriense  is  known 
from  numerous  permanent  water  habitats 
between  about  300  and  1200  m  elevation. 
Kinosternon  flavescens  is  also  known 
from  this  basin  (Iverson,  1979). 

Rio  Sonora,  Arizona  and  Sonora 
(SNRA).  —  Like  the  Rfo  Magdalena,  the 
Rio  Sonora  rises  near  the  Arizona-Sonora 
border  and  disappears  (below  Hermosillo) 
into  coastal  sands  (Tamayo  1964:102). 
Kinosternon  sonoriense  is  locally  very 
abundant  in  this  basin  in  permanent  water 
habitants  between  at  least  200  and  1200  m 
elevation.  Kinosternon  flavescens  is  the 
only  other  freshwater  turtle  known  from 
this  basin  (Iverson,  1979). 

Rio  Yaqui  basin  (excluding  the  Papigo- 
chic  drainage),  Arizona,  Chihuahua,  and 
Sonora  (YAQ).  —  Because  of  the  zoo- 
geographic  dissimilarity  of  the  Yaqui 
basin  west  of  the  Continental  Divide  (Rios 
Yaqui,  Moctuzuma,  Bavispe,  and  Aros) 
and  those  east  of  the  Divide  (Rios  Papigo- 
chic  and  Tomochic)  (Meek,  1904;  Miller, 
1958),  and  because  obvious  differences 
were  observed  early  in  the  study  between 
the  turtles  of  the  K.  hirtipes  group  on 
either  side  of  that  Divide  in  those  rivers 
{Kinosternon  sonoriense  in  streams  to  the 
west,  K.  hirtipes  in  those  to  the  east),  the 
Yaqui  sample  was  divided  into  a  Plateau 
portion  (hereafter  called  the  Rfo  Papi- 
gochic  sample)  and  a  non-Plateau  portion 
(hereafter  restrictively  called  the  Rfo 
Yaqui  sample).  Kinosternon  sonoriense  is 
known  only  from  headwater  populations 
in  permanent  water  situations  between 
1200  and  2000  m  elevation  in  southeastern 
Arizona  and  adjacent  northeastern  Son- 
ora, and  western  Sonora  near  the  Chihua- 
hua border.  A  specimen  from  El  Novillo, 
Sonora,  catalogued  as  A',  sonoriense 
(UAZ   36505)   but    unseen    by   me   (not 


mapped  on  Fig.  3,  but  mapped  in  Iverson, 
1976)  is  probably  misidentified  since  it  is 
the  only  record  of  K.  sonoriense  in  the 
lower  Ri'o  Yaqui.  Additional  field  work  in 
this  basin  is  badly  needed. 

The  freshwater  turtles,  Kinosternon 
integrum,  K.  flavescens,  K.  alamosae, 
and  Pseudemys  scripta  also  occur  in  the 
Yaqui  basin  (Legler  and  Webb,  1970; 
Iverson,  1978,  1979;  Berry  and  Legler, 
1980),  but  the  microsympatry  of  any  pair 
of  species  in  the  Yaqui  basin  has  not  been 
established 

Rio  Fuerte  basin.  Chihuahua  and 
Sinaloa.  —  Three  specimens  of  Kinoster- 
non sonoriense  (identification  verified) 
collected  by  Wilmer  Tanner  bear  the  lo- 
cality data  "Cerocahui,  Chihuahua" 
(question  mark  in  Fig.  3).  As  mapped  by 
Tanner  and  Robison  (1960),  and  Conant 
(1978:466),  the  locality  Hes  along  a  tribu- 
tary of  the  Rio  Fuerte  (Pacific  drainage). 
No  other  members  of  the  K.  hirtipes 
group  are  known  from  this  basin,  whereas 
K.  integrum  is  abundant  at  lower  eleva- 
tions (Berry,  1978;  Iverson,  unpublished). 
Contreras-Balderas  (1975)  suggested  that 
the  fish  Notropis  (Codoma)  ornatus  (pri- 
marily an  inhabitant  of  the  Mexican  Pla- 
teau) may  also  inhabit  the  headwaters  of 
the  Rfo  Fuerte.  This  would  indicate  hist- 
orical faunal  interchange  (perhaps  stream 
capture)  between  the  Plateau  and  the 
upper  Rfo  Fuerte,  and  might  have  permit- 
ted K.  hirtipes,  but  not  K.  sonoriense,  to 
reach  the  Fuerte.  The  Cerocahui  locality 
must  therefore  remain  problematical  until 
this  rarely  visited  area  in  southwestern 
Chihuahua  is  better  studied. 

Rio  Casas  Grandes  Interior  Basin,  Chi- 
huahua (CSGR).  —  The  headwaters  of 
the  Rio  Casas  Grandes  are  in  the  Sierra 
Madre  Occidental,  very  close  (25  km 
radius)  to  the  headwaters  of  the  Rfo 
Bavispe  (Yaqui)  and  Ri'o  Papigochic 
basins.  In  fact,  headwater  streams  of  the 
Bavispe  and  Casas  Grandes  reach  within  6 
km  of  one  another  at  about  2000  m, 
southwest  of  Pacheco,  Chihuahua.  The 
basins  are  there  separated  by  a  divide  less 
than  200  meters  high. 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Tamayo  (1962:475)  provided  a  photo- 
graph of  the  Rio  Casas  Grandes,  presum- 
ably south  of  the  town  of  that  name,  and 
Goldman  ( 1 95 1 : 1 1 9- 1 22)  describes  several 
habitats  in  this  basin.  Below  (north)  the 
town  of  Nueva  Casas  Grandes,  the  river  is 
diverted  for  agricultural  purposes  and  can 
at  best  be  called  intermittent.  The  river 
terminates  in  Laguna  Guzman  (1180  m; 
photographs  in  Henrickson,  1977)  but 
seldom  (only  during  floods)  does  that 
Laguna  receive  water  via  the  Rfo  Casas 
Grandes.  K.  sonoriense  apparently  occurs 
only  above  (south  oO  Nueva  Casas 
Grandes  (1475  m),  up  to  an  elevation  of 
between  1500  and  1600  m  in  the  Rio 
Piedras  Verdes  near  Colonia  Juarez. 

My  trapping  operations  were  unsuccess- 
ful on  13  May  1977  in  the  main  channel  of 
the  Rfo  Casas  Grandes  at  a  bridge  on 
Highway  2  between  Janos  (ca.  10  km  N) 
and  Ascension;  only  catfish  (Ictalurus  sp.) 
were  trapped  in  apparently  permanent 
pools  even  though  locals  told  me  that 
"rock"  turtles  lived  in  the  stream.  Inde- 
pendently, Conant  (1978:488)  took  only 
catfish  and  bullfrogs,  and  collected  no 
turtles  in  his  traps  during  two  days  of  field 
work  at  the  same  site. 

On  1  August  1980,  the  Rio  Piedras 
Verdes  above  Colonia  Juarez  was  very 
shallow  (average  depth,  10-20  cm;  maxi- 
mum depth,  0.5  m)  and  slowly  moving.  In 
one  hour,  two  K.  sonoriense  were  collect- 
ed by  hand  in  shallow  water  and  two  more 
were  taken  in  traps  set  in  the  deepest 
areas.  Roger  Conant  (pers.  comm.)  trap- 
ped eight  K.  sonoriense  near  this  same  lo- 
cality on  19  August  1974  when  the  river 
was  in  flood. 

No  other  aquatic  turtles  are  known 
from  the  Casas  Grandes  basin;  Van  De- 
vender  and  Van  Devender's  (1975)  Chrys- 
emys  picta  record  was  based  on  specimens 
actually  from  the  Rio  Santa  Maria  basin. 

Rio  Santa  Maria  interior  drainage,  Chi- 
huahua (STMR).  —  The  Rio  Santa  Maria 
rises  in  the  Sierra  Madre  Occidental  very 
close  to  the  Rfo  Papigochic  basin,  and 
flows  northward  across  the  desert  floor  in 
northwestern  Chihuahua.  The  river  bed 


terminates  in  Laguna  de  Santa  Maria 
(1172  m),  and  like  Laguna  Guzman  sel- 
dom receives  water  from  its  confluent 
stream.  The  Laguna  de  Santa  Marfa 
(photographed  in  Henrickson,  1978)  is 
separated  from  the  Laguna  de  Guzman  by 
a  divide  of  not  more  than  61  m  elevation 
(Goldman,  1951:123).  Geological  evi- 
dence suggests  the  lakes  were  continuous 
during  Wisconsin  time  when  the  water 
level  reached  at  least  1225  m  (Axtell  1978: 
509). 

Kinosternon  hirtipes  reaches  its  north- 
ernmost distribution  in  the  Rio  Santa 
Maria,  and  is  common  in  permanent  wa- 
ter situations  throughout  the  basin  be- 
tween at  least  1400  and  1600  m  elevation. 
On  12  May  1977,  in  a  tributary  of  the 
Rio  Santa  Maria,  southeast  of  Galeana, 
two  assistants  and  I  captured  nearly  100 
individuals  of  K.  hirtipes  by  hand  (most 
of  which  were  subsequently  released)  in 
less  than  two  hours,  primarily  by  feeling 
under  stream  banks.  Seven  man  hours  of 
hand  collecting  on  2  August  1980  at  the 
same  locality  produced  140  turtles,  which 
were  measured,  marked  and  released  as 
part  of  an  ecology  study.  Habitats  near 
that  location  were  described  by  Van 
Devender  and  Van  Devender  (1975). 
Semmler  et  al.  (1977)  reported  similar  suc- 
cess in  the  Rio  Santa  Maria,  also  near 
Galeana. 

Both  K.  flavescens  (Iverson,  1979)  and 
Chrysemys  picta  (H.  M.  Smith  and 
Taylor,  1950a;  Iverson,  field  notes;  Roger 
Conant,  pers.  comm.)  also  occur  in  the 
Santa  Maria  basin. 

Rio  Carmen  (^  Rio  Santa  Clara)  interi- 
or drainage,  Chihuahua  (CRMN).  —  The 
headwaters  of  the  Rfo  Carmen  lie  in  the 
Sierra  Madre  west  of  the  Sierra  del  Nido. 
The  river  once  flowed  (at  least  during 
floods)  to  the  Lago  de  Patos  (1175  m) 
near  Villa  Ahumada,  but  since  the  con- 
struction of  a  dam  (Presa  de  Las  Lajas) 
just  south  of  Ricardo  Flores  Magon  ( =  El 
Carmen)  the  river  no  longer  flows  south 
past  Ricardo  Flores  except  in  cement 
irrigation  flumes.  When  visited  on  11-12 
May  1977,  the  remnants  of  the  riparian 


No. 


Kinosternon  Biosystematics 


woodland  below  the  dam  were  still  in  evi- 
dene,  but  rapidly  disappearing  (see  also 
Conant  1977:488).  Sixty  trap  hours  along 
the  shores  of  the  Presa  yielded  no  turtles. 
The  rocky  shoreline  lacks  aquatic  vegeta- 
tion and  the  continually  changing  shore- 
line (evident  from  water  marks  on  the 
rocks)  presented  habitats  which  were  un- 
doubtedly generally  unsuitable  for  Kino- 
sternon turtles.  However,  Conant  (1978: 
473)  indicated  that  he  recently  obtained 
K.  hirtipes  in  the  impoundment. 

Records  for  K.  hirtipes  are  available 
from  the  Rfo  Carmen  below  (north  of)  the 
dam  at  Ricardo  Flores  (15(X)-1600  m),  up 
(South)  to  the  region  near  Santa  Clara 
(1800  m).  Kinosternon  flavescens  is  the 
only  other  aquatic  turtle  known  from  this 
basin  (Iverson,  1979). 

Rio  El  Sauze  (=  Encinillas)  interior 
drainage.  Chihuahua  (SAUZ).  —  The 
Rfo  El  Sauz  rises  in  the  eastern  slopes  of 
the  Sierra  del  Nido,  and  flows  intermit- 
tently to  the  desert  floor  in  the  vicinity  of 
the  town  of  Sauz.  It  then  flows  intermit- 
tently northward,  paralleling  the  Sierra 
del  Nido,  until  it  disappears  into  the 
ground  about  100  km  north-northwest  of 
Ciudad  Chihuahua. 

According  to  Hubbs  (in  Hubbs  and 
Springer  1957:299;  and  in  Miller  1961: 
393)  the  entire  Sauz  Valley  went  dry  in 
1947;  however,  Minckley  and  Koehn 
(1965)  recorded  an  apparently  permanent, 
though  artificial  pond  (with  fishes)  in  the 
Sauz  Valley  in  1964  and  Contreras-Bal- 
deras  (1974:182)  reported  fish  collections 
made  in  1964  and  1968. 

Kinosternon  hirtipes  has  been  collected 
from  at  least  four  localities  in  the  Sauz 
Valley  between  1500  and  1700  m;  how- 
ever, we  trapped  none  at  either  of  two 
sites  of  apparently  permanent  water  (con- 
taining fishes)  near  Sauz  on  1 1  May  1977. 
Local  children  at  that  time  confirmed  the 
occurrence  of  turtles  in  the  stream,  but 
said  they  were  uncommon;  they  were  also 
unsuccessful  at  securing  any  for  us. 

The  only  other  aquatic  turtle  supposed- 
ly recorded  from  this  basin  is  Sterno- 
therus  odoratus,  but  the  single  record  re- 


mains problematical  (Moll  and  Williams, 
1963;  Conant  and  Berry,  1978).  Although 
unknown,  K.  flavescens  may  occur  in  this 
basin  (Iverson,  1979). 

Alamito  Creek  drainage,  Presidio  Co., 
Texas  (TEX).  —  Alamito  Creek  is  an 
ephemeral  tributary  of  the  Rio  Grande, 
east  and  north  of  Presidio,  Texas.  K.  hir- 
tipes is  known  from  only  two  permanent 
ponds  in  this  drainage  in  Texas  (about 
1050  m).  Conant  and  Berry  (1978:11-15) 
elaborated  on  the  specific  localities  and 
field  work  in  the  area.  This  species  has  not 
been  found  in  the  Rio  Grande  itself  or  its 
American  tributaries  in  southwest  Texas, 
but  populations  may  reside  in  permanent 
Mexican  tributaries  between  Presidio  (or 
Ojinaga,  Chihuahua)  and  the  Big  Bend  of 
the  Rio  Grande. 

Kinosternon  hirtipes  does  not  likely 
occur  today  above  Presidio  in  the  Rio 
Grande  since  even  by  1919  that  river  was 
sometimes  completely  dry  between  the 
New  Mexico  border  and  the  mouth  of  the 
Rfo  Conchos  (Udden,  Baker,  and  Bose 
1919:23). 

Kinosternon  flavescens  is  the  only  other 
aquatic  turtle  occurring  in  the  Alamito 
drainage  with  K.  hirtipes;  based  on 
museum  records,  the  two  species  occur 
microsympatrically  in  at  least  one  of  the 
spring-fed  ponds  in  this  basin  (Iverson, 
1979). 

Rio  Conchos  drainage.  Chihuahua  and 
Durango  (CNCH).  —  The  Rio  Conchos 
is  the  major  tributary  of  the  Rio  Grande 
(=  Rio  Bravo),  accounting  for  18%  of 
the  latter's  total  flow  (Tamayo  1964:89). 
With  its  headwaters  in  the  Sierra  Madre  in 
southwestern  Chihuahua  and  extreme 
northern  Durango,  it  drains  more  of  the 
state  of  Chihuahua  than  any  other  single 
river. 

More  specimens  of  K.  hirtipes  have 
been  collected  in  the  Rfo  Conchos  basin 
than  in  any  other  basin.  The  species  is 
known  from  the  mouth  of  the  Conchos 
(ca.  800  m;  the  lowest  altitudinal  record 
for  the  species)  near  Ojinaga  (Legler, 
1960)  to  its  more  accessible  headwaters  (in 
the  Rfo  Florido)  near  Las  Nieves  in  north- 


10 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


ern  Durango  (ca.  1800  m).  Bushnell 
(1971:332)  provided  a  photograph  of 
Lago  Boquillo  on  the  Rfo  Conchos,  a  lo- 
cality from  which  K.  hirtipes  is  known. 
Kinosternon  flavescens,  Pseudemys 
scripta,  and  Trionyx  spiniferus  are  also 
known  from  the  Conchos  basin  (Legler, 
1960;  H.  M.  Smith,  et  al.,  1963;  Webb, 
1973;  and  Iverson,  1979). 

Laguna  Bustillos  interior  drainage, 
Chihuahua  (BVST).  —  The  small  Laguna 
Bustillos  basin  (2720  square  km)  is  in  the 
foothills  of  the  Sierra  Madre  west  of  Ciu- 
dad  Chihuahua.  The  Laguna  itself  lies  at 
approximately  19(X)  m.  A  single  collection 
(UMMZ;  4  specimens)  of  K.  hirtipes  is 
available  from  a  tributary  27.4  km  north 
of  Ciudad  Cuauhtemoc.  I  trapped  along  a 
clear,  shallow,  permanent,  though  inter- 
mittently flowing  stream  with  an  adjacent 
cattle  tank  (pond)  24.3  km  north  of 
Cuauhte'moc  on  Hwy.  28  on  10  May  1977. 
These  two  localities  probably  are  the 
same,  since  I  could  find  no  other  tribu- 
taries on  the  road  north  from  Cuauhte'- 
moc. No  turtles  were  captured,  although 
the  microhabitat  seemed  adequate  in 
some  of  the  deeper  areas  (maximum,  0.5 
m).  Populations  of  K.  hirtipes  may  exist 
in  the  Laguna  itself  or  in  this  tributary 
where  it  leaves  the  mountains  to  the  west 
(and  thus  presumably  has  more  flow). 
Further  field  work  is  warranted  to  deter- 
mine if  K.  hirtipes  is  still  extant  in  the 
Bustillos  basin.  No  other  aquatic  turtle  is 
known  from  the  basin. 

Rio  Papigochic,  Chihuahua  (PAP).  — 
As  mentioned  above  in  regard  to  the  Rfo 
Yaqui  basin,  the  plateau  portion  of  that 
basin,  the  Rio  Papigochic,  is  here  consid- 
ered a  separate  sample  area.  The  Papigo- 
chic arises  in  the  Sierra  Madre  west  of 
Ciudad  Chihuahua  and  flows  northwest- 
ward to  the  vicinity  of  Yepdmera,  where  it 
turns  sharply  southward  for  about  25  air- 
line km  to  the  confluence  of  the  Rfo 
Tom6chic  and  another  sharp  turn  to  the 
northwest.  It  continues  in  that  direction 
for  almost  150  airHne  km  before  heading 
southwestward  to  its  confluence  with  the 
Rio  Aros  (tributary  to  the  Yaqui)  just  in- 


side the  Sonora  border.  Because  of  this 
anomalous  drainage  pattern  and  because 
the  faunal  affinities  of  the  Papigochic  are 
with  the  Rfo  Conchos  and  not  the  Rfo  Ya- 
qui (sensu  stricto),  zoogeographers 
believe  the  Rfo  Papigochic  was  until  re- 
cently (prehistorically)  a  tributary  of  the 
former  stream  (Meek,  1904;  Miller,  1958; 
among  others). 

Kinosternon  hirtipes  is  known  from 
both  the  Tomochic  and  Papigochic  from 
elevations  of  1200  to  at  least  2000  m.  Like 
Van  Devender  and  Lowe  (1977),  I  found 
K.  hirtipes  very  common  near  Yepomera. 
On  10  May  1977,  5  traps  set  in  a  broad, 
ponded  stretch  of  a  Papigochic  tributary 
yielded  25  turtles  in  two  hours.  No  other 
turtle  species  were  collected  (or  are  known 
from  the  basin),  but  Tom  Van  Devender 
(pers.  comm.)  reports  that  natives  near 
Yepdmera  told  him  of  a  "tortuga  pinta" 
that  lives  in  the  Papigochic  basin.  The  va- 
lidity of  the  report  and  the  identity  of  the 
turtle  (perhaps  Chrysemys  picta)  are  un- 
confirmed. 

Rio  Nazas  interior  drainage,  Durango 
and  Coahuila  (NAZ).  —  More  of  the 
state  of  Durango  is  drained  by  the  Rio 
Nazas  than  any  other  single  drainage  sys- 
tem. The  Nazas  rises  in  the  Sierra  Madre 
in  western  Durango  as  two  major  tribu- 
taries, the  north-flowing  Rio  Santiago 
and  the  south-flowing  Rfo  Tepehuanes. 
The  confluence  of  these  two  streams, 
about  20  airline  km  northwest  of  Santiago 
Papasquiaro  begins  the  Rio  de  Ramos. 
The  major  northern  tributary,  the  Rfo  del 
Oro  rises  in  the  Sierra  in  northwestern 
Durango  and  joins  the  Rfo  Ramos  near  El 
Palmito  below  which  it  is  called  the  Rfo 
Nazas.  The  Nazas  then  flows  eastward 
across  the  Chihuahuan  desert  (photo- 
graphed in  Spieth,  1950:34),  formerly  as 
far  as  the  Bdlson  de  Mayran  in  southwest- 
ern Coahuila.  Diversion  of  the  waters  for 
agriculture  near  Torredn  has,  however, 
severed  the  Nazas-Mayran  connection 
(Conant,  1963,  provides  an  excellent  de- 
scription of  the  topography  of  this  basin). 

Trapping  on  the  Rio  Nazas  west  of  Tor- 
redn (78  trap  hours,  5  -  6  May  1977,  3 


No.  I 


Kinosternon  Biosystematics 


11 


Kinosternon;  156  trap  hours,  23  -  24  July 
1980,  6  Kinosternon)  and  near  El  Palmito 
(98  trap  hours,  7-8  May,  1977,  5  Kinoster- 
non) by  me,  and  near  El  Palmito  (30  trap 
hours,  20-21  July  1976,  5  Kinosternon). 
Roger  Conant  (pers.  comm.)  indicated 
that  K.  hirtipes  is  uncommon  in  the  river 
itself.  The  absence  of  a  large  series  from 
any  one  locality  in  the  Nazas  basin  sup- 
ports this  statement.  The  species  is  known 
from  between  1100  (Lerdo)  and  1400  m 
(El  Palmito). 

Pseudemys  scripta  was  collected  at  all 
three  locations  I  visited  and  that  species 
probably  is  sympatric  with  K.  hirtipes  in 
most  of  the  Nazas  drainage.  K.  flavescens 
is  the  only  other  aquatic  turtle  known 
from  this  basin  (Iverson,  1979). 

Viesca  interior  basin,  Coahuila 
(VSCA).  —  Only  a  single  collection  of  A". 
hirtipes  (and  Pseudemys  scripta)  is  known 
from  the  small  area  south  of  the  city  of 
Viesca,  Coahuila  (symbol  J  in  Fig.  4). 
Bryce  Brown  seined  8  Kinosternon  and  2 
Pseudemys  scripta  on  4  June  1961  from  2 
drying  ponds  south  of  Viesca  at  about 
1 100  m.  Natives  informed  Brown  that  the 
ponds  were  remnants  of  a  once  active  hot 
spring  (pers.  comm.).  At  my  suggestion, 
ichthyologist  Robert  Rush  Miller  visited 
the  Viesca  area  in  the  spring  of  1978  and 
verified  (pers.  comm.)  the  fact  that  a 
spring  did  once  exist  along  the  mountains 
southwest  of  the  city,  but  he  could  find  no 
permanent  aquatic  habitats  suitable  for 
turtles  or  native  fishes. 

We  visited  the  area  on  23  July  1980  and 
an  elderly  Viesca  resident  showed  us  the 
locations  of  the  extinct  springs  (8  total;  1 
hot)  and  confirmed  the  lack  of  permanent 
surface  water  today.  He  told  us  that  the 
springs  had  gone  dry  "about  20  years 
ago"  but  that  prior  to  that  time  there  had 
been  much  water,  with  many  turtles, 
snakes,  and  fish.  The  only  permanent 
water  near  Viesca  of  which  anyone  there 
knew  was  a  spring  near  the  small  town  of 
La  Pena,  about  20  miles  to  the  East.  We 
visited  that  spring  on  23  July  1980  (as 
Miller  did  in  1978),  found  only  introduced 
fish  and  no  Kinosternon,  and  failed  to 


locate  the  Pseudemys  script a-WV.^  turtles 
locals  told  us  "used  to  occur"  in  the  two 
imy  presas  that  remain.  The  Viesca  turtle 
populations  must  therefore  be  considered 
extinct. 

The  specimens  of  Pseudemys  scripta 
from  near  Viesca  are  very  similar  to  those 
in  the  Rio  Nazas  immediately  to  the  west. 
The  Kinosternon,  however,  show  little  af- 
finity with  Nazas  specimens  and  in  fact 
have  the  most  unique  morphology  of  any 
member  of  the  hirtipes  group  (see  later). 

Rio  Aguanaval  interior  drainage,  Coa- 
huila, Durango  and  Zacatecas  (AGUN). 
—  The  Rio  Aguanaval  rises  on  the  Mex- 
ican Plateau  in  the  mountains  northwest 
of  the  city  of  Zacatecas  and  flows  inter- 
mittently northward  across  the  Chihua- 
huan  Desert.  It  once  emptied  into  the  La- 
guna  de  Viesca  in  southwestern  Coahuila, 
before  its  diversion  for  agricultural  pur- 
poses (Conant  1963,  1969). 

K.  hirtipes  is  the  only  turtle  known  to 
occur  in  the  Aguanaval  basin  and  has 
been  collected  only  in  the  headwaters  im- 
mediately northwest  of  Fresnillo  between 
2000  and  2200  m.  Natrix  erythrogaster 
alta,  endemic  to  the  Aguanaval  system,  is 
likewise  known  only  from  the  headwaters 
(Conant,  1969:46). 

Laguna  de  Santiaguillo  interior  drain- 
age, Durango  (STGO).  —  The  Laguna 
Santiaguillo  is  isolated  at  just  under  2000 
m  in  the  Sierra  Madre  northwest  of 
Canatlan,  Durango.  Only  two  collections 
(four  specimens)  of  K.  hirtipes  have  been 
made  in  the  Santiaguillo  basin,  both  ap- 
parently from  the  same  tributary  to  the 
Laguna  near  the  village  of  Guatimape.  On 
7  May  1977,  this  tributary  was  temporari- 
ly reduced  to  isolated  pools  (maximum 
depth  1  m)  in  the  stream  bed.  Trapping 
and  seining  produced  three  K.  hirtipes.  A 
presa  with  permanent  abundant  water  was 
subsequently  located  about  0.7  km  up- 
stream from  the  highway  bridge,  but  was 
not  sampled;  it  probably  supports  a  good 
population  of  A',  hirtipes.  No  other  turtle 
is  known  from  the  basin. 

Rio  Mezquital  drainage,  Durango 
(MEZ).  —  The  Rk)  Mezquital  is  the  ma- 


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Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


jor  tributary  of  the  Rio  San  Pedro  (Pacif- 
ic drainage),  and  drains  that  portion  of 
the  Mexican  Plateau  near  Ciudad  Du- 
rango,  Durango  (Albritton  1958:  Conant 
1963).  Both  K.  hirtipes  and  K.  integrum 
occur  in  the  Plateau  portion  of  the  Rio 
Mezquital,  but  only  AT.  integrum  is  known 
from  the  Pacific  coastal  plain  portion  of 
the  Mezquital-San  Pedro  system  (Iverson, 
unpublished).  Because  of  the  inaccessibili- 
ty of  the  area  south  and  southeast  of  Ciu- 
dad Durango,  the  lower  Hmit  of  the  range 
of  K.  hirtipes  in  the  Mezquital  is  uncer- 
tain. The  locality  farthest  downstream  is 
at  Mezquital  on  the  Rio  Mezquital,  south- 
east of  Ciudad  Durango  at  about  1100  m. 
The  species  reaches  its  highest  known  ele- 
vation in  this  basin  (2600  m)  at  Otinapa. 
Conant  (1978:467,  473)  correctly  ques- 
tioned the  record  (H.  M.  Smith  and  Tay- 
lor, 1950a:26)  of  K.  sonoriense  from 
Durango,  Durango  (this  basin);  it  was  ap- 
parently based  on  a  specimen  of  K.  hir- 
tipes. 

The  Rfo  Mezquital  and  its  Plateau  trib- 
utaries near  Ciudad  Durango  contain 
much  permanent  water  even  at  the  end  of 
the  dry  season.  On  6  May  1977  at  the  Ri'o 
La  Sauceda  (tributary  to  the  Mezquital) 
bridge  on  Hwy  40  (Figure  5),  60  K.  hir- 
tipes (pre-dominately  juveniles  and  sub- 
adults)  entered  11  traps  in  just  three 
hours.  The  most  productive  traps  had 
been  set  along  steep,  undercut  dirt  banks; 
traps  in  areas  of  gently  sloping  shoreHnes 
were  unproductive.  Six  traps  set  for  45 
minutes  at  the  same  locality  on  25  July 
1980  produced  14  K  hirtipes.  Based  on 
museum  records,  K.  hirtipes  is  very 
common  in  the  Mezquital  system  (148 
specimens),  whereas  K.  integrum  is  un- 
common (1  am  aware  of  but  14  specimens); 
a  single  collection  of  turtles  from  0.8  km 
N  Graceros  contains  12  K.  hirtipes  (KU 
68733-36,  68738-45)  and  a  single  K. 
integrum  (KU  68737). 

El  Salto  area,  Rio  Acaponeta  basin, 
Durango  (SALT).  -  K.  hirtipes  appar- 
ently occurs  in  the  Rfo  Acaponeta  only  in 
its  headwaters  northeast  of  El  Saho 
(Symbol  B  in  Fig.  4).  On  25  July  1980,  at 


a  shallow  (maximum  depth  0.75  m)  ap- 
parently permanent  stream  9.7  road  km 
ENE  El  Salto  (about  2400  m)  in  a  moun- 
tain meadow  surrounded  by  pine-oak 
woodland,  six  K.  hirtipes  were  collected  in 
12  trap  hours  and  three  more  were  taken 
by  hand.  K.  integrum  is  very  common  in 
the  lower  Rio  Acaponeta  basin  (Berry 
1978;  Iverson,  unpublished),  but  is 
unknown  from  the  headwaters  region. 

Southwestern  San  Luis  Potosi 
(interior?)  basin  (SLP).  -  In  an  isolated 
portion  of  the  Rio  Santa  Maria  drainage 
basin  (Panuco,  i.e.,  Atlantic  drainage 
southwest  of  Villa  de  Reyes  (symbol  C  in 
Fig.  4),  K.  hirtipes  and  K.  integrum  co- 
occur  abundantly.  Iverson  and  Berry 
(1979)  argued  that  this  population  of  K. 
hirtipes  is  the  resuh  of  an  introduction.  I 
continue  to  support  that  view,  especially 
since  K.  integrum  has  been  collected  at 
numerous  localities  in  the  Rfo  Santa 
Maria  system  (see  list  in  Iverson  and 
Berry,  1979),  yet  K.  hirtipes  is  known  only 
from  the  Laguna  de  las  Rusias  (  =  Presa 
El  Refugio;  ca.  1900  m)  area.  As  de- 
scribed by  Iverson  and  Berry  (1979:320) 
the  only  remaining  aquatic  habitat  found 
on  11  June  1978  was  a  small  permanent 
stream  that  was  diverted  entirely  for 
agriculture  within  2  km  below  the  broken 
dam.  The  Arroyo  below  the  dam  was 
lined  with  seepage  springs  and  quaking 
ground.  The  stream  varied  from  one  to 
four  m  wide  (x  ^  1)  and  averaged  only 
0.25  m  deep  (over  a  soft  mud  bottom  at 
least  1  m  deep).  The  water  was  quite  clear 
along  most  of  its  length,  but  odor  and 


Figure  5.  Rio  La  Sauceda  (Rio  Mezquital  basin)  at 
Highway  40  near  city  of  Durango,  6  May  1977. 


No.  1 


Kinosternon  Biosystematics 


13 


refuse  in  the  water  indicated  its  use  as  a 
human  sewage  effluent.  Kinosternon  was 
abundant  in  the  stream  on  11  June;  13  K. 
hirtipes  and  3  K.  integrum  were  dipnetted 
or  trapped  in  one  hour. 

Rio  Aguascalientes  drainage,  Aguas- 
calientes  (AGUAS).  -  Because  the 
distance  between  the  two  clusters  of 
localities  for  K.  hirtipes  in  the  Rfo  Aguas- 
caHentes- Verde  system  is  so  great,  I  have 
arbitrarily  divided  tTie  system  into  two 
parts:  the  Rio  Aguascalientes  basin  in  the 
state  of  Aguascalientes  and  the  remainder 
of  the  Rio  Verde,  primarily  in  Jalisco. 

Collection  data  for  museum  specimens 
indicate  that  Kinosternon  hirtipes  and  K. 
integrum  co-occur  in  the  Rio  Aguas- 
cahentes  between  at  least  about  1900  and 
2000  m  elevation.  Oswaldo  Mooser  (pers. 
comm.)  indicated  that  K.  hirtipes  is  much 
less  common  than  K.  integrum  in 
Aguascalientes,  The  fact  that  21  museum 
specimens  of  K.  integrum  are  available 
from  ten  localities  in  Aguascalientes 
whereas  15  specimens  of  K.  hirtipes  are 
known  from  only  five  localities  (Iverson, 
unpubl.)  support  his  contention. 
Mooser's  field  observations  also  indicate 
that  the  former  occurs  only  in  permanent 
water  situations,  wereas  the  latter  is 
common  in  those  situations  as  well  as 
temporary  aquatic  habitats.  I  have  not 
visited  the  Rfo  Aguascalientes  basin. 

Rio  Verde  drainage,  Jalisco 
(VERD).  -  Draining  most  of  northeastern 
Jalisco,  the  Rio  Verde  empties  near 
Guadalajara  into  the  Rio  Grande  de 
Santiago,  which  flows  through  the  Sierra 
madre  Occidental  to  the  Pacific  Ocean.  K. 
integrum  occurs  throughout  the  entire 
Verde-Santiago  system  (Berry  1978;  Iver- 
son, unpublished),  but  within  the  Rio 
Verde  system  (excluding  Aguascalientes) 
K.  hirtipes  is  known  from  only  three 
localities.  At  least  two  are  permanent 
water  situations,  their  permanence 
enhanced  by  the  construction  of  dams. 

On  16  June  1978,  I  collected  four  K. 
integrum  and  three  K.  hirtipes  in  one  hour 
at  the  most  southerly  known  locality  in 
this  basin  Gust  over  1800  m),  a  tributary 


of  the  Rfo  Verde  north  of  the  city  of  Valle 
de  Guadalupe.  At  that  time  the  stream 
was  reduced  to  a  series  of  isolated,  well- 
vegetated  pools  (one  to  two  m  across  and 
P  1  m  deep)  in  the  channel  below  a  large 
stone  dam.  The  impounded  reservoir 
(Presa  Canada  Grande)  was  unvegetated 
and  reduced  to  a  small  (50  m  X  50  m), 
deep  (2  m  *),  muddy  pond  immediately 
behind  the  dam.  The  heads  of  literally 
hundreds  of  Kinosternon  were  visible  at 
the  pool's  surface  on  that  day.  The  isolat- 
ed pools  in  the  stream  channel,  however, 
contained  few  turtles,  usually  only  one 
per  pool.  Turtles  were  alsoablmdant  here 
on  10  May  1981,  when,  gravid  female  K. 
hirtipes  were  seen  basking. 

The  other  locality  (ca.  2000  n\)  below 
the  Presa  el  Cuarenta  on  the  Rio  del 
Cuarenta  (=  Rfo  de  Lagos  =  Rio  San 
Juan  de  los  Lagos)  near  the  village  of 
Paso  de  Cuarenta,  was  also  visited  on  16 
June  1978.  The  large  cement  dam  im- 
pounds a  huge,  muddy,  unvegetated  res- 
ervoir. Because  of  the  apparent  lack  of 
turtle  habitat  along  the  shoreline,  the 
presa  was  not  trapped.  Below  the  dam, 
however,  were  numerous  rocky-shored  (in 
the  main  channel  and  well  vegetated, 
mud-shored  overflow  and  seepage  pools. 
The  more  vegetated  ponds  were  most  pro- 
ductive, and  eight  trap  settings  produced 
15  K.  hirtipes  and  two  K.  integrum  in  one 
hour. 

K.  hirtipes  is  probably  much  more 
common  in  the  Rio  Verde  than  is  indi- 
cated by  the  paucity  of  locality  records. 
Additional  field  work  should  verify  this 
prediction.  Of  particular  interest  is  the 
downstream  limit  of  A",  hirtipes  in  the  Rfo 
Verde. 

Rio  Grande  de  Santiago  drainage. 
-  This  river  links  the  Lago  de  Chapala 
with  the  Pacific  Ocean  and  passes  through 
the  Sierra  Madre  Occidental.  Tanner  and 
Robison  (1960)  reported  the  collection  of 
three  unidentified  Kinosternon  from  7.5 
mi.  north  of  Magdalena,  Jalisco  (1000  m 
elevation)  in  this  basin.  One  of  the 
included  specimens  (BYU  14630)  is  un- 
questionably   K.     sonoriense    and     the 


14 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


locality  data  for  that  specimen  must  be 
considered  in  error.  The  other  two  speci- 
mens supposedly  collected  at  the  same 
locality  (BYU  14631-32)  are  presently 
unlocatable  (Tanner,  pers.  comm.).  I 
doubt  the  natural  occurrence  of  a  K. 
sonoriense-WkQ  member  of  the  K.  hirtipes 
species  group  in  the  Rio  Grande  de 
Santiago  below  Guadalajara.  Of  those 
major  systems  joining  the  Rio  Santiago 
below  Lake  Chapala,  only  the  Verde 
harbors  K.  hirtipes  (see  above).  K. 
integrum  however,  ranges  throughout  the 
Lerma  and  Santiago  basins  and  all  their 
tributaries,  both  on  the  Plateau  and  off 
(Iverson,  unpublished). 

Conant's  (1978:465;  Map  10)  two  most 
southerly  records  for  K.  hirtipes  in 
Durango  were  erroneously  plotted  in  the 
Rio  Atengo  drainage,  a  tributary  to  the 
Rio  Santiago  (Roger  Conant,  pers. 
comm.);  the  records  belong  in  the  Rio 
Mezquital  basin  near  Ciudad  Durango. 

Rio  Lerma  drainage,  Jalisco,  Guana- 
juato, Michaocan,  and  Mexico.  —  Most 
of  the  southern  portion  of  the  Mexican 
Plateau  is  drained  by  the  Rio  Lerma  and  its 
tributaties.  The  Lerma  originates  in  the 
springs  and  lakes  in  the  southern  end  of 
the  Toluca  Valley  at  over  2400  m  (Gold- 
man  1951:185,   305,   plates  59  and  60; 
Tamayo,  1964:104;  Romero,  1965),  and 
flows    northward    and    then    westward 
across    the    southern    Plateau    to    Lake 
Chapala.  The  basic  physiography  of  the 
river  along  most  of  its  course  is  discussed 
by    Barbour    (1973:541)    and    Tamayo 
(1962).  Like  the  Rio  Grande  de  Santiago, 
the  Rio  Lerma' s  south  bank  tributaries 
are  not  extensive  (the  river  flows  parallel 
to  and  immediately  north  of  the  Sierra 
Volvanica  Transversal),  whereas  several 
of  those  on  the  north  bank  are  very  large 
(notably  the  Rios  Turbio  and  de  la  Laja). 
Because  the  Lerma  basin  is  well  over 
400  airUne  km  long,  I  chose  to  subdivide  it 
for  analysis  of  its  resident  turtle  popu- 
lations. I  have  followed  Barbour  (1973: 
540)  in  his  division  of  the  Rio  Lerma  basin 
into    four    physiographic    regions:    the 
Valley  of  Toluca  (TOL;  above  the  canyon 


below  Temascalcingo  in  the  state  of  Mex- 
ico), the  Maravati'o  basin  (MAR;  from 
near  Temascalcingo,  Mexico  through 
Michoac^n  and  Guanajuato  to  the  rapids 
near  Salvatierra,  Guanajuato),  the  Baji'o 
(BAJ;  from  near  Salvatierra,  Guanajuato 
to  the  region  between  Piedad  and  Yure'- 
cuaro,  Michoac^n),  and  the  Lake 
Chapala  basin  (CHAP;  in  Jalisco  and 
Michoacan.  [The  reader  is  referred  to 
Barbour  (1973)  for  discussion  of  these 
physiographic  provinces.]  I  have  also 
considered  the  turtles  in  the  lower  Rio 
Lerma  tributary,  the  Rio  Duero  (DUER) 
(historically  a  tributary  of  Lake  Chapala; 
see  Tamayo,  1962:404),  as  a  separate  pop- 
ulation for  purposes  of  analysis.  Each  of 
these  subdivisions  is  discussed  separately. 
K.  hirtipes  and/or  K.  integrum  are  the 
only  aquatic  turtles  presently  known  to 
occur  in  these  basins  (But  Pleistocene 
fossils  of  Pseudemys  cf.  scripta  are 
known  from  near  Lake  Chapala;  Tom 
Van  Devender,  pers.  comm.). 

Valley  of  Toluca  basin,  Mexico 
(TOL).  -  No  turtles  are  known  from  the 
Toluca  basin,  but  K.  hirtipes  probably 
occurs  in  the  springs  and  lakes  near  the 
Rio  Lerma  headwaters  in  southeast  Mex- 
ico state. 

Maravatio  basin,  Mexico,  Michoacan 
and  Guanajuato  (MAR).  -  Only  a  single 
broken  K.  hirtipes  shell  (KU  43637)  is 
available  from  this  basin,  and  this  region 
is  thus  unrepresented  in  subsequent  analy- 
sis. No  K.  integrum  are  known  from  the 
Maravatio  (Iverson,  unpublished),  but 
both  species  probably  occur  throughout 
the  basin. 

Bajio  basin,  Guanajuato  and  Micho- 
acan (BAJ).  -  This  basin  includes  the 
drainages  of  the  Lerma  tributaries,  the 
Rio  de  la  Laja  and  Rio  Turbio.  Both  K. 
hirtipes  and  K.  integrum  occur  in  the 
Bajio  up  to  at  least  1900  m  (north  of  San 
Miguel  de  Allende;  Iverson,  unpublished). 
On  12  June  1978  I  sampled  two  marshy 
areas  near  the  Rio  de  la  Laja  between  San 
Miguel  de  Allende  and  Dolores  Hidalgo, 
Guanajuato.  An  hour  at  each  locality  pro- 
duced three  K.  hirtipes  (two  by  hand;  one 


No.  1 


Kinosternon  Biosystematics 


15 


trapped)  and  two  K.  hirtipes  (one 
trapped,  one  seined)  respectively. 

Rio  Duero  drainage,  Michoacdn 
(DUER).  -  K.  hirtipes  is  known  from  only 
one  locality  in  this  drainage.  At  spring- 
fed,  cypress-lined  Lago  Came'cuaro  (1700 
m),  east  of  Zamora  (Symbol  G  in  Fig.  4), 
I  found  K.  hirtipes  very  abundant  on  14 
June  1978;  17  trap  hours  produced  20  K. 
hirtipes  and  one  K.  integrum.  The  latter 
species  is  known  from  several  other  locali- 
ties in  the  Rio  Duero  and  likely  occurs 
throughout  the  basin. 

Villa  Victoria  basin,  Mexico  (state) 
(VILLA).  -  K.  hirtipes  has  been  collected 
at  only  three  localities  within  a  3  km 
radius  in  this  basin  (part  of  the  Rio  Balsas 
basin;  see  later)  at  about  2500  m;  K.  inte- 
grum is  therein  unknown.  K.  hirtipes  is 
apparently  not  common  in  the  basin  as 
evidenced  by  our  collection  of  only  two 
specimens  in  134  trap  hours  at  four  locaH- 
ties  below  the  Presa  Villa  Victoria  on 
21-22  June  1979. 

Lago  de  Chapala  basin,  Jalisco  and 
Michoacdn  (CHAP).  -  Lake  Chapala  (el- 
evation 1525  m)  is  80  km  long,  east  to 
west  and  covers  about  1685  km^  (Debuen 
1945;  Deevey  1957;  see  photographs  in 
Tamayo,  1962).  Average  depth  is  only  8  m 
(Tamayo  1964:105),  and  maximum  depth 
is  probably  9.8  m  (Cole  1963:413).  The 
Rio  Lerma  flows  into  the  extreme  eastern 
end  of  the  lake  and  the  Rk)  Grande  de 
Santiago  exits  the  lake  about  15  km  north 
of  the  Lerma's  mouth.  There  are  no  other 
large  confluent  streams.  All  records  but 
one  (Jiquilpan;  Duellman,  1961)  of  K. 
hirtipes  from  the  Chapala  basin  are  from 
along  its  shores.  K.  integrum  has  also 
been  commonly  collected  along  the  lake 
shores  (Berry  1978;  Iverson,  unpublished). 
Trapping  along  the  south  shore  near 
Tuxcueca  on  15  June  1978  produced  no 
turtles  whatsoever,  although  J.F.  Berry 
(pers.  comm.)  obtained  a  series  oi K.  inte- 
grum at  the  same  locality  in  June  1975. 

The  once  extensive  marshes  at  the  east- 
ern end  of  Lake  Chapala  probably  sup- 
ported dense  populations  of  Kinosternon 
turtles,  but  drainage  operations  have  un- 


fortunately nearly  eliminated  this  habitat 
(Goldman,  1951:173-174). 

Laguna  de  Zapotldn  interior  drainage, 
Jalisco  (ZAPO).  -  The  Zapotlan  basin  lies 
north  of  Ciudad  Guzman,  Jalisco  in  the 
Sierra  Volcanica  Transversal.  Only  K. 
hirtipes  is  known  from  the  basin  and  all 
specimens  apparently  originated  at  the 
southern  end  of  the  lake  (ca.  1 500  m)  near 
Ciudad  Guzman.  Gadow's  (1908)  record 
of  Cinosternum  integrum  from  this  basin 
must  therefore  be  based  on  K.  hirtipes. 

San  Juanico  Valley  interior  drainage, 
Michoacdn  (SNJ).  -  The  Valley  of  San 
Juanico  (north  of  Cotija,  Michoacan)  was 
until  recently  an  isolated,  interiorly 
drained  basin,  formed  prehistorically  by 
the  damming  of  a  northward-flowing  trib- 
utary of  the  Chapala  basin  by  a  lava  flow 
(Alvarez  1963,  1972;  Barbour  1973).  The 
construction  of  the  Presa  San  Juanico 
across  the  valley's  southern  end  has  en- 
larged Lake  San  Juanico,  and  directed  its 
effluents  southward  to  the  Balsas  system 
(Alvarez  1972:158;  Barbour  1973;  pers. 
observ.). 

Turtles  of  the  Kinosternon  hirtipes 
group  are  the  only  turtles  known  from  the 
valley  behind  and  above  the  Presa  San 
Juanico  (ca.  1500  m;  Symbol  H  in  Fig.  4). 
Field  work  by  Clyde  Barbour  (pers, 
comm.)  and  my  own  field  crew  (75  trap 
hours,  14-15  June  1978;  180  trap  hours, 
6-7  May  1981)  in  i\iQ  presa  (Figure  6)  have 
produced  only  7  specimens,  three  Hving 
(one  seined  by  Barbour;  two  trapped  by 
me)  and  four  articulated  shells  (by  my 
crew).  Ichthyological  field  work  in  the 
presa  on  three  dates  in  1962  and  1963  by 
Alvarez  (1963)  apparently  produced  no 
turtles. 

The  diversion  of  effluents  from  the  San 
Juanico  Valley  to  the  Balsas  appears  to  be 
permitting  K.  integrum  (known  through- 
out the  Balsas  system;  Berry,  1978;  Iver- 
son, unpublished)  to  expand  its  range 
toward  the  presa.  Although  no  K.  inte- 
grum are  known  from  above  the  presa, 
and  although  we  obtained  no  turtles  3  km 
below  the  dam  in  1978  in  one  of  the  two 
effluent   irrigation    ditches    (during   one 


16 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


hour  of  seining  and  12.5  trap  hours),  a 
single  K.  integrum  observed  sunning 
along  the  other  ditch  (ca.  100  m  below  the 
dam)  was  seined.  An  additional  juvenile 
K.  integrum  (TUL  19504)  is  also  known 
from  one  of  the  effluent  ditches  where  it 
crosses  the  Cotija-Tocumbo  road  (dis- 
tance below  dam  uncertain).  K.  integrum 
likely  will  soon  invade  Lake  San  Juanico. 

In  June  1978,  the  lake  itself  was  very 
low;  much  of  the  muddy  bottom  was  ex- 
posed due  to  evaporation  and  diversion 
for  agriculture  (Fig.  6).  A  small  water 
hyacinth  population  represented  the  only 
obvious  vegetation  in  the  muddy  lake.  In 
1981  the  lake  was  even  lower  and  new 
ditches  were  draining  the  lake  even  more. 
No  turtles  were  collected  and  their  exist- 
ence seems  tenuous. 

Lago  de  Cuitzeo  interior  basin,  Micho- 
acan  CUIT).  -  Lago  de  Cuitzeo  (just  over 
1800  m)  is  the  largest  interiorly  drained 
natural  lake  in  Mexico.  It  is  fed  primarily 
by  the  Rio  Grande  de  Morelia,  which 
heads  in  the  mountains  east  of  P^tzcuaro 
and  flows  southeastward  to  its  confluence 
with  the  southeastern  shore  of  the  lake 
(Camacho,  1925).  The  lake  is  very  shal- 
low, and  has  been  known  to  be  nearly  dry 
(Debuen,  1943).  Aquatic  vegetation  is 
accordingly  uncommon.  When  visited  on 
12  June  1977,  the  lake  level  was  very  high 
and  therefore  not  trapped  for  turtles. 
Based  solely  on  museum  specimens,  K. 
hirtipes  (one  specimen)  is  much  less  com- 
mon than  K.  integrum  (35  specimens,  7 
localities;  Iverson,  unpublished).  The 
single  available  specimen  is  a  poorly  pre- 
served male. 

Lago  de  Pdtzcuaro,  interior  basin, 
Michoacan  (PATZ).  -  Lago  de  Patzcuaro 
(2,035  m;  Symbol  I  in  Fig.  4;  see  phot- 
graphs  in  Tamayo,  1962:493  and  Solor- 
zano  Preciado  1961:55)  has  been  well- 
studied  limnologically  (summary  in  Cole, 
1963),  but  its  turtles  have  been  only  in- 
frequently mentioned  (Duellman,  1961; 
Altini,  1942)  or  completely  ignored 
(Martin  del  Campo,  1940).  It  has  a  sur- 
face area  of  only  about  1 1 1  km^  and  has  a 
maximum  depth  of  15  m  (DeBuen,  1944). 


Figure  6.  Presa  San  Juanico,  Michoacdn,  looking 
northwestward  from  dam,  on  15  June  1978. 
Reservoir  was  much  reduced  due  to  irrigation  de- 
mands and  the  dry  season.  Kinosternon  hirtipes 
magdalense  was  collected  along  the  dredged  canal 
in  foreground. 

Emergent  vegetation  (primarily  Scirpus)  is 
common  along  the  shoreline,  especially 
the  southern  margin  (Goldman,  1951:195, 
plate  58;  Barbour,  1973:543),  where  the 
mats  often  extend  out  20  m  or  more  from 
the  shoreline  (pers.  observ.). 

On  12-13  June  1978,  the  lake  was  quite 
clear;  however,  we  saw  no  turtles  during 
the  day  or  night  in  shallow  water  (<  1  m) 
in  narrow  strips  of  shoreline  on  the  south- 
eastern shore  where  emergent  vegetation 
has  been  removed  for  docks  (Fig.  7). 
Kinosternon  hirtipes  (the  only  turtle 
known  from  the  lake)  were,  nevertheless, 
for  sale  the  next  day  in  the  Patzcuaro 
(city)  market,  an  apparently  frequent  oc- 
currence. 

The  smaller  (8  km^)  higher  (2120  m), 
younger,  and  deeper  (maximum  depth  45 
m)  Lago  de  Zirahuen  (DeBuen,  1943, 
1944)  immediately  to  the  southwest,  is 
believed  to  have  been  historically  drained 
by  a  tributary  of  the  Rio  Lerma  flowing 
through  the  Lake  Patzcuaro  and  Lake 
Cuitzeo  basins  (DeBuen,  1943).  No  turtles 
are  known  from  Lake  Zirahuen. 

Rio  Balsas  drainages,  Michoacan  and 
Puebla  (BALS).  -  Turtles  of  the  hirtipes 
group  have  been  recorded  from  only  three 
localities  in  the  Rio  Balsas  basin,  the  river 
sytem  which  drains  most  of  southern 
Mexico    south   of  the   Sierra   Volcanica 


No.  1 


Kinosternon  Biosystematics 


17 


Transversal.  The  Villa  Victoria  (VILLA) 
localities  have  already  been  discussed. 
Duellman  (1961)  recorded  K.  hirtipes 
from  the  Balsas  on  the  basis  of  UIMNH 
24707  from  8  km  W  Ciudad  Hidalgo, 
Michoacan  (ca.  2200  m;  Rio  Tuxpan 
Valley;  Symbol  D  in  Fig.  4).  I  have  exam- 
ined the  specimen  and  believe  it  to  be  a 
female  K.  integrum.  However,  another 
specimen  from  the  same  locality  (AMNH 
62257)  is  unquestionably  a  female  K. 
hirtipes.  In  addition,  I  collected  a  single 
male  K.  hirtipes  in  128  trap  hours  in  a 
stream  of  approximately  the  same  locality 
on  4-5  May  1981.  Kinosternon  integrum 
occurs  throughout  the  Tuxpan  (and 
Balsas)  system  (Berry,  1978;  Iverson,  un- 
published); but  K.  hirtipes  is  now  definite- 
ly known  in  the  Tuxpan  only  near  Ciudad 
Hidalgo. 

The  only  other  supposed  Balsas  speci- 
men of  K.  hirtipes  is  an  adult  male  (UU 
12096)  from  a  tributary  of  the  Rfo 
Atoyac,  4.5  km  S  Molcaxac  (just  below 
2000  m;  Symbol  E  in  Fig.  4).  The  identifi- 
cation is  correct,  but  I  question  the  valid- 
ity of  the  data  for  three  reasons.  First, 
because  of  the  numerous  highway  access- 
es to  the  RTo  Atoyac  drainage  and  the  fact 
that  I  know  of  at  least  88  specimens  of  K. 
integrum  (73  of  which  I  have  seen)  from 
13  localities  in  the  Atoyac-Balsas  system- 
in  the  state  of  Puebla,  additional  speci- 


Figure  7.  Southeastern  shore  (foreground)  of  Lago 
de  Pitzcuaro,  Michoacan,  13  June  1978. 


mens  o[  K.  hirtipes  would  likely  have  been 
collected  if  the  species  did  occur  in  that 
system.  Second,  nine  trap  hours  at  the 
Molcaxac  locality  on  3  May  1981  pro- 
duced 23  K.  integrum  and  no  K.  hirtipes. 
Third,  based  on  field  numbers  and  collec- 
tion dates,  Clyde  Barbour  collected  K. 
hirtipes  along  the  Rfo  Lerma  in  Jalisco 
(UU  12120)  on  7  May  1969  and  K.  inte- 
grum in  the  Rio  Turbio  in  Guanajuato 
(UU  12083-84)  on  8  May  and  in  the  Rfo 
Villeto  in  San  Luis  Potosf  (UU  12085)  on 
12  May]  immediately  before  he  collected 
near  Molcaxac  (18  May  1969).  Of  the 
eleven  turtles  recorded  as  collected  near 
Molcaxac,  ten  are  definitely  K.  integrum. 
I  submit  that  through  a  mixup,  the  Mol- 
caxac locality  datum  was  mistakenly 
applied  to  the  eleventh  specimen,  and  that 
the  specimen  possibly  originated  some- 
where in  the  Lerma  basin  (Rfos  Lerma  or 
Turbio?)  where  Barbour  also  collected. 

Valley  of  Mexico  interior  basin 
(VALLE).  -  The  physiography  and  histor- 
ical geology  of  the  Valley  of  Mexico  in 
which  Mexico  City  lies  has  been  well- 
studied  (Bryan  1946,  1948;  De  Terra  et 
al.,  1949;  Arellano,  1953;  Sokoloff  and 
Lorenzo,  1953;  Zeevaert,  1953;  Foreman, 
1955;  Hibbard,  1955;  Maldonado-Koer- 
dell,  1955;  Sears  and  Clisby,  1955; 
Mooseretal.,  1956;  Mooser,  1957,  1963; 
Deevey,  1957;  Lorenzo,  1958;  Bernal, 
1959;  Bribiesca  Castrejon,  1960;  White, 
1962;  Golomb,  1965;  Bradbury,  1971; 
and  an  excellent  summary  in  Barbour 
1973:537).  The  entire  basin  is  about  24  km 
wide  and  113  km  long  (Foreman,  1955) 
and  covers  about  8000  km-  (Maldonado- 
Koerdell,  1955:15).  At  the  time  of  the 
Spanish  conquest  (ca.  1520),  the  Valley  of 
Mexico  was  one  of  the  largest  interior 
drainage  basins  in  the  Transverse  Vol- 
canic Arc,  supporting  five  large  spring- 
fed  lakes  (Tamayo,  1964;  De  Terra  et  al., 
1949;  among  others).  So  extensive  were 
the  lakes  at  that  time,  that  the  early  city  of 
Mexico  had  been  built  on  an  island  and 
the  Spaniards  were  forced  to  build  ships 
in  order  to  besiege  the  city  (Huntington 
1914:96).     Tremendous    fluctuations    in 


18 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


water  level  in  the  Valley  prompted  drain- 
age operations  in  the  late  16th  century, 
and  by  1608  some  of  the  Valley's  water 
was  diverted  northward  to  the  Rfo  Tula 
(Atlantic  drainage;  Tamayo,  1964; 
Barbour,  1973:540).  This  artificial  drain- 
age system  was  finally  completed  in  1900 
(Huntington,  1914:97;  Bribiesca  Castrejon, 
1960)  and  they  only  sizeable  lakes  in  the 
Valley  today  are  Zumpango  (2243  m 
elevation)  and  Texcoco  (2236  m)  (Bar- 
bour, 1973).  Only  about  lOiVo  of  the 
Valley  floor  is  covered  with  water 
(Foreman,  1955).  For  discussions  of  the 
changing  conditions  of  the  lakes  since 
about  1500  AD,  see  Bribiesca  Castrejon 
(1960).  Aquatic  habitats  in  the  Valley 
were  discussed  and  photographed  by  Gadow 
(1908:6)  and  Goldman  (1951: 138-39,  plate 

56).  ,     , 

Apparently  the  first  record  of  a  turtle 

from  the  Valley  of  Mexico  is  Wagler's 
(1830)  description  of  Cinosternon  hirtipes 
(see  later  justification).  Numerous  liter- 
ature records  (See  synonymies)  and  muse- 
um specimens  confirm  the  presence  of  K. 
hirtipes  in  the  Valley  of  Mexico.  Kino- 
sternon  integrum  has  possibly  been  col- 
lected in  the  Valley  only  three  times. 
FMNH  116521  bears  only  the  data  "Dis- 
trito  Federal".  Data  associated  with  SM 
9722-23  indicate  they  were  purchased  in 
the  Xochimilco  market  on  11  June  1962. 
Fourteen  turtles  purchased  for  me  by 
Gustavo  Casas-Andreu  in  the  Xochimilco 
market  in  August  1977  are  all  K.  inte- 
grum. The  merchants  told  him  the  turtles 
were  from  the  "Valle  de  Mexico". 
Because  there  is  no  verifed  record  of  the 
occurrence  of  K.  integrum  in  the  Valley 
before  1962,  and  because  all  three  subse- 
quent records  are  apparently  from  mark- 
ets, I  strongly  doubt  the  natural  occur- 
rence of  K.  integrum  in  the  Valley  of 
Mexico.  K.  integrum  is,  however,  very 
abundant  southeast  of  the  Valley  in 
Puebla  and  Oaxaca  (Berry,  1978;  personal 
observation;  see  discusssion  under  Rfo 
Balsas),  and  may  have  been  imported  to 
the  Valley  for  sale  in  the  markets.  Support 


for  such  an  hypothesis  comes  from 
Berry's  (1978:83,  Fig.  17)  discriminant 
analysis  of  data  from  the  turtles  sent  to 
me  by  Casas  Andreu  (UF  41651-64), 
which  clearly  showed  their  affinities  to  be 
with  turtles  from  the  Upper  Rfo  Balsas 
(Rio  Mexcala)  and  the  Upper  Rfo  Papalo- 
apan  (Rfo  Santa  Domingo  basin,  Puebla 
and  Oaxaca).  Whatever  their  true  origin, 
K.  integrum  will  likely  soon  establish 
itself  in  the  Valley.  A  study  of  the  inter- 
action of  that  species  with  the  native  K. 
hirtipes  would  be  significant. 

Mittermeier  (1971:16)  found  Pseud- 
emys  scripta  elegans,  obviously  intro- 
duced, in  the  markets  of  Mexico  City, 
where  he  was  told  that  the  species  had 
been  introduced  into  ponds  near  Mexico 
City.  No  other  turtle  is  known  from  the 
Valley  of  Mexico. 

Unrepresented  or  unsampled  basins. 
-  Several  other  isolated  and/or  interior 
drainage  basins  within  or  adjacent  to  the 
range  of  A',  hirtipes  should  be  investigated 
for  that  species.  These  include  the  Laguna 
de  Babicora  (2100  m),  northwest  of 
Gomez  Farias,  Chihuahua;  the  Laguna  de 
Los  Mexicanos  (2100  m),  south  of 
Cuauhtemoc,  Chihuahua;  the  Laguna  de 
Zapacu  and  Presa  de  Copandaro,  near 
Zacapu  in  northern  Michoacan;  the  crater 
lakes  of  the  Llanos  of  Pueblo  (see  Alva- 
rez, 1949);  the  Rio  Mezquitic  (=  Rfo 
Balanos),  tributary  to  the  Rio  Grande  de 
Santiago  and  accessible  near  Valparaiso, 
Zacatecas;  the  Laguna  de  Sayula  (about 
1300  m),  north  of  Sayula,  Jalisco;  and  the 
Laguna  de  San  Marcos  (ca.  1300  m),  near 
Zacoalco,  Jalisco.  K.  integrum  occurs  in 
the  latter  three  basins  (Iverson,  un- 
published), where  it  is  the  only  turtle 
species  known. 

CHARACTERS 

Nineteen  shell  measurements  were 
made  with  dial  calipers  on  museum  speci- 
mens of  the  hirtipes  species  group  from 
drainage  basins   discussed   above.   Only 


No.  I 


Kinoslernon  Biosyslematics 


19 


data  from  specimens  over  80  mm  cara- 
pace length  (except  three  females)  with  the 
full  complement  of  measurements  were 
used  in  the  morphometric  analyses;  vari- 
ous ratios  of  characters  were  also  em- 
ployed to  minimize  ontogenetic  variation 
(see  STATISTICAL  TECHNIQUES). 
Character  means  and  ranges  by  popula- 
tion (Appendices  1  and  2)  and  taxon  (Ap- 
pendices 3  and  4)  for  each  sex  are  avail- 
able from  NAPS'. 

Mensural  characters  recorded  and  their 
abbreviations  follow  [Methods  of 
measurement  were  given  by  Iverson 
(1977a);  midline  plastral  scute  measure- 
ments were  always  made  on  the  animal's 
right  side.]:  carapace  length  (CL),  cara- 
pace width  (CW),  maximum  plastral 
length  (PL),  plastral  widths  measured  at 
the  lateral  edges  of  the  seams  between  the 
humeral,  pectoral,  abdominal,  femoral, 
and  anal  laminae  (WA,  WB,  WC,  and 
WD  respectively),  bridge  length  (BL), 
gular  length  (GL),  gular  width  (GW), 
interhumeral  seam  length  (IH),  inter- 
pectoral  seam  (IP),  interabdominal 
(lAB),  interfemoral  (IF),  interanal  (IAN), 
first  vertebral  width  (VW)  and  length 
(VL),  maximum  length  of  plastral  fore- 
lobe  (FL),  and  maximum  length  of  plas- 
tral hindlobe  (HL).  The  ratios  of  each 
character  to  CL  as  well  as  the  ratios  of 
IH,  IP,  lAB,  IF,  and  IAN  to  PL  (total,  23 
ratios)  were  employed  in  the  analysis.  In 
some  analyses,  the  number  of  variables 
was  reduced  to  those  thirteen  (excluding 
the  ratios  CW,  HL,  PWD,  IH,  IP,  lAB, 
IF,  IAN,  VW  and  VL  to  CL)  with  the 
greatest  variation  in  the  species  group. 


,See  NAPS  document  03915 for  20  pages  of  supple- 
mentary material.  Order  form  NAPS,  c/o  Micro- 
fiche Publications,  P.O.  Box  3513,  Grand  Central 
Station,  New  York,  NY  10017,  USA.  Remit  in 
advance  for  each  NAPS  accession  number.  Insti- 
tutions and  organizations  may  use  purchase  orders 
when  ordering;  however,  there  is  a  billing  charge  of 
$5.00  for  this  service.  Mack  checks  payable  to 
Microfiche  Publications.  Photocopies  are  $5.00. 
Microfiche  are  $3.00  each.  Outside  the  United  States 
and  Canada,  postage  is  $3.00  for  a  photocopy  and 
$1.00  for  a  fiche. 


Those  analyses  are  noted  in  the  text.  Sexes 
(males  have  long  tails  and  scale  patches  on 
the  hind  legs)  were  always  analyzed  sep- 
arately. 

Relative  shell  height  has  been  used  to 
distinguish  Kinosternon  hirtipes  from  K. 
sonoriense  (e.g.,  Ernst  and  Barbour, 
1972;  Wermuth  and  Mertens,  1961);  how- 
ever, the  character  is  difficult  to  measure 
consistently  and  preliminary  analysis 
revealed  it  would  not  reliably  separate  the 
two  taxa.  It  has  therefore  not  been  used  in 
this  analysis. 

Qualitative  characters  also  recorded  in- 
cluded relative  head  size,  plastral  color, 
and  shell  carination  as  well  as  the  follow- 
ing. 

Nasal  scale.  -  As  described  by  Conant 
and  Berry  (1978:3),  adult  kinosternids 
have  a  patch  of  cornified  epithelium 
which  extends  from  the  dorsal  margin  of 
the  rostrum  for  a  variable  distance  poster- 
iorly on  the  dorsum  of  the  head.  A 
drawing  of  the  shape  and  extent  of  the 
nasal  scale  on  each  individual  turtle  was 
made. 

Chin  barbels.  -  The  number,  relative 
sizes,  and  locations  of  chin  and  neck 
barbels  were  recorded. 

Head  pattern.  -  Although  often  quite 
variable  and  difficult  to  describe,  an  at- 
tempt was  made  to  qualify  head  patterns. 
The  procedure  involved  photographing 
the  heads  of  as  many  specimens  as  possi- 
ble (over  500  total  head  photographs 
available)  for  later  simultaneous  exam- 
ination and  comparison. 

STATISTICAL  TECHNIQUES 

Character  ratios  were  employed  in  the 
statistical  analyses  despite  recent  criticism 
of  their  use  by  Atchley  et  al.  (1975,  1976). 
This  decision  is  based  on  arguments  in 
favor  of  their  use  by  Corruccini  (1977), 
Nussbaum  (1976),  Dodson  (1978),  Heyer 
(1978),  and  Iverson  (1979),  as  well  as  the 
articulate  demonstration  by  Berry  (1978) 
that,  for  at  least  one  other  Kinosternon 
species  group,  the  use  of  ratios  as  input 
variables  in   both   multiple  discriminant 


20 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


analysis  and  distance  (D-)  analysis  pro- 
duced results  almost  identical  to  those 
obtained  by  using  residual  values  from  re- 
gression analysis  as  input  variables  (the 
standardization  technique  recommended 
by  Atchley  et  al.,  1976).  My  own  unpub- 
lished data  on  other  kinosternid  species 
also  support  the  effectiveness  and  reliabil- 
ity of  multivariate  analyses  using  char- 
acter ratios  for  at  least  this  family  of 
turtles.  Furthermore,  I  attach  no  statist- 
ical significance  to  multivariate  output 
generated  from  ratios.  The  output  is  only 
used  as  a  tool  to  pinpoint  distinctive 
samples,  and  to  suggest  the  characters 
most  important  to  those  distinctions. 

Simple  statistics  were  performed  using 
the  Statistical  Analysis  System  (Service, 
1972).  Standard  deviations  accompany 
mean  character  ratios  only  as  a  relative 
measure  of  dispersion;  no  statistical  sig- 
nificance is  implied.  Step-wise  discimi- 
nant  analyses  (see  Gould  and  Johnston, 
1972,  and  Sneath  and  Sokal,  1973,  for 
review  of  the  procedure)  utilized  the  Bio- 
medical Programs  BMD07M  (W.J. 
Dixon,  1973),  and  BMDP7M  (W.J. 
Dixon,  1977).  Cluster  analyses  employed 
the  NT-SYS  (Rohlf  and  Kispaugh,  1972) 
and  BMDP2M  programs.  Specific  appli- 
cations of  these  analyses  are  outlined  un- 
der MORPHOMETRIC  ANALYSIS. 

Because  of  the  pronounced  sexual  di- 
morphism in  members  of  the  K.  hirtipes 
species  group,  sexes  are  considered  sep- 
arately in  all  cases.  Turtles  from  basin 
samples  represented  by  only  one  or  two 
individuals  of  either  sex  were  included  in 
the  analysis  as  unknowns,  and  assigned  to 
the  most  phenetically  similar  sample  by 
discriminant  analysis. 

BIOSYSTEMATIC  TACT 

Like  many  evolutionary  biologists  be- 
fore me,  the  problem  of  interpreting  the 
genetic  (and  taxonomic)  relationships  of 
closely-related,  allopatric  vertebrate  pop- 
ulations is  a  perplexing  one  (see  Inger, 
1961;  Amadon,  1966,  1968;  Mayr, 
1970:210-211;  Amadon  and  Short,  1976). 
The  propensity  (perhaps   restriction)  of 


members  of  the  K.  hirtipes  group  for 
permanent  water  habitats,  coupled  with 
the  geographical  isolation  of  inhabited 
river  basins  due  to  historical  geology  and 
desertification  have  produced  at  least 
thirty  allopatric  populations  of  members 
of  this  group.  Many  of  these  populations 
differ  notably  from  geographically  adjac- 
ent populations,  but  are  quite  similar  to 
other  populations  far  removed  (see 
RESULTS).  Interpreting  such  complex 
variational  patterns  is  difficult. 

In  this  paper  I  have  taken  a  conserva- 
tive approach  to  the  species-subspecies 
dilemma.  Within  a  species  morphologi- 
cally distinct,  isolated  (i.e.  allopatric)  pop- 
ulations are  afforded  only  subspecific 
status  even  though  additional  data 
(especially  breeding  information)  may 
show  that  some  are  full  biological  species. 
The  genetic  relationships  of  the  popula- 
tions so  named  are  unfortunately  clouded 
by  this  taxonomic  designation  [Amadon 
and  Short,  1976,  define  "megasubspecies" 
and  "allospecies"  in  an  attempt  to 
counteract  this  confusion];  however, 
complementary  studies  of  protein  varia- 
tion now  in  progress  should  perhaps 
further  clarify  the  specific-subspecific 
(i.e.  genetic)  relationships  of  these  turtle 
populations. 

LITERATURE 

Because  far  more  than  half  the  litera- 
ture records  of  kinosternid  turtles  in  Mex- 
ico are  in  error  I  had  to  assume  that  every 
literature  record  was  incorrect  until  per- 
sonally verified  by  examination  of  the 
respective  specimens  or  by  analysis  of  dis- 
tributional information  (for  example,  in 
cases  where  only  one  kinosternid  occurs  in 
a  particular  basin).  I  have  therefore 
attempted  to  substantiate  every  literature 
record  for  any  kinosternid  from  through- 
out the  range  of  the  members  of  the  K. 
hirtipes  group  (or  stated  as  being  from 
that  range),  and  any  member  of  the  Kino- 
sternon  hirtipes  group  (i.e.  K.  sonoriense 
or  K.  hirtipes).  Complete  chronological 
synonymies  were  then  compiled  for  K. 
sonoriense  and  K.  hirtipes,  and  each  ref- 


No.  1 


Kinosternon  Biosystematics 


21 


erence  was  annotated  to  indicate  the  reas- 
ons for  its  inclusion.  Copies  of  this 
annotated  synonymy  have  been  deposited 
in  the  Florida  State  Museum  Herpetology 
Library  (University  of  Florida),  and  are 
available  from  the  author  as  well.  Most  of 
this  information  appears  in  this  paper  in 
the  SYSTEMATICS  Section,  with  some 
elaboration  in  the  next  section. 

RESULTS  AND  DISCUSSION 

Literature  Corrections 

Because  of  the  past  difficulty  in  the 
discrimination  of  the  K.  scorpioides 
group  (including  K.  integrum  and  K. 
alamosae)  from  the  K.  hirtipes  group 
(including  K.  hirtipes  and  K.  sonoriense), 
the  literature  on  Mexican  Kinosternon  has 
accumulated  so  many  errors  of  identifica- 
tion that  it  is  almost  unusable.  The  fol- 
lowing literature  corrections  (ordered  by 
the  valid  taxon  with  which  the  K.  hirtipes 
group  member  was  confused)  are  an 
attempt  to  bring  some  order  and  accuracy 
to  the  error-plagued  literature. 
Sternotherus  odoratus. 

Apart  from  Meek's  El  Sauz,  Chihua- 
hua specimen  of  Sternotherus  odoratus 
(see  discussions  in  Moll  and  Williams, 
1963  and  Conant  and  Berry,  1978:15), 
that  species  (or  its  nomenclatural  equiva- 
lent) has  been  frequently,  though  errone- 
ously, recorded  from  Mexico.  Duges 
(1869:143)  was  apparently  the  first  to 
record  "Ozotheca  (odorata?)"  from 
"Guanajuato  y  Mexico",  but  in  a  subse- 
quent list  (1888:106)  he  apparently 
changed  his  identification  to  Cinosternon 
pensylvanicum.  Because  K.  hirtipes 
occurs  in  both  Guanajuato  and  Mexico 
states,  because  it  is  more  similar  to  S. 
odoratus  than  is  K.  integrum  (the  only 
other  turtle  recorded  from  those  two 
states),  and  because  Duges  was  apparently 
not  aware  of  Wagler's  (1830)  description 
of  A",  hirtipes  from  "Mexico"  (the  species 
is  not  on  his  1869  list),  Duges'  Ozotheca 
record  was  almost  certainly  based  on 
Kinosternon  hirtipes.  The  following 
orthographic  variations  of  Duges'  record 


were  apparently  based  on  his  1869  list  and 
are  thus  considered  synonymous  (in  part) 
with  K.  hirtipes:  Ozotheca  odorata, 
Velasco  (1890b:54,  1891:52,  1892b:40, 
1893b:81,  1894:40,  1896a:30,  1898:62); 
Ozothea  odorata,  Velasco  (1892a:76, 
1892c:79,  1895:38,  1896b:37);  and  Ozho- 
teca  odorata,  Garcias-Cubas  (1884:179) 
and  Velasco  (1890a:35,  1893a:64, 
1897:41).  In  addition,  Conant  and  Berry 
(1978:15)  have  clarified  Brown's 
(1950:230)  record  of  Sternotherus 
odoratus  from  Presidio  Co.,  Texas;  the 
specimen  on  which  the  record  was  based  is 
TCWC  650,  the  holotype  of  Kinosternon 
murrayi  Glass  and  Hartweg  195 1(=  K. 
hirtipes  murrayi).  The  last  erroneous 
record  is  that  of  Altini  (1942:159)  for 
Kinosternon  odoratum  in  Veracruz;  based 
on  his  specimen  description,  it  is  appar- 
ently referable  to  K.  leucostomum. 
Kinosternon  subrubrum. 

Testudo  pensilvanica  Gmelin  (1788: 
1042)  has  been  recognized  as  a  synonym 
of  Kinosternon  subrubrum  (Lacepede, 
1788:132)  at  least  since  1917  (see  Iverson, 
1977b).  Prior  to  that  time,  however,  the 
former  specific  name  had  an  active  his- 
tory in  the  Mexican  herpetological  litera- 
ture, despite  the  fact  that  the  species  does 
not  range  in  Mexico.  Lichtenstein 
(1856:2)  was  the  first  to  apply  the  name 
(as  Cinosternon  pensylvanicum)  to  speci- 
mens from  Mexico  in  the  Berlin  Museum. 
Over  the  next  50  years,  no  fewer  than  22 
papers  recorded  that  species  name  (or 
orthographic  variations  thereoO  for  speci- 
mens from  Mexico.  Based  on  the  greater 
similarity  of  T.  pensilvanica  (i.e.  K. 
subrubrum)  to  K.  hirtipes  than  to  K. 
integrum,  and  the  fact  that  most  of  these 
references  are  based  on  specimens  from 
Guanajuato  (where  integrum  and  hirtipes 
co-occur)  and/or  the  Valley  of  Mexico 
(where  only  hirtipes  naturally  occurs;  see 
MATERIALS  AND  METHODS),  the 
following  binomials  and  references  most 
probably  refer  to  K.  hirtipes:  Cinosternon 
pennsylvanicum,  Duges  (1888:106;  1890, 
in  Velasco  1890b:291;  1895:5;  1896a:lv; 
sylvanicum,    Bocourt    (1876:5),    Herrera 


22 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


(1890:330;  1891:46;  1893:339;  1904:5), 
(1890:330;  1891:46;  1893:339;  1904:5), 
Herrera  and  Lope  (1899:281),  Westphal- 
Castelnau  (1872:278),  and  Strauch 
(1890:88);  Cinosternonus  pensylvanicum, 
Herrera  (1899:28;  for  discussion  see  H.M. 
Smith  and  R.B.  Smith,  1975:86);  Cino- 
sternum  pennsilvanicum,  Cope 
(1900:1229);  Cinosternum  pennsylvan- 
icum,  Gadow  (1905:209);  Cynosternon 
pensylvanicum,  Herrera  and  Lope 
(1899:131);  Cynosternon  pennsylvani- 
cum,  Herrera  (1893:342);  and  Kinostern- 
um  pennsilvanicum,  Cope  (1896:1021). 

The  failure  of  these  person  to  recognize 
their  specimens  as  K.  hirtipes  Wagler  is 
probably  a  consequence  of  the  lack  of  a 
nuchal  scute  by  Wagler's  only  type  spec- 
imen (see  later).  Unaware  that  a  missing 
nuchal  scute  (actually  worn  away)  is  an 
uncommon,  though  natural  anomaly, 
A.M.C.  Dumeril  and  G.  Bibron 
(1834:370),  A.H.A.  Dumeril  (1870:25), 
Bocourt  (1876:50)  and  Duges  (1888:106) 
used  the  absence  of  that  scute  as  the  key 
character  in  identifying  hirtipes. 

Several  additional  orthographic  vari- 
ations were  not,  however,  based  on  K. 
hirtipes.  Gadow's  (1905:194)  record  of 
Cinosternum  pennsylvanicum  from  Guer- 
rero must  be  based  on  K.  integrum  if  the 
datum  is  correct,  because  it  is  the  only 
Kinosternon  found  there. 

Lampe's  (1901:185)  description  of 
Cinosternum  pensylvanicum  from  north 
Mexico  makes  it  clear  he  is  referring  to 
Kinosternon  subrubrum  hippocrepis 
(probably  from  Texas). 

Siebenrock's  (1905:465)  erroneous 
record  of  Testudo  pensylvanica  from 
Veracruz  is  possibly  based  on  a  specimen 
of  A',  herrera i. 

Cinosternon  hippocrepis  (another 
synonym  of  K.  subrubrum;  see  Iverson, 
1977b)  was  erroneously  recorded  from 
Sonora  by  Strauch  (1865:100,  184)  pre- 
sumably based  on  a  specimen  of  A^.  sonor- 
iense. 
Kinosternon  flavescens. 

Several  K.  flavescens  records  are  in  part 
based   on   members   of  the  K.    hirtipes 


species  group.  Most  of  these  have  been 
previously  discussed  (Iverson,  1978).  In 
addition.  Cooper  (1870:66)  recorded 
Platythyra  flavescens  from  the  Colorado 
River  Valley  along  the  California  border 
(precise  locality  unknown).  1  have  else- 
where (Iverson,  1978:477)  questioned  the 
existence  of  A',  flavescens  in  the  Colorado 
River  basin  and  here  suggest  that 
Cooper's  record  was  almost  certainly 
based  on  K.  sonoriense. 
Kinosternon  scorpioides  group. 

The  true  identity  of  the  species  of  Kino- 
sternon occuring  on  Marfa  Madre  Island 
in  the  Tres  Marfas  Islands  off  the  coast  of 
Nayarit  has  plagued  herpetologists. 
Gunther  (1885:15)  first  recorded  and  fig- 
ured K.  hirtipes  from  the  island,  but  the 
same  specimens  were  called  K.  integrum 
by  Boulenger  (1889:42).  Both  Strauch 
(1890:91)  and  Stejneger  (1899:64)  sup- 
ported Boulenger's  view,  yet  Gadow 
(1905:209)  advocated  Gunther's  original 
designation.  Siebenrock  (1906:96)  was  the 
next  to  support  Boulenger's  position. 
H.M.  Smith  and  Taylor  (1950a:25) 
avoided  the  problem  by  recording  both 
species  from  the  islands.  Zweifel 
(1960:94)  next  addressed  the  problem  in 
his  study  of  the  herpetofauna  of  the 
islands.  In  collaboration  with  Norman 
Hartweg,  he  finally  corrected  the  record; 
K.  integrum  is  the  only  species  of  the 
genus  occurring  in  the  Tres  Marfas. 
Wermuth  and  Mertens  (1961  :Fig.  13,  p. 
20)  reproduced  Gunther's  (1885)  figures 
and  recommitted  the  latter's  error.  Casas 
Andreu  (1967:44)  likewise  repeated  the 
error,  apparently  following  Smith  and 
Taylor  (1950a). 

Hardy  and  McDiarmid  (1969:218)  were 
next  to  discuss  the  problem  and  they 
supported  Hartweg,  Zweifel,  and 
Boulenger's  position.  In  what  I  hope  is 
the  final  chapter  in  this  prolonged  story,  I 
can  only  repeat  and  emphatically  support 
Hartweg's  opinion  (in  Zweifel  1960:95) 
that  K.  integrum  is  "the  only  species  of 
the  genus  that  gets  to  the  Tres  Marias." 
Kinosternon  hirtipes  group. 

Garman  (1887:16)  erroneously  record- 


No.  1 


Kinosternon  Biosystematics 


23 


ed  Cinosternum  hirtipes  from  San  Luis 
Potosi,  Mexico.  Taylor  (1952:793)  re- 
peated that  record,  listing  it  as  "possibly 
doubttui".  Ihe  specimen  on  which 
Carman's  record  was  based  (MCZ  4545), 
from  the  mountains  of  Alvarez,  is  un- 
questionably AT.  integrum.  The  occurrence 
of  K.  hirtipes  in  the  state  of  San  Luis 
PotosC  has  thus  been  verified  at  only  one 
other  locality  (see  MATERIALS  AND 
METHODS). 

J.R.  Dixon  et  al.  (1972:228)  recorded 
K.  hirtipes  from  Cadereyta,  Queretero  on 
the  basis  of  AMNH  71570.  That  speci- 
men, an  articulated  shell,  is  referrable  to 
K.  integrum;  K.  hirtipes  does  not  occur  in 
that  part  of  Quere'taro, 

Liner  (1964:221)  recorded  the  deposi- 
tion in  the  Tulane  collections  of  K. 
hirtipes  he  collected  in  Guanajuato 
(precise  locality  not  published).  TU  17563 
(adult  male)  from  that  collection,  from  2 
mi.  N.  Ojo  de  Agua,  is  not  K.  hirtipes, 
but  K.  integrum. 

Four  papers  (Martin  del  Campo, 
1937:265;  Caballero  y  Caballero,  1938a: 
103,  1938b:448;  Casas  Andreu,  1967:45) 
erroneously  recorded  K.  hirtipes^  from 
Tasquillo,  Hidalgo,  lying  in  the  Rio  Tula 
basin.  Because  only  K.  integrum  occurs  in 
that  basin,  those  records  must  pertain  to 
that  species.  Similarly  Caballero  y  Cabal- 
lero's  (1940a:225)  record  of  K.^  hirtipes 
from  Uruapan,  Michoacan  (Rio  Balsas 
basin)  is  based  on  K.  integrum,  since  only 
the  latter  species  occurs  in  that  area. 

Altini  (1942:154)  recorded  Kinosternon 
hirtipes  from  Lake  Patzcuaro  and  Lake 
Chapala,  Mexico,  and  Lake  Pete'n,  Cuat- 
emala;  the  species  occurs  in  both  of  the 
Mexican  lakes,  but  clearly  does  not  occur 
in  Guatemala.  He  also  recorded  K.  leuco- 
stomum,  K.  cobanum  (=  K.  acutum), 
and  K.  triliratum  (=  K.  scorpioides)  from 
Lake  Pet  en.  All  three  of  those  species  are 
known  from  the  Pete'n  region:  Claudius 
angustatus  and  Staurotypus  triporcatus. 
Which  of  the  latter  two  species  was  mis- 
identified  by  Altinas  K.  hirtipes  cannot 
be  determined  by  the  data  available  to  me. 
In  the  same  paper,  Ahini  also  erroneously 


recorded  K.  leucostomum  from  Mexico's 
Lake  Chapala;  the  species  is  not  found 
there.  Because  only  K.  integrum  and  K. 
hirtipes  occur  in  Lake  Chapala  and 
because  Altini  also  recorded  K.  hirtipes 
from  the  lake  (presumably  correctly  iden- 
tified), his  K.  leucostomum  record  is 
probably  based  on  K.  integrum.  Clearly, 
an  examination  of  Altini's  specimens 
(presumably  at  the  University  of  Bologna) 
will  be  necessary  to  rectify  these  misiden- 
tifications. 

Based  on  my  discussions  with  the 
author,  it  is  clear  that  Wiewandt's 
(1971:34;  and  Wiewandt  et  al.,  1972:162) 
records  of  A^.  sonoriense  from  Sonora,  3.5 
miles  W.  Alamos,  were  based  on  speci- 
mens of  the  recently  described  K. 
alamosae  (K.  sonoriense  does  not  occur 
there).  Similarly,  as  explained  by  Berry 
and  Legler  (1980),  Herenghi's  (1969) 
Sonora  K.  hirtipes  are  also  referable  to  K. 
alamosae. 

Morphometric  Analysis 

An  NTSYS  cluster  analysis  was  per- 
formed early  in  the  study  (1977)  employ- 
ing population  means  for  all  23  variables 
as  OTU's  (males  and  females  separately). 
Two  major  phenetic  groups  were  evident 
in  both  the  male  (Fig.  8)  and  female  dis- 
tance phenograms.  The  first  group  in  each 
analysis  included  the  Casas  Grandes, 
southwest  New  Mexico,  Magdalena, 
Sonora,  Yaqui,  and  Sonoyta  samples 
(i.e.,  the  populations  of  K.  sonoriense  as 
previously  recognized;  Iverson,  1976  and 
1978),  and  the  Aguascalientes  sample  in 
the  male  analysis  and  the  Nazas  sample  in 
the  female  analysis.  Of  later  significance 
is  the  fact  that  the  Sonoyta  sample  was  the 
most  distinct  of  the  sonoriense  population 
in  both  the  male  and  female  analysed.  The 
second  group  in  each  case  included  all 
populations  from  the  Rio  Santa  Maria  in 
Chihuahua  south  and  eastward  to  sout- 
ceral  Mexico  (representing  populations  of 
K.  hirtipes). 

In  both  analyses  the  main  hirtipes 
cluster  was  divided  phenetically  into  two 
subgroups;  however,  the  included  samples 


24 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


were  different  in  each  analysis.  For  both 
sexes,  one  subgroup  included  all  of  the 
northern-most  K.  hirtipes  samples  (Santa 
Marfa,  Carmen,  Sauz,  and  Texas);  but  for 
the  males  the  subgroup  also  included  sev- 
eral of  the  southern-most  populations 
(Valley  of  Mexico,  Villa  Victoria,  and 
Bajio)  and  for  the  females  it  included 
another  northern  population  (Conchos),  a 
central  population  (Mezquital),  and  two 
southern  populations  (Patzcuaro  and 
Chapala).  No  other  obvious  morpho- 
geographic  correlations  or  discontinuities 
were  noted  in  these  preliminary  clusters, 
but  many  of  the  sample  sizes  on  which  the 
means  were  based  were  quite  small.  The 
level  of  differences  between  clusters 
and/or  samples  were  generally  higher  in 
females  than  males,  substantiating  my 
subjective  observation  that  there  is  less 
variation  among  females. 

The  final  male  (Fig.  9)  cluster  analysis 
(BMDP2M)  of  population  means  for  all 
23  variables  for  all  samples  with  N  >  2 
(Appendices  1  and  2)  suggests  that  six 
groups  were  evident.  In  declining  order  of 
distinctiveness  they  are  the  1)  Viesca, 
2)  Sonoyta,     3)  Villa    Victoria,    4)  K. 


Figure  8.  Preliminary  NT-SYS  cluster  (based  on 
the  distance  matrix,  with  complete  averaging  and 
low  values  considered  similar)  of  population 
means  of  all  23  variables  for  males  of  the  K. 
hirtipes  species  group  (including  K.  sonoriense). 
Abbreviations  as  in  text.  N  >  3  for  all  samples 
but  SLP  and  PATZ  (N   =  2  each). 


Figure  9.  BMDP2M  cluster  of  population  means  of 
all  23  character  ratios  (Appendices  1  and  2)  for 
males  of  the  K.  hirtipes  species  group  (including 
K.  sonoriense).  Abbreviations  as  in  text.  N  >  4 
for  all  samples  but  VSCA  (N  =  2)  and  TEX 
(N  =  3).  Numbers  are  amalgamation  distances 
(i.e.,  distance  between  the  clusters  joined). 


sonoriense,  except  Sonoyta,  plus  Aguas- 
calientes,  5)  Duero,and6)  the  remaining 
K.  hirtipes  samples.  In  the  final  female 
analysis,  the  nine  most  distinct  groups  are 
the  1)  San  Juanico  (but  N  =  only  2), 
2)  Sonoyta,  3)  Viesca  (no  female  Villa 
Victoria  sample  was  included),  4)  Balsas 
(N  =  only  2),  5)  Zapotlan  (N  =  2), 
6)  Santiaguillo  (N  =  2),  7)  Duero,  8)  K. 
sonoriense  except  Sonoyta,  plus  Nazas 
and  Verde,  and  9)  the  remaining  K. 
hirtipes  samples.  Again,  the  female  anal- 
ysis differences  were  not  at  as  low  levels  as 
the  males'. 

Stepwise  discriminant  analysis  of  popu- 
lations with  N  >  2,  based  on  all  23 
character  ratios,  produced  plots  of  popu- 
lation means  on  the  first  two  (most  im- 
portant) canonical  axes  for  males  and 
females  (Figure  10).  Two  groups  separate 
along  the  first  canonical  axis  in  both  anal- 
yses: 1)  the  seven  K.  sonoriense  samples 
and  2)  the  K.  hirtipes  samples.  Within  the 
cluster  of  K.  hirtipes  sample  means,  two 
patterns  are  apparent  (especially  along  the 


No.  1 


Kinosternon  Biosystematics 


25 


second  canonical  axis).  First,  the  popula- 
tions are  arranged  from  northern-most 
(Santa  Man'a;  Carmen)  to  central  (Mex- 
quital;  Aguascalientes)  to  southern-most 
(Duero,  Vajio,  Patzcuaro,  and  Chapala); 
except  the  Valley  of  Mexico  sample, 
which  falls  with  the  northern  populations. 
Second,  there  appears  to  be  a  weak 
phenetic  break  in  this  clinal  arrangement 
1)  in  males,  between  populations  north  of 
and  including  the  Nazas  (plus  the  Valley 
of  Mexico)  and  those  south  and  east  of 
that  basin  and  2)  in  females,  (less 
distinctly),  between  populations  north  of 
and  including  the  Conchos  (plus  the 
Valley  of  Mexico)  and  those  south  and 
east.  Other  morpho-geographical  dis- 
continuities include  the  relative  isolation 
of  Duero,  Patzcuaro,  and  Aguascalientes 
samples  (and  the  proximity  of  the  latter  to 
the  K.  sonoriense  samples)  in  the  male 
plot,  and  the  relative  isolation  of  the 
Zapotlan  (but  N  ==  2),  Valley  of  Mexico, 
and  Sonoyta  samples  in  the  female  plot. 

The  character  ratios  most  important  for 
discrimination  of  the  samples  were  deter- 
mined in  the  stepwise  discriminant  anal- 
ysis  in  two  ways:    1)  by  the  highest   F 
values  calculated  for  each  variable  before 
any  were  entered  into  the  discriminant 
function  and  2)  by  the  order  in  which  the 
variables  were  entered  into  the  function. 
The   first   (most   important)   variable   in 
each  is  always  the  same,  but  the  remaining 
frequently  are  not,  especially  if  character 
information  is  redundant  in  two  or  more 
variables.   For  males  the  five  most  im- 
portant    variables     were     PWB/CL, 
PWA/CL,  GW/CL,  BL/CL,  and  IP/PL 
by    F   value,    and    PWB/CL,    IAN/PL, 
IP/CL,  and  BL/CL  by  order  of  inclusion. 
For  females,  they  were  IP/PL,  IP/CL, 
IF/CL,  AN/PL,  and  IF/PL,  and  IP/PL, 
IH/CL,    IF/CL,    BL/CL,    and    FL/CL, 
respectively.  The  analysis  reveals  that  K. 
sonoriense  in  general  has  smaller  inter- 
pectoral  seam  lengths,  larger  interfemoral 
seam  and  gular  scute  lengths,  and  a  wider 
plastron,  gular  scute,  and  bridge  (see  also 
Appendices    3    and    4).    Bivariate    plots 
(Figures    11-14)   of  the  most   important 


characters  (by  F  value)  illustrate  that  K. 
sonoriense  is  both  geographically  and 
morphologically  disjunct.  Because  they 
are  the  most  morphometrically  distinct 
groups  within  the  species  group,  because 
they  are  nowhere  known  to  hybridize, 
because  there  is  no  evidence  of  introgres- 
sion  (based  on  the  morphometric  calcula- 
tions) in  the  two  most  geographically 
proximate  populations  (Casas  Grandes 
and  Santa  Maria),  and  because  several 
other  morphological  characters  (see  later) 
also  show  a  sharp  phenetic  break  between 


OUfB 

**'• 

•                            •CHAT 

•  2»fO 

»&UN 

• 

VSCA*        , 

»GUAS 

• 

»AO 

CNCH      •  '*^ 

• 

v»ilf         SauZ     It! 

• 

StM» 

CSG« 

CIMN 

$TmI 

SAU7 

ca**N 

v«.. 

• 

•' 

C^«:M 

fcAJ 

c^ 

HA2%   aOUAS 

• 

<^'»«,M*OC 

•                          M(Z          • 

AGUN» 

,;»»o 

Figure  10.  Plots  on  the  first  (k,)  and  second  (k,) 
canonical  axes  of  the  population  means  of  males 
(above)  and  females  (below)  of  the  Kinosternon 
hiriipes  species  group  (including  K.  sonoriense). 
Abbreviations  as  in  text.  Analysis  based  on  all  23 
character  ratios  for  populations  with  N  >  2.  First 
two  axes  account  for  30.4  and  20.6%  of  the  total 
variation,  respectively,  in  males;  and  23.8  and 
18.0%,  respectively,  in  females. 


26 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


the  two  morphometric  groups,  K. 
sonoriense  and  K.  hirtipes  are  considered 
valid  species  as  previously  defined  (Iver- 
son,  1976,  1978;  Wermuth  and  Mertens, 
1977).  Therefore,  populations  of  each 
species  were  analyzed  separately. 
Variation  within  K.  sonoriense. 

The  above  analyses  (see  especially 
Figures  9,  10  and  12-14)  suggest  that  the 
Sonoyta  sample  is  the  most  distinct  of  the 
populations  of  K.  sonoriense.  Stepwise 
discriminant  analyses  of  the  seven 
sonoriense  populations  with  N  >  4  (Fig- 
ure 15),  employing  13  variables  (those 
indicated  as  the  most  important  in  the 
overall  analyses)  clearly  support  this 
suggestion. 

Those  plots  also  suggest  additional 
variational  patterns.  Although  most  of 
the  non-Sonoyta  samples  are  morpholog- 
ically very  homogeneous  (note  cluster 
overlap  in  Figure  15),  both  the  male  and 
female  Yaqui  sample  contain  some 
apparently  anomalous  individuals.  In  the 
male  plot,  all  but  one  Yaqui  specimen  lie 
within  the  main  cluster.  The  outlier  (BYU 


39 

J 

V,U 

\ 

NAZ 

* 

*SAU 

J 

<  36 

< 

^ 

ui 

SIMR 

*" 

'Sez 

'"' 

.^ 

Figure  11.  Graph  of  percent  posterior  width  of 
plastral  forelobe/carapace  length  (PWB/CL) 
versus  percent  plastral  width  at  humero-pectoral 
seam/carapace  length  (PWA/CL)  for  males  of  all 
populations  of  the  Kinosternon  hirtipes  species 
group  (including  Kinosternon  sonoriense).  Only 
population  means  are  plotted.  Abbreviations  as 
in  text. 


Figure  12.  Graph  of  percent  posterior  width  of 
plastral  forelobe/carapace  length  (PWB/CL) 
versus  percent  gular  width/carapace  length 
(GW/CL)  for  males  of  all  populations  of  the 
Kinosternon  hirtipes  species  group  (including 
Kinosternon  sonoriense).  Only  population  means 
are  plotted.  Abbreviations  as  in  text. 


14629)  is  the  only  male  available  from  the 
entire  Ri'o  Bavispe  portion  of  the  Yaqui 
basin  and  the  only  specimen  available 
from  that  locality.  The  significance  of  its 
apparent  distinctiveness  must  await  the 
collection  of  additional  material.  The 
most  important  characters  in  the  discrim- 
ination are  IAN/PL,  GW/CL,  IF/PL, 
PL/CL  and  GL/CL  by  F-values,  and 
IAN/PL,  GW/CL,  GL/CL,  IP/PL,  and 
PWC/CL  by  order  of  inclusion. 

In  the  female  plot  all  but  one  of  the 
Yaqui  specimens  lie  outside  the  main 
cluster,  and  separate  fairly  well  along  the 
second  canonical  axis.  Only  a  specimen 
from  the  Rio  Gavilan  (MVZ  46646),  a 
tributary  of  the  Rfo  Bavispe,  falls  within 
the  main  cluster.  Although  the  recorded 
locality  is  clearly  in  the  Gavilan-Bavispe 
drainage  (see  map  in  Marshall,  1957),  the 
possibility  exists  that  the  specimen 
actually  originated  in  the  Rio  Casas 
Grandes  basin;  the  two  drainages  inter- 
digitate  near  the  locality.  The  outlying 
cluster  of  female  Yaqui  specimens  is 
problematical,  especially  since  it  includes 
specimens  from  the  same  localities  that 
clustered  in  the  main  group  in  the  male 


No.  1 


Kinosternon  Biosystematics 


27 


VMLI 

CliNN 

•CMW 

iim 

IWHM 

Til 

^ITOO 

• 

SAUl 

•  AOUU 

MI 

«CIUCi 

• 

•CNCM 

lAJ 

CMM           ,,yyj,f                                   »l»  • 

N» 

,, 

Vu 

SNBA                        AOUH 

,«. 

no> 

• 

^«T. 

Figure  13.  Graph  of  percent  interanal  seam 
length/carapace  length  (lAN/CL)  versus  percent 
interpectoral  seam  length/carapace  length 
(IP/CL)  for  females  of  all  populations  of  the 
Kinosternon  hirtipes  species  group  (including 
Kinosternon  sonoriense).  Only  population  means 
are  plotted.  Abbreviations  as  in  text. 


analysis.  Only  additional  field  work  in  the 
Yaqui  basin  can  clarify  these  variational 
anomalies.  The  most  important  charac- 
ters in  the  discrimination  are  IF/PL, 
lAN/CL,  PL/CL,  IH/PL,  and  PWB/CL 
by  F-values  and  IF/PL,  PL/CL,  IH/PL, 
PW/CL,  and  IP/PL  by  order  of  inclu- 
sion. 

Two  morphotypes  are  thus  recogniz- 
able within  K.  sonoriense,  represented  by 
Sonoyta  basin  turtles  and  the  other 
samples,  respectively.  The  Sonoyta  basin 
turtles  have  shorter  plastra,  longer  inter- 
femoral  seams,  shorter  interanal  seams, 
wider  first  vertebrals,  and  narrower  gulars, 
than  the  other  populations. 

Variation  within  K.  hirtipes 

Cluster  analysis  (BMDP2M)  of  popu- 
lation means  for  all  23  variables,  suggest 
that  three  main  groups  exist  for  males  and 
females  (Fig.  16).  For  males,  the  most  dis- 
tinct is  the  Viesca  sample  (but  N  =  2), 
followed  by  the  Villa  Victoria  sample  and 
all  remaining  hirtipes  populations.  Within 
the  last  group,  the  only  relationship  be- 
tween phenetics  and  geography  is  a  small 


SNIA* 

JN«»^ 

»Ao' 

MAGO,           •AGON 

.NAZ 

«MT2 
'STGO 

SWNM* 

*CMCM 

uuz 

SNJ. 

STMI 

•stf 

CHAT 

.CtMN 

•miii 

Figure  14.  Graph  of  percent  interpectoral  seam 
length/carapace  length  (IP/CL)  versus  percent 
interfemoral  seam  length/carapace  length 
(IF/CL)  for  females  of  all  populations  of  the 
Kinosternon  hirtipes  species  group  (including 
Kinosternon  sonoriense).  Only  population  means 
are  plotted.  Abbreviations  as  in  text. 


subgroup  including  the  Patzcuaro,  San 
Juanico,  Chapala,  and  Duero  samples. 
Contrary  to  the  results  of  the  discriminant 
analysis  which  included  K.  sonoriense 
(Fig.  10),  no  clinal  relationships  or 
phenetic  breaks  between  northern  and 
southern  populations  are  evident.  For 
females,  the  most  distinct  group  is  the  San 
Juanico  sample  (but  N  =  2),  followed  by 
the  Viesca  sample  and  the  remaining 
populations.  No  obvious  phenetic- 
geographic  relationships  are  suggested 
within  the  last  group.  However,  Villa 
Victoria  was  not  represented  in  the  female 
analysis  since  only  one  female  specimen  is 
known  from  that  basin.  Raw  data  for  that 
specimen  (Appendix  2)  suggest  it  might  be 
as  distinct  as  the  male  sample  (see  later). 

Stepwise  discriminant  analysis  of  K. 
hirtipes  populations  with  N  i  2,  based 
on  all  23  character  ratios,  produced  the 
canonical  plots  in  Figure  17.  The  male 
analysis  reveals  the  same  north-south 
clinal  pattern  along  the  first  axis,  the  same 
phenetic  break  between  the  Nazas  and 
Aguanaval  basins,  and  the  same  similarity 
between  the  Valley  of  Mexico  population 
and  northern  populations  that  the  earlier 


28 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


overall  discriminant  analysis  illustrated 
(Fig.  10).  It  further  suggests  the  distinct- 
iveness of  the  Villa  Victoria  population 
and  possibly  also  the  Patzcuaro  sample. 
The  female  analysis  again  reveals  the 
general  north-south  clinal  pattern  along 
the  first  axis  (ahhough  no  Nazas- 
Aguanaval  break  is  evident),  and  also 
suggests  the  distinctiveness  of  the  Viesca, 
Zapotlan,  San  Juanico  and  possibly  Duero 
populations. 


Figure  15.  Plots  on  the  first  (k,)  and  sec  ond  (k2) 
canonical  axes  of  population  means  (solid  dots) 
of  males  (above)  and  females  (below)  of  K. 
sonoriense.  Lines  connect  the  most  dispersed 
values  about  the  population  mean.  Population 
mean  symbols  are  1,  SNTA;  2,  SWNM;  3,  GILA 
and  WILL;  4,  MAGD;  5,  SNRA;  6,  CSGR;  and 
7,  YAQ.  Individual  Yaqui  turtles  are  marked 
with  Y  (Yaqui  River  proper)  or  B  (Bavispe  River). 
Analysis  based  on  13  character  ratios.  First  two 
axes  account  for  71 .0  and  10.2%  of  the  variation, 
respectively,  in  males;  and  47.9  and  23.1%,  re- 
pectively,  in  females. 


The  most  important  characters  in  the 
male  discriminant  analysis  are  BL/CL. 
GL/CL,  PWC/CL,  IP/CL,  and  IAN/PL, 
based  on  F-values  and  BL/CL,  GL/CL, 
PWC/CL,  IAN/PL,  and  PWD/CL  based 
on  order  of  inclusion.  For  the  females,  the 
important  characters  are  IP/CL,  BL/CL, 
IP/PL,  PWB/CL,  and  FL/CL,  and 
IP/CL,  BL/CL,  IH/CL,  FL/CL,  and 
PWB/CL,  respectively.  Bivariate  plots  of 
the  most  important  characters  (Figures 
18-21;  see  also  Figures  11-14)  do  not 
suggest  that  a  significant  phenetic  break 
exists  between  northern  and  remaining 
populations.  However,  as  in  the  previous 
analyses,  they  again  indicate  the  distinct- 
iveness of  several  samples,  including  the 
Valley  of  Mexico,  Viesca,  San  Juanico, 
Patzcuaro  samples,  and  possibly  also  a 
group  of  three  geographically  adjacent 
samples  occupying  the  ancestral  Lake 
Chapala  basin  (Tamayo,  1964:108): 
Chapala,  Zapotlan  and  Duero  (see  Figs. 
18  and  21). 

MORPHOMETRIC  CONCLUSIONS 

The  numerous  analyses  have  strongly 
suggested  that  1)  K.  hirtipes  and  K.  sono- 
riense are  distinct  morphometrically,  2) 
the  Sonoyta  sample  within  K.  sonoriense 
is  morphometrically  distinct,  3)  there  is 
tremendous  variation  within  K.  hirtipes, 
and  4)  the  most  morphometrically  distinct 
populations  of  hirtipes  are  the  San  Juan- 
ico, Viesca,  Patzcuaro,  Villa  Victoria, 
and  Valley  of  Mexico  samples  and 
possibly  also  the  combined  Chapala-Za- 
potlan-Duero  samples. 

As  detailed  above,  the  basic  patterns  of 
morphometric  variation  in  the  K.  hirtipes 
species  group  involve  several  character 
complexes,  the  most  important  of  which 
are  1)  body  size  (see  later),  2)  relative  plas- 
tron size  (measured  primarily  as  PL, 
PWA,  PWB,  PWC,  and/or  PWD),  3) 
relative  bridge  length,  4)  the  relative 
lengths  of  the  gular,  interhumeral  seam, 
and  interpectoral  seam  to  the  forelobe 
length  (the  forelobe  length  itself  is  not  as 
important),  and  5)  (of  much  less  impor- 
tance) the  relative  lengths  of  the  inter- 


No.  1 


Kinoslcrnon  Biosyslcmatics 


29 


femoral  and  interanal  seams  to  the  hind- 
lobe  length  (the  hindlobe  length  is  also  not 
as  important).  Other  characters  clearly 
are  important  in  individual  population 
comparisons,  but  these  complexes  are  ap- 
parently the  most  important  when  consid- 
ering variation  in  the  group  as  a  whole. 

Variation  in  relative  plastron  size  is 
much  greater  in  males  than  females. 
Females  appear  to  be  much  more  conser- 
vative regarding  plastral  reduction  or 
modification.  For  relative  male  plastron 
size  there  is  somewhat  of  a  continuum 
from  the  relatively  large  plastron  of  K. 
sonoriense  (Fig.  22b)  to  the  small  plastra 
of  San  Juanico  and  Viesca  populations 
(Fig.  22,  1  and  m)  with  the  remaining 
populations  somewhat  intermediate.  For 
females,  the  range  is  from  the  medium- 
size  plastra  of  the  San  Juanico  and  Viesca 
populations  (Fig.  22n)  to  the  relatively  ex- 
tensive plastra  of  the  remaining  popula- 
tions. Plastral  reduction,  typically  cor- 
related with  an  increase  in  aquatic  habits 
among  turtles  (Zangerl,  1939:386;  Berry, 
1977;  Iverson,  MSI)  and  presumably  an 
adaption  thereto  (Iverson,  MSI),  is  con- 
sidered derived. 


Relative  bridge  length  is  extremely  vari- 
able in  the  genus  Kinosternon.  Males  vir- 
tually always  have  shorter  bridges  than  fe- 
males. For  males,  bridge  length  ranges 
from  short  in  San  Juanico,  Pa'tzcuaro, 
Valley  of  Mexico,  and  Viesca  turtles  to 
medium  length  in  the  other  populations. 
For  females,  it  ranges  from  medium 
length  in  San  Juanico,  Patzcuaro,  and 
Valley  of  Mexico  turtles  to  long  (but  not 
as  long  as  some  members  of  the  K.  scor- 
pioides  group)  in  the  remaining  popula- 
tions. Its  reduction  is  not  necessarily  cor- 
related with  plastral  reduction  [for  exam- 
ple, Patzcuaro  turtles  have  medium 
(male)  or  large  (female)  plastra  and  rela- 
tively short  bridges].  I  consider  relatively 
reduced  bridge  length  in  males  or  females 
a  derived  character,  both  because  of  its 
rarity  in  this  species  group  and  because 
many  of  the  most  specialized  members  of 
the  genus  have  short  bridges. 

The  components  of  the  plastral  fore- 
lobe  are  quite  variable  in  this  species 
group,  but  because  the  interhumeral  seam 
length  is  essentially  of  medium  length  in 
all  samples  (except  perhaps  in  the  Villa 
Victoria  basin),  variational  patterns  are 


^ 


•-c 


-c 


STMR 

CRMN 

VALLE 

PAP 

CHAP 

DUER 

PATZ 

AGUN 

CNCH 

AGUAS 

SLP 

MEZ 

VERD 

NAZ 

ZAPO 

SAUZ 

VSCA 

SNJ 


I       I      I       I      I »— I 1      I       I 


)  O  O  >OCJ  CJ  NJ    ~- 


Figure  16.  BMDP2M  cluster  of  population  means  of  all  23  character  ratios  (Appendices  1  and  2)  for  male 
(top)  and  female  K.  hirtipes.  Abbreviations  as  in  text.  For  males,  N  >  4  for  all  samples  except  VSCA 
(N  =  2);  for  females.  N  >  5,  except  SNJ  (N  =  2)  and  ZAPO  (N  =  2).  Numbers  are  amalgamation 
distances  (i.e.,  distance  between  the  clusters  joined). 


30 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


dominated  by  the  relative  lengths  of  the 
gular  and  the  interpectoral  seam. 
Patzcuaro,  Viesca,  and  San  Juanico  tur- 
tles have  very  short  gulars  (Fig.  22,  i-n), 
whereas  the  remaining  populations  have 
medium  to  long  gulars  {K.  sonoriense 
having  the  longest).  Patzcuaro  and  San 
Juanico  turtles  (and  possibly  Villa  Vic- 
toria) have  the  longest  interpectoral  seams 
as  well  (Fig.  22,  i-1);  K.  sonoriense,  the 
shortest;  and  the  remaining  populations 
have   intermediate   lengths.    Since   most 


Figure  17.  Plots  on  the  first  (k,)  and  second  (kj) 
canonical  axes  of  the  population  means  (solid 
dots)  of  males  (above)  and  females  (below)  of 
Kinosternon  hirtipes  (excluding  K.  sonoriense). 
Abbreviations -as  in  text  for  females;  but  further 
shortened  for  males.  Polygons  in  male  plot 
enclose  total  dispersion  of  each  population. 
Analysis  based  on  all  23  character  ratios  for 
populations  with  N  >  2.  First  two  axes  account 
for  33.7  and  16.0%  of  the  total  variation,  re- 
spectively, in  males;  and  27.4  and  14.5%, 
respectively,  in  females. 


Kinosternon  have  interpectoral  seams  of 
medium  length  (frequently  used  to  diag- 
nose the  genus;  e.g.,  Conant,  1975), 
deviations  from  that  condition  are  consid- 
ered derived.  For  similar  reasons,  the  con- 
ditions of  reduced  and  extensive  gular 
scutes  are  considered  derived. 

The  plastral  hindlobe  components  do 
not  show  as  much  variation  as  the  other 
complexes,  but  a  few  patterns  are  evident. 
Hindlobe  length  is  greater  in  K.  sono- 
riense and  Villa  Victoria  turtles  than  in  re- 
maining populations.  The  interfemoral 
seam  is  relatively  shorter  in  the  Valley  of 
Mexico  sample  (Fig.  22,  e-f)  than  in  all 
other  samples;  and  the  interanal  seam  is 
relatively  longer  in  that  sample  and  the 
Chapala-Zapotlan  combined  sample,  and 
shorter  in  the  Viesca  sample.  Variation  in 
these  characters  within  K.  sonoriense 
(i.e.,  shorter  interanal  and  longer  inter- 
femoral seams  in  Sonoyta  turtles)  has 
already  been  discussed.  Each  of  these 
deviations  from  the  modal  hindlobe  con- 
dition found  in  the  group  are  considered 
derived. 

Other  Morphological  Characters 
Nasal  scale.  —  The  cornified  epidermal 

shield  (here  called  the  nasal  scale)  found 


Figure  18.  Graph  of  percent  gular  length/carapace 
length  (GL/CL)  versus  percent  bridge 
length/carapace  length  (BL/CL)  for  males  of 
populations  of  Kinosternon  hirtipes.  Only  popu- 
lation means  are  plotted.  Abbreviations  as  in 
text. 


No. 


Kinosternon  Biosystematics 


31 


• 

•  DUEH 

■ 

CH«P 

«UN« 

ilu;  4  Mf; 

•s,.         ^ 

'•CNCM 

ZAPO 

•^ 

., 

•p.i/ 

VSCA* 

Figure  19.  Graph  of  percent  interpectoral  seam 
length/carapace  length  (IP/CL)  versus  percent 
bridge  length/carapace  length  (BL/CL)  for  males 
of  populations  of  Kinosternon  hirtipes.  Only 
population  means  are  plotted.  Abbreviations  as 
in  text. 


on  the  anterior  dorsum  of  the  head  of  all 
subadult  and  adult  kinosternid  turtles  has 
received  little  attention  (but  see  Sieben- 
rock,  1907)  until  recently  (Conant  and 
Berry,  1978).  Cornification  of  the  scale 
begins  near  the  rostrum  in  the  juveniles  of 
these  turtles  and  spreads  posteriorly  with 


age.  Development  is  usually  more  rapid 
laterally  than  medially,  such  that  older 
juveniles  or  subadults  may  have  V-shaped 
scales  even  though  adults  might  have  tri- 
angular to  rhomboidal  scales.  Due  to  its 
distinctive  shape  in  the  adults  of  kino- 
sternid turtle  populations,  it  is  an  impor- 
tant taxonomic  character.  Intraspecific 
variation  in  nasal  scale  shape  has  already 
been  noted  in  Kinosternon  subrubrum. 
Ernst  et  al.  (1974)  have  shown  that  the 
subspecies  Kinosternon  subrubrum  stein- 
dachneri  usually  (81*7o)  has  a  head  scale 
that  is  furcate  posteriorly  (their  "divided 
nasal")  whereas  turtles  of  both  other  sub- 
species (hippocrepis  and  subrubrum)  usu- 
ally (90  to  97%)  have  a  non-furcate,  bell- 
shaped  scale  (posterior  margin  convex). 

Variation  within  the  K.  hirtipes  species 
group  is  considerable  but  intrapopulation 
variation  is  insignificant.  Three  distinctive 
nasal  scale  shapes  are  evident  in  inter- 
population  comparisons  of  adults  of  this 
group.  Populations  from  the  Rios  Casas 
Grandes  and  Yaqui  (excluding  the  Papi- 
gochic)  and  those  westward  and  north- 
ward (i.e.,  populations  of  K.  sonoriense 
as  previously  recognized;  Iverson,  1976), 
and  the  population  from  the  Valley  of 


DUER 

CHAP* 

■ 

•aGun 

■ 

VSCA^ 

Zapo         J  ^^^ 

.C.MN 

• 

--• 

*"" 

I 

-'. 

Figure  20.  Graph  of  percent  interpectoral  seam 
length/carapace  length  (IP/CL)  versus  percent 
bridge  length/carapace  length  (BL/CL)  for 
females  of  populations  of  Kinosternon  hirlip  s. 
Only  populations  means  are  plotted.  Abbrevi- 
ations as  in  text. 


Figure  21.  Graph  of  percent  gular  length/carapace 
length  (GL/CL)  versus  percent  bridge 
length/carapace  length  (BL/CL)  for  females  of 
populations  of  Kinosternon  hirtipes.  Only 
population  means  are  plotted.  Abbreviations  as 
in  text. 


32 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


No. 


Kinosternon  Biosystematics 


33 


Figure  22.  Plastral  variation  in  members  of  the  Kinosternon  hirtipes  species  group:  Gila  River  basin 
Kinosternon  sonoriense,  JBI  563  female  (a)  and  JBI  386  male  (b);  Rio  Papigochic  basin  K.  hirtipes 
murrayi,  UF  40391  female  (c)  and  UF  40396  male  (d);  Valley  of  Mexico  A',  h.  hirtipes,  UMMZ  99458 
female  (e)  and  UMMZ  80357  male  (0;  Lake  Chapala  basin  K.  h.  chapalaense,  UMMZ  97123  female  (g) 
and  UMMZ  97128  male  holotype  (h);  Lake  Patzcuaro  K.  h.  tarascense,  UF  43505  female  (i  and  j; 
illustrating  plastral  scute  staining  and  loss  of  stain  with  scute  shedding)  and  UF  43506  male  (k);  Presa  San 
Juanico  A',  h.  magdalense.  UF  45035  male  holotype  (1);  Viesca  A',  h.  megacephahim  SM  1 1464  female  (m) 
and  SM  9823  male  (n). 


34 


Tulane  Studies  in  Zoology  and  Botany 


Vol.23 


Mexico  exhibit  a  triangular,  rhomboidal 
or  bell-shaped  nasal  scale  as  adults  (Fig. 
23:  a  and  d).  Turtles  from  the  Zapotlan, 
Lake  Chapala,  and  Rfo  Duero  basins  pos- 
sess a  nasal  scale  that  typically  is  crescent- 
shaped  and  extremely  reduced  in  size.  It 
nearly  always  lies  completely  anterior  to 


the  orbits  in  turtles  from  the  former  two 
basins  (Fig.  23:  c),  but  may  reach  to  mid- 
orbit  in  Ri'o  Duero  turtles  (Fig.  23:  0-  All 
remaining  populations  of  this  group  have 
a  nasal  scale  deeply  furcate  behind  (Fig. 
23:  b  and  e;  but  see  Synthesis). 

Because  most  of  the  members  of  the 


Figure  23.  Nasal  scale  variation  in  members  of  the  Kinosternon  hiriipes  species  group:  A",  sonoriense,  JBI  697 
(a);  A',  hiriipes  murrayi,  UF  40396  (b);  A',  h.  chapalaense,  UMMZ  97130  paratype  (c);  A',  h.  hiriipes, 
UMMZ  99449  (d);  A.  h.  larascense,  UF  43505  paratype  (e);  A'.  //.  chapalaense  x  A',  h.  murravi,  UMMZ 
97135  (f). 


No.  1 


Kinosicrnon  Biosystemalics 


35 


genus  Kinosternon  have  triangular  or  bell- 
shaped  nasal  scales  as  adults  (A',  dunni,  K. 
angustipons,  K.  acutum,  K.  baurii,  most 
A',  subrubrum,  and  all  members  of  the  A. 
scorpioides  and  A.  leucosiomum  species 
groups),  and  because  1  believe  that  the 
furcate  condition  in  A.  subrubrum  slein- 
dachneri  is  derived  from  the  bell-shaped 
condition  found  in  a  A.  subrubrum  subru- 
brum-WkQ  ancestor  (see  also  Ernst  et  al, 
1974),  1  consider  the  large  nonfurcate 
shape  to  be  the  primitive  adult  condition 
in  the  genus  Knujsternon.  Therefore,  the 
condition  in  A.  sonuriense  and  Valley  of 
Mexico  A.  luriipes  is  considered  primi- 
tive, whereas  the  nasal  scale  reduction  in 
remaining  populations  of  A.  hirtipes  is 
considered  derived. 

Chin  Barbels.  —  Variable  numbers  of 
barbels  are  present  on  the  chin  and/or 
gular  region  of  all  kinosternid  turtles; 
however,  two  basic  patterns  exist  in  the  K. 
hirtipes  species  group.  In  the  first,  char- 
acteristic of  all  populations  of  K.  sono- 
riense  as  previously  defined  (Iverson, 
1976,  1978),  3  or  4  pairs  of  barbels  are 
present  and  the  largest  2  pairs  are  sub- 
equal  and  relatively  long  (  <  one  half 
orbit  diameter)  with  one  pair  mentally  lo- 
cated and  the  other  at  the  level  of  mid- 
tympanum. 

Populations  of  K.  hirtipes  are  charac- 
terized by  the  presence  of  at  least  two 
pairs  of  chin  barbels,  the  largest  two  pairs 
both  located  on  the  chin  with  the  anterior 
pair  decidedly  the  largest  (yet  P  half  orbit 
diameter).  Because  the  typical  Kinoster- 
non condition  is  one  with  two  mental 
pairs  of  barbels,  the  condition  in  K.  sono- 
riense  is  considered  derived  and  that  of  AT. 
hirtipes  ancestral. 

Head  color.  —  Head  patterns  in  this 
group  are  extremely  variable,  even  within 
populations.  Patterns  vary  nearly  con- 
tinuously from  broadly  mottled  (common 
in  K.  sonoriense;  Fig.  24b;  Conant  and 
Berry,  1978,  Fig.  7)  to  medium  or  finely 
reticulated  (as  in  most  populations  of  K. 
hirtipes  from  Chihuahua  to  Mexico  City; 
Fig.  24:  e,  f,  g,  and  n;  Conant  and  Berry, 
1978,  Fig.  7)  to  finely  spotted  (common  in 


female  Patzcuaro  A.  hirtipes;  Fig.  24:  h). 
Whatever  the  general  pattern,  however, 
the  lateral  markings  are  more  or  less  lon- 
gitudinally oriented,  such  that  two  yellow, 
cream,  or  white  lateral  stripes  (one  ex- 
tending across  the  temporal  region,  poste- 
riorly from  the  posterodorsal  margin  of 
the  orbit;  and  the  other  extending  poste- 
riorly from  the  posteroventral  margin  of 
the  orbit,  along  the  posterodorsal  margin 
of  the  maxillary  sheath  to  the  angle  of  the 
jaw)  are  vaguely  to  very  well  developed. 
The  more  ventral  of  those  stripes  is  almost 
always  apparent,  no  matter  how  finely 
reticulated  or  spotted  the  pattern,  or  me- 
lanistic  the  head  coloration.  Most  of  this 
general  range  of  pattern  variation  may 
occur  in  a  single  population;  however, 
females  usually  have  less  dark  pigment  on 
the  head,  have  finer  mottling  or  reticula- 
tions, and  are  more  likely  to  be  spotted 
(compare  Fig.  24:  a  versus  b  or  e  versus  f; 
see  also  Conant  and  Berry,  1978,  Fig.  7). 
The  jaw  sheaths  are  also  variably 
pigmented,  but  in  general  the  more  dark 
pigment  on  the  head,  the  more  darkly  pig- 
mented (streaked)  are  the  jaw  sheaths. 

The  only  two  significant  deviations 
(considered  derived  conditions)  from  this 
general  (considered  primitive)  color 
scheme  are  in  K.  hirtipes  from  the  Lake 
Chapala  and  Zapotlan  basins  and  the  Val- 
ley of  Mexico  basin.  Turtles  from  the  lat- 
ter basin  have  typical  amounts  of  dark 
pigment  but  most  specimens  have  both 
light  lateral  head  stripes  very  well-defined 
(Fig.  24:  c  and  d).  In  the  former  two 
basins,  the  dark  markings  are  generally 
broad,  but  the  overall  amount  of  dark 
pigment  is  significantly  reduced  (compare 
Figure  24b  versus  i-1);  in  other  popula- 
tions broadness  of  marking  is  correlated 
with  abundance  of  dark  pigment.  In  addi- 
tion, in  Chapala  and  Zapotlan  turtles,  the 
lateral  temporal  head  stripe  is  typically 
bordered  ventrally  by  a  broad  dark  stripe 
and  the  ventral  stripe  is  bordered  dorsally 
by  a  similar  dark  stripe.  The  general 
appearance  is  one  of  two  dark  stripes 
rather  than  two  light  ones  (Fig.  24:  i-1). 

Although  their  nasal  scales  are  similar 


36 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


No.  1 


Kinosternon  Biosystematics 


37 


K.  -  * ''  ^^ Y*^ '  ''*"; 


W 


Figure  24.  Head  pattern  variation  in  members  of  the  Kinosternon  hirtipes  species  group:  Gila  River  Basin 
Kinosternon  sononense,  JBl  563  female  (a)  and  JBI  387  male  (b);  Valley  of  Mexico,  A',  hirtipes  hirtipes, 
UMMZ  99458  female  (c)  and  UMMZ  80357  male  (d);  Rio  Papigochic  A',  h.  murrayi.  UF  40391  female  (e) 
and  UF  40395  male  (0;  Patzcuaro  basin  A.  h.  tarastense,  UF  43596  female  paratype  (g)  and  AMNH  82628 
female  (h);  Lake  Chapala  basin,  A.  h.  chapalaense,  UMMZ  97128  male  holotype  (i),  UMMZ  97127  male 
paratype  (j),  UMMZ  97123  female  paratype  (k),  and  UMMZ  97130  male  paratype  (1);  Viesca,  Coahuila 
K.  h.  megacephatum,  SM  11462  female  paratype  (m,n). 


38 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


to  Chapala-Zapotlan  turtles,  specimens 
from  the  Rfo  Duero  have  much  darker 
head  pigment  as  in  more  northerly  and 
easterly  populations. 

Head  size.  —  Only  one  population  de- 
viates from  the  typical  (clearly  primitive) 
condition.  Turtles  from  the  Viesca  basin 
have  distinctly  enlarged  heads  (especially 
females)  with  greatly  expanded  alveolar 
surfaces  (Fig.  24:  m-n). 

Plastral  staining.  —  Although  the  plas- 
tron of  members  of  the  K.  hirtipes  group 
is  typically  cream,  yellow,  or  yellow 
orange,  it  may  be  variably  stained  dark 
brown  to  black.  The  turtles  from  Lake 
Patzcuaro  have  plastra  consistently  (and 
uniquely?)  stained  red-brown.  At  least  in 
that  population  the  color  is  due  to  envi- 
ronmental staining  since  the  color  is  shed 
with  the  scute  (Fig.  22:  i-j). 

Shell  carination.  —  Due  to  sexual  dif- 
ferences and  considerable  ontogenetic 
change,  quantification  of  variation  in  the 
development  of  keels  in  members  of  this 
group  is  difficult.  In  general,  adult  K. 
sonoriense  are  much  more  obviously  tri- 
carinate  than  K.  hirtipes.  The  latter 
species  often  appears  almost  unicarinate, 
the  former,  very  rarely.  The  dorsum  of 
the  shell  thus  has  a  flatter  appearance  in 
K.  sonoriense  than  in  K.  hirtipes. 

Body  size.  —  Average  carapace  lengths 
of  males  and  females  in  Appendix  1  and  2 
reveal  that  females  average  larger  than 
males  in  populations  of  K.  sonoriense, 
whereas  males  average  larger  in  most  pop- 
ulations of  K.  hirtipes.  There  is  also  con- 
siderable variation  in  body  size  among 
populations  of  K.  hirtipes.  The  most  ob- 
vious deviations  from  the  mode  are  in  tur- 
tles from  the  Viesca  and  San  Juanico 
basins.  Although  these  basins  both  have  a 
small  sample  size,  I  believe  the  data  truly 
reflect  the  small  size  of  the  inhabitant  tur- 
tles. Patzcuaro  turtles  also  tend  to  be 
smaller  than  the  mode,  although  not  so 
distinctly.  A  more  complete  analysis  of 
population  and  sexual  variation  in  body 
size  in  K.  hirtipes  is  in  preparation. 

Nuchal-neural  bone  contact.  —  Berry 
and    Legler    (1980:11)    report    that    the 


nuchal  bone  contacts  the  first  neural  bone 
in  73%  of  the  K.  sonoriense  and  only  4% 
of  the  K.  hirtipes  they  examined  (sample 
sizes  not  reported).  However,  only  38.1<^o 
of  the  K.  sonoriense  (N  =  42)  and 
10.2%  of  the  K.  hirtipes  (N  =  98)  I 
examined  have  nuchal-neural  contact. 
The  discrepancy  between  our  results  for 
K.  sonoriense  is  possibly  due  to  their 
smaller  sample  size,  but  the  character  is 
obviously  of  only  limited  value  in  distin- 
guishing the  two  species. 

Synthesis  Of  Character  Variation 

Analysis    of   geographic    variation   in 
morphological  characters  in  the  Kinoster- 
non  hirtipes  species  complex  supports  the 
recognition  of  two  allopatric  species,  both 
previously    recognized    (Wermuth    and 
Mertens,  1977;  among  many  others):  K. 
sonoriense  and  K.  hirtipes.  Analysis  of 
populations  of  K.  sonoriense  suggests  the 
existence  of  two  distinct  morphotypes, 
represented  by  1)  the  population  inhabit- 
ing the  Rio  Sonoyta  basin  and  2)  the 
remaining  populations  previously  recog- 
nized as  K.  sonoriense  (Fig.  3).  Stepwise 
discriminant  analysis  of  those  two  sam- 
ples using  only  13  morphometric  charac- 
ters is  capable  of  distinguishing  100%  of 
the  males  and  98.6%  of  the  females.  Be- 
cause the  holotype  oi  K.  sonoriense  was 
collected  in  the  Gila  River  basin  (Iverson, 
1976)  the  Rfo  Sonoyta  population  is  here 
described  as  a  new  subspecies. 

Patterns  of  geographic  variation  in 
morphology  within  Kinosternon  hirtipes 
suggest  the  existence  of  several  undes- 
cribed  taxa  (Fig.  4).  The  most  distinct 
morphological  samples  in  this  highly  vari- 
able species  are  the  1)  Viesca,  2)  San  Juan- 
ico, 3)  Pa'tzcuaro,  4)  Valley  of  Mexico,  5) 
Chapala-Zapotlan,  6)  possibly  the  Duero 
sample,  7)  possibly  the  Villa  Victoria  pop- 
ulation (see  below)  and  8)  the  remaining 
populations  of  K.  hirtipes.  Stepwise  dis- 
criminant analysis  of  the  seven  samples, 
excluding  the  Villa  Victoria  population 
(see  below),  using  all  23  morphometric 
variables,  was  able  to  classify  turtles  into 
the  correct  morphotype  at  least  75%  of 


No.  1 


Kinosternon  Biosystematics 


39 


the  time  for  any  given  morphotype  of 
either  sex.  San  Juanico  and  Viesca  turtles 
were  always  classified  correctly,  and  only 
one  Valley  of  Mexico  turtle  was  misclassi- 
fied  (a  female,  into  sample  8,  above).  A 
single  male  and  one  female  from  Patz- 
cuaro  were  misclassified  (into  San  Juan- 
ico, in  both  cases).  Two  males  and  one 
female  Duero  turtle  were  misclassified 
into  the  Chapala-Zapotlan  sample;  and 
two  other  male  Duero  turtles  were  mis- 
classified in  the  Patzcuaro  sample. 
Chapala-Zapotlan  turtle  misclassification 
included  three  males  and  one  female  into 
the  Patzcuaro,  and  one  female  into  the 
Viesca  sample.  The  large  and  highly  vari- 
able sample  of  the  remaining  K.  hirtipes 
populations  included  the  following  mis- 
classifications:  12  males  and  13  females 
into  the  Chapala-Zapotlan  sample;  7 
males  and  6  females  into  the  Valley  of 
Mexico  sample;  9  males  and  two  females 
into  the  Duero  sample;  seven  females  into 
the  Patzcuaro  sample  and  three  females 
into  the  Viesca  sample. 

Based  on  the  various  morphological 
analyses,  I  conclude  that  at  least  the  fol- 
lowing samples  should  be  recognized  tax- 
onomically:  1)  Viesca,  2)  San  Juanico,  3) 
Patzcuaro,  4)  Chapala-Zapotlan,  5)  Val- 
ley of  Mexico,  and  6)  the  remaining  popu- 
lations of  K.  hirtipes  (perhaps  excluding 
the  Villa  Victoria  sample).  I  tentatively 
consider  the  Duero  population  as  inter- 
grading  between  samples  4  and  6.  The 
first  four  samples  have  not  been  named 
and  are  described  here.  The  holotype  of 
Kinosternon  hirtipes  murrayi  clearly  be- 
longs in  the  last  group  and  hence  that 
group  should  bear  that  trinomen.  The  sta- 
tus of  the  Valley  of  Mexico  and  Villa  Vic- 
toria samples  are  not  as  clear. 

Several  of  the  early  analyses  (see  Fig- 
ures 9,  11,  16,  and  17)  suggested  that  the 
male  Villa  Victoria  sample  was  morpho- 
metrically  distinct.  Unfortunately,  only 
one  female  is  known  from  that  basin,  and 
although  not  as  distinct  (see  Appendix  2), 
it  does  exhibit  some  of  the  characters 
which  seem  to  distinguish  the  males  (long- 
er hindlobe,  shorter  interhumeral  seam, 


longer  interpectoral  seam,  and  longer  first 
vertebral  scute).  However,  the  complete 
lack  of  geographically  proximate  com- 
parative material  from  the  entire  upper 
Lerma  system  and  the  near  lack  of  mate- 
rial from  the  Balsas  drainage  system  (one 
female  from  45  miles  to  the  west)  make  a 
decision  regarding  the  distinctiveness  of 
this  population  difficult.  I  have  therefore 
tentatively  included  the  population  with 
those  of  K.  h.  murrayi  until  field  work  in 
the  upper  Balsas  and  Lerma  basins  can 
clarify  distribution  and  morphological 
variation  in  those  regions. 

Even  less  clear  is  the  correct  allocation 
of  the  holotype  o{  Kinosternon  hirtipes,  a 
very  old,  worn  male  specimen,  with  no 
associated  data  except  "Mexico"  (see 
SYSTEMATIC  ACCOUNTS).  Plastral 
erosion  makes  clear  morphometric  alloca- 
tion impossible  (Fig.  25).  In  addition,  the 
shape  of  the  nasal  scale  (Fig.  25)  is  some- 
what intermediate  between  a  V-shaped 
condition  of  A^.  h.  murrayi  and  the  rhom- 
boidal  condition  of  Valley  of  Mexico  tur- 
tles. Schmidt  (1953)  restricted  the  type- 
locality  to  "lakes  near  Mexico  City"  (in 
the  Valley  of  Mexico)  but  without  varia- 
tional analyses.  Because  the  correct  al- 
location can  only  be  solved  by  field  work 
in  the  Valley  of  Mexico  and  adjacent  ba- 
sins, I  tentatively  follow  Schmidt  (1953)  in 
the  application  of  the  name  K.  h.  hirtipes 
to  the  populations  in  that  Valley. 

Systematic  Accounts 

A  chronological  list  of  all  synonyms 
and  orthographic  variations  thereof  is 
given  for  each  valid  taxon.  Each  ortho- 
graphic combination  appears  with  refer- 
ence to  its  first  use,  including  author,  date 
and  pagination  (referenced  in  Literature 
Cited).  Selected  subsequent  usages,  espe- 
cially those  incorrect  or  of  taxonomic  sig- 
nificance, and  including  all  pre- 1930  ref- 
erences, are  included  in  species  and  sub- 
species synonymies.  Most  citations  are  al- 
so parenthetically  annotated.  Localities 
and  location  of  all  available  specimens  are 
also  included. 


40 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Figure  25.  Nasal  scale  shape  (top)  and  plastron 
shape  (bottom)  in  holotype  of  Kinosternon 
hinipes  (ZSM  1374/0). 


Kinosternon  sonoriense     LeConte 
Sonora  Mud  Turtle 

Kinosternum  sonoriense  LeConte,  1854: 
184  [type-locality,  "Tucson,  in 
Sonora",  Arizona.  Type,  a  male,  col- 
lected by  Dr.  J.  L.  LeConte  (author's 
son)  and  placed  in  Philadelphia  Acad- 
emy of  Sciences;  presently  unlocat- 
able];  Troschel,  1855:415. 

Kinosternon  sonoriense  Gray,  1855:79 
(first  use  of  this  combination;  Tucson); 
Stejneger,  1902:149  (Fort  Huachuca 
and  Babacomari  creek,  Arizona);  Ruth- 
ven,  1907:594  (Sabino  Canyon,  Santa 
Catalina  Mountains,  Arizona); 
Mearns,  1907:117;  Van  Denburgh  and 
Slevin,  1913:396  (Gila  River  and  its 
tributaries;  8  specific  localities);  Grin- 
nell  and  Camp,  1917:200  (in  part;  result 


of  incorrect  synonymy  of  K.  flavescens 
with  A^.  sonoriense;  lower  Colorado 
River,  California);  Stejneger  and  Bar- 
bour, 1917:112  (in  part;  southern  New 
Mexico  and  Arizona  into  southeastern 
California;  northern  Mexico);  Schmidt, 
1922:618;  Van  Denburgh,  1922:967 
(Arizona,  18  localities;  California,  2 
localities;  and  Sonora,  5  localities); 
Pratt,  1923:238  (in  part;  western  Texas 
[=  K.  hirtipes]  into  southern  Califor- 
nia); Van  Denburgh,  1924:229  (New 
Mexico;  "Fort  Union"  locality  in  er- 
ror, see  Iverson,  1978);  Strecker  and 
Williams,  1927:15  (in  part;  Bexar  Co., 
Texa§  locality  based  on  K.  flavescens); 
Storer,  1930:430;  Ditmars,  1936:397  (in 
part;  southwestern  Texas  records  based 
on  K.  flavescens);  Dunn,  1936:472  (in 
part;  Chihuahua  locality  based  on  K. 
hirtipes);  Pickwell,  1947:60  (in  part; 
southwestern  Texas  record  based  on  K. 
flavescens);  Brown,  1950:228  (in  part; 
Texas  localities  based  on  K.  flavescens); 
H.  M.  Smith  and  Taylor,  1950a:26  (in 
part;  western  Texas  localities  based  on 
K.  flavescens;  Chihuahua  and  Durango 
localities  based  on  K.  hirtipes);  Carr, 
1952:90  (in  part;  Texas  records  based 
on  K.  flavescens);  Schmidt,  1953:91  (in 
part;  Texas  records  based  on  K. 
flavescens;  erroneously  restricted  type- 
locality  of  the  synonym  Kinosternum 
henrici  to  Las  Cruces,  New  Mexico); 
Mertens  and  Wermuth,  1955:336  (in 
part;  Texas  records  based  on  K.  flaves- 
cens; Chihuahua  and  Durango  records, 
on  K.  hirtipes);  Cagle,  in  Blair  et  al., 
1957:281  (in  part;  Texas  records  based 
on  K.  flavescens);  Gijzen  and 
Wermuth,  1958:44  (in  part;  photograph 
apparently  K.  integrum);  Wermuth  and 
Mertens,  1961:27  (in  part;  Texas  rec- 
ords based  on  K.  flavescens  ;  Chihua- 
hua and  Durango  records  based  on  K. 
hirtipes);  Casas  Andreu,  1965:382  (in 
part;  Chihuahua  and  Durango  records 
based  on  A',  hirtipes);  Stebbins,  1966:82 
(in  part;  Texas  records  based  on  K. 
flavescens;  Durango  records  based  on 
K.  hirtipes);  Casas  Andreu  1967:51  (in 


No.  1 


Kinosternon  Biosystematics 


41 


part;  Chihuahua  and  Durango  records 
based  on  K.  hirtipes);  Pritchard,  1967: 
37  (in  part;  Coahuila  records  incorrect; 
Texas  records  based  on  K.  flavescens); 
Cochran  and  Coin,  1970:136  (in  part; 
Texas  records  based  on  K.  flavescens); 
Legler  and  Webb,  1970:163  (in  part; 
Chihuahua  records  based  on  K. 
hirtipes);  Wiewandt,  1971:34  (in  part; 
southern  Sonora  records  based  on  K. 
alamosae);  Wiewandt,  Lowe  and 
Larson,  1972:162  (in  part;  southern 
Sonora  records  based  on  K.  alamosae); 
Ernst  and  Barbour,  1972:64  (in  part; 
Chihuahua  and  Durango  records  based 
on  A',  hirtipes);  Hambrick,  1976:291  (in 
part;  Texas  records  invalid);  Iverson, 
1976:1  (in  part;  upper  Ri'o  Yaqui  rec- 
ords in  Chihuahua  based  on  K. 
hirtipes);  Wermuth  and  Mertens,  1977: 
10;  Conant  and  Berry,  1978:1;  Iverson, 
1978:476;  H.  M.  Smith  and  R.  B. 
Smith,  1980:156;  Berry  and  Legler, 
1980:1. 

Thyrosternum  sonoriense  Agassiz,  1857: 
428;  Blair,  1859:3,  Troschel,  1860:270; 
Carman,  1885:8. 

Cinosternum  sonoriense  Agassiz,  1857: 
Plate  V,  fig.  8-11;  Cope,  1875:52, 
Coues,  1875:589;  Yarrow,  1883:31; 
Gunther,  1885:13;  Boulenger,  1889:40; 
Siebenrock,  1907:551;  Siebenrock, 
1909:444. 

Kinosternum  henrici  LeConte,  1859:4 
(type-locality,  "New  Mexico").  Type,  a 
male,  collected  by  Dr.  T.C.  Henry  and 
placed  in  Philadelphia  Academy  of 
Sciences  (ANSP  83).  Locality  data  with 
type  is  "Gila  River,  New  Mexico." 
Type-locality  erroneously  restricted  by 
Schmidt  (1953:91)  to  vicinity  of  Las 
Cruces;  Cope,  1880:13  (in  part;  Texas 
record  based  on  K.  flavescens). 

Thyrosternum  henrici  Troschel,  1860: 
270;  Carman,  1884:8. 

Cinosternon  henrici  Strauch,  1862:41; 
Strauch,  1865:101;  Strauch,  1890:89 
(in  part;  Dallas,  Texas  record  based  on 
K.  subrubrum). 

Cinosternon  sonoriense  Strauch,  1862:41; 
Strauch,  1865:100. 


Thylosternum  sonoriense  Muller,1865: 
598. 

Kinosternon  punctatum  Gray,  1870:67  (in 
part;  Sonora;  eastern  United  States  rec- 
ords based  on  K.  subrubrum) . 

Swanka  henricii  Gray,  1870:69. 

Platythyra  flavescens  Cooper,  1870:66 
(possibly  in  part;  see  Iverson,  1978; 
Colorado  River  Valley). 

Cinosternum  henrici  Cope,  1875:52; 
Yarrow,  1875:583;  Coues,  1875:590; 
Yarrow,  1883:31;  Boulenger,  1889:40; 
Ditmars  1907:26;  Strecker,  1915:10  (in 
part;  Texas  records  based  on  K.  flaves- 
cens); Malnate,  1971:353. 

Aromochelys  carinatus  Yarrow,  1875:582 
(in  part;  Arizona);  Coues,  1875:589  (in 
part;  Arizona). 

Cinosternum  flavescens  Yarrow,  1883:31 
(in  part;  "Utah"  and  "Fort  Mora", 
specimens  actually  K.  sonoriense,  see 
Iverson,  1978). 

Cinosternum  hirtipes  Gunther,  1885:15 
(in  part;  result  of  his  synonymy  of  K. 
henrici  LeConte  with  K.  hirtipes  Wag- 
ler);  Cope,  1887:23  (in  part;  result  of 
his  synonymy  of  C.  henrici  with  C.  hir- 
tipes); Gadow,  1905:209  (in  part;  Ari- 
zona and  New  Mexico). 

Cinosternon  integrum  Strauch,  1890:91 
(in  part;  result  of  his  synonymy  of  C. 
hirtipes  Gunther  with  C.  integrum 
LeConte). 

Kinosternon  flavescens  Van  Den  burgh, 
1922:972  (in  part;  Ft.  Verde  and 
Graham  Co.  records  actually  K.  sono- 
riense; see  Iverson,  1978);  LaRivers, 
1942:66  (in  part;  Nevada;  see  Iverson, 
1978);  Stebbins,  1966:82  (in  part; 
northwest  Arizona;  see  Iverson,  1978). 

Kinosternon  sp.  Little,  1940:264  (Roose- 
velt Reservoir  and  Sallymae  Creek,  Gila 
Co.,  Arizona);  Tanner  and  Robison, 
1960:59  (in  part;  specimens  are  K. 
sonoriense  but  locality  doubtful). 

Kinosternon  sonoriensis  Bogert  and 
Ohver,  1945:396;  Smith  and  Buechner, 
1947:10;  H.  M.  Smith,  Wiiliams  and 
Moll,  1963:207. 

Kinosternon  hirtipes  H.  M.  Smith  and  E. 
H.  Taylor,  1950a:25  (in  part;  Arizona). 


42 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Kinosternon    sonorensis    Weise,     1962: 

165. 
Kinosternon  seonoriense  Berry  and  Shine, 

1980:189. 

Type.  Lost;  see  synonymy. 

Content.  Two  subspecies,  one  new,  are 
described:  K.  s.  sonoriense  and  K.  s.  long- 
ifemorale. 

Diagnosis.  A  Kinosternon  of  the 
hirtipes  species  group  with:  1.)  the  adult 
nasal  scale  large  and  triangular,  rhomboi- 
dal  or  bell  shaped  (not  furcate  behind);  2.) 
usually  three  or  four  pairs  of  relatively 
long  chin  or  neck  barbels  (at  least  one  pair 
more  than  half  orbit  diameter);  3.)  male 
plastron  relatively  wide  (PWB  42-53%  of 
CL;  X  =  47.2%);  4.)  first  neural  often 
(38.1%)  in  contact  with  nuchal  bone;  5.) 
the  female  generally  larger  than  the  male; 
and,  6.)  populations  confined  to  Arizona, 
California,  New  Mexico,  Sonora,  western 
Chihuahua,  and  possibly  Baja  California. 

Description.  The  adult  carapace  gen- 
erally is  tricarinate  with  the  medial  keel 
most  apparent;  some  turtles  possess  well- 
defined  keels,  others  have  only  the  medi- 
an keel  present  with  mere  hints  of  the  two 
lateral  keels,  and  still  others  have  a  virtu- 
ally smooth  shell.  The  algae  covered  shells 
of  some  individuals  are  extremely  rugose 
and  densely  pock-marked  (Fig.  26;  found 
in  15  of  164  turtles  by  Hulse,  1976:347),  a 
condition  perhaps  induced  by  the  algae 
(the  condition  is  known  for  no  other  kino- 


Figure  26.  Articulated  shell  (without  scutes)  of  adult 
K.  sonoriense  (JBI  800)  from  Sonora,  near 
Cucurpe  (Rio  Sonora  basin).  Note  rugosity. 


sternid).  The  average  female  is  larger  than 
the  average  male.  Maximum  female  size  is 
175  mm  carapace  length;  males  155  mm. 
The  nasal  scale  is  not  furcate  behind  in 
adults.  The  first  vertebral  scute  usually 
touches  the  second  marginal.  The  axillary 
is  nearly  always  in  broad  contact  with  the 
inguinal,  and  the  inguinal  contacts  the 
eighth  marginal.  The  ninth  marginal  scute 
is  not  elevated  above  the  preceding  marg- 
inals. The  tenth  marginal  is  higher  than 
the  ninth  marginal  and  the  eleventh  mar- 
ginal may  or  may  not  be  elevated  to  the 
height  of  the  posterior  portion  of  the 
tenth  marginal.  Interpectoral  seam  length 
is  less  than  one-half  of  gular  length.  The 
nuchal  bone  often  contacts  the  first  neural 
bone.  The  carapace  is  brown  to  olive  in 
color,  the  seams  darker.  Well-developed 
transverse  plastral  hinges  border  the  ab- 
dominal scutes.  The  male  plastron  is  rel- 
atively extensive  (PWB  =  42  to  53%  CL). 
The  plastron  is  yellow  to  brownish  with 
darker  brown  seams.  The  bridge  area  is 
dark  brown.  The  skin  is  dark  gray  and  the 
head  and  neck  bear  cream  colored  mot- 
tlings  that  tend  to  form  at  least  one  pair  of 
stripes  extending  back  from  the  orbit,  one 
above  and  the  other  below  the  typanum 
after  intersecting  the  angle  of  the  jaw.  A 
yellow  or  cream  stripe  also  extends  from 
the  palmar  surface  of  each  foot  to  the 
base  of  the  hmb  along  its  posterior  sur- 
face in  some  adults.  Three  to  four  pairs  of 
relatively  long  chin  or  neck  barbels  usual- 
ly are  present.  Mature  females  possess 
short,  stubby  tails,  with  a  small  terminal 
spine,  whereas  males  possess  long,  thick- 
ened tails  with  a  large  terminal  spine  and  a 
patch  of  elevated  (tubercular),  acute, 
nonimbricated  scales  on  the  posterior  sur- 
face of  the  crus  and  thigh  of  each  hind 
leg. 

Remarks.  Iverson  (1976)  has  synthe- 
sized most  of  the  hterature.  Additional 
important  references  include  Hulse 
(1976);  Morafka  (1977);  Bowler  (1977); 
Conant  and  Berry  (1978);  Iverson  (1978); 
H.M.  Smith  and  R.B  Smith  (1980);  Berry 
and  Legler  (1980);  and  Iverson  and  Wey- 
man  (MS). 


No.  1 


Kinosternon  Biosystematics 


43 


Kinosternon  sonoriense  is  the  largest 
Kinosternon  in  which  the  females  are  gen- 
erally larger  than  the  males.  Perhaps  con- 
comitantly it  produces  the  largest  number 
of  eggs  per  clutch  of  any  kinosternine  — 
up  to  at  least  eight  (Hulse,  1974;  Iverson, 
unpubhshed  data).  I  have  observed  copu- 
lation in  the  field  near  Fort  Huachuca, 
Arizona  (Gila  River  basin)  on  4  May 
1974,  much  later  than  the  March-April 
records  of  Hulse  (1974).  The  smallest  tur- 
tles I  measured  were  22.3  mm  CL  (20.0 
mm  PL),  23.9  mm  CL  (18.3  mm  PL),  and 
25.7  mm  CL.  In  the  southern  part  of  its 
range,  this  turtle  is  apparently  active  year 
round;  I  have  observed  activity  at  Quito- 
baquito  Pond,  Arizona  on  several  occa- 
sions in  January. 
Range.  Kinosternon  sonoriense  occurs 

in  the  United  States  from  the  Lower  Colo- 
rado and  Gila  rivers  in  Arizona  and  New 
Mexico,  southward  to  and  including  the 
Rio  Yaqui  basin  west  of  the  continental 
divide,  and  eastward  through  the  Rfo 
Casas  Grandes  basin  of  northwestern  Chi- 
huahua. It  is  known  from  between  at  least 
43  and  2042  m  elevation.  The  species  may 
also  occur  in  the  Rio  Fuerte  (see  MATER- 
IALS AND  METHODS). 

Specimens  examined  and  Additional 
Records.  See  Locality  list. 

Etymology.  The  specific  name  sonor- 
iense refers  to  the  Sonoran  Biotic  Prov- 
ince, wherein  the  turtle  is  found. 

Kinosternon  sonoriense  sonoriense 
(LeConte) 

Sonora  Mud  Turtle 

Synonymy.  See  species  synonymy,  ex- 
cept those  references  in  synonymy  of  K. 
sonoriense  longifemorale. 

Holotype.  Lost;  see  species  account. 

Diagnosis.  A  subspecies  of  K.  sonori- 
iense  with  1)  a  relatively  long  interanal 
seam  (male  x  lAN/CL,  19.5'^o;  female  x  , 
23.0%);  2)  a  relatively  short  interfemoral 
seam  (male  X,  IF/CL,  10.1%;  female  x  , 
10.1%);  3)  a  first  vertebral  scute  of  medi- 
um width  (male  x  ,  VW/CL,  24.4%;  fe- 
male X  ,  25.5%);  and  4)  a  relatively  wide 


gular  scute  (male  x ,  GW/CL,  20.0%;  fe- 
male X  ,  19.4%). 

Range.  Kinosternon  s.  sonoriense  is 
definitely  known  from  the  Bill  Williams, 
lower  Colorado,  Gila,  Sonora,  Magda- 
lena,  Yaqui,  southwest  New  Mexico,  and 
Casas  Grandes  basins  of  Arizona  and 
New  Mexico,  and  Sonora  and  western 
Chihuahua,  Mexico. 

Specimens  examined  and  Additional 
Records.  See  locality  list. 

Etymology.  See  species  account. 

Kinosternon  sonoriense  longifemorale 

ssp.  nov. 

Sonoyta  Mud  Turtle 

Kinosternon  sonoriense  M earns,  1907: 1 17 
(Sonoyta);  Van  Denburgh,  1922:969. 
(Sonoyta  River  three  miles  from  Sonoy- 
ta); Stebbins,  1966:83  (Quitobaquito 
Spring);  Hulse,  1974:15,  94  (Quitoba- 
quito Spring);  H.M.  Smith  and  R.B. 
Smith,  1980:160  (3  localities  in  Sonoyta 
basin). 

Holotype.  USNM  21710,  young  male, 
preserved  whole,  from  "artificial  pond 
fed  by  springs",  Sonoyta,  Sonora,  Mex- 
ico (31°5rN,  112°50'W);  collected  15 
January  1894,  app^arently  by  E.A. 
Mearns. 

Paratypes.  USNM  21709  and  21711, 
topotypic  male  and  female,  preserved 
whole,  and  USNM  21708,  aduh  female, 
preserved  whole,  from  Sonoyta  River,  3 
mi.  from  Sonoyta,  collected  on  22  Janu- 
ary 1894  by  B.A.  Wood;  UAZ  27987  and 
27996,  adult  male  and  female,  respective- 
ly, preserved  whole,  Quitobaquito 
Springs,  Organ  Pipe  Cactus  National 
Monument,  Pima  County,  Arizona,  col- 
lected on  14  May  1967  and  10  April  1965, 
respectively,  by  R.D.  Krizman  and  T.J. 
Cox,  respectively;  and  UF  47719  and 
47720  (Fig.  27),  skeletal  aduh  male  and 
female,  respectively  from  Quitobaquito 
Pond,  Pima  County,  Arizona,  collected 
on  19  January  1976  by  John  B.  Iverson. 
Diagnosis.  A  subspecies  of  K.  sonori- 
ense with  1)  a  relatively  short  interanal 
seam  (male  x  ,  lAN/CL,  14.4%;  female 


44 


Tulune  Studies  in  Zoology  and  Boiany 


Vol.23 


Figure  27.  Plastron  of  female  Kinoslernon 
sonoriense  longifemorale  (UF  47720)  from 
Quitobaquito  Pond,  Pima  County,  Arizona. 
Note  short  interanal  and  long  interfemoral 
seams. 


X,  18.5%);  2)  a  relatively  long  interfem- 
oral seam  (male  x ,  IF/CL,  12.8%;  female 
X,  13.5%);  3)  a  wide  first  vertebral  scute 
(male  x,  VW/CL,  28.9%;  female  x, 
28.8%);  and  4)  a  relatively  narrow  gular 
scute  (male  x,  GW/CL,  17.7%;  female 
X,  17.8%). 

Range.  Kinoslernon  sonoriense  longi- 
femorale is  known  only  from  the  Ri'o 
Sonoyta  basin  in  Arizona  and  Sonora, 
Mexico. 

Specimens  examined  and  Additional 
Records.  See  locality  list. 

Etymology.  The  subspecific  name  long- 
ifemorale  is  from  the  Latin  longiis  (long) 
and  femoralis  (of  the  femur;  here  the 
femoral  scute)  and  refers  to  the  long  inter- 
femoral seam  which,  along  with  the  short 
interanal  seam,  diagnoses  the  taxon. 

Kinoslernon  hiriipes  (Wagler) 
Rough-footed  Mud  Turtle 

Cinosternon  hiriipes  Wagler,  1830:137, 
plate  5,  fig.  29-30  (Type-locality,  "Mex- 
ico", restricted  to  "lakes  near  Mexico 
City"  by  Schmidt  1953:89,  but  see  RE- 
MARKS under  A'.  /?.  hiriipes).  Holo- 
type,  Miinchen  Museum  (Germany) 
1374/0,  a  male,  collected  by  Baron  Kar- 


winski,  collecting  date  unknown.  Type- 
locality  incorrectly  restricted  to  "Maz- 
atlan,  Sinaloa"  by  H.M.  Smith  and 
E.H.  Taylor  1950b:25;  see  discussion  in 
Hardy  and  McDiarmid,  1969:70,  218); 
Wagler,  1833:plate  30;  Fitzinger, 
1835:125;  A.M.C.  Dumeril  and  Bibron, 
1834:370;  A.M.C.  Dumeril  and  Dum- 
eril, 1851:17;  Gray,  1855:46  (in  part; 
"Brazils"  record  in  error);  Strauch, 
1862:41;  Strauch,  1865:101;  A.H.A. 
Dumeril,  1870:25;  Westphal-Castelnau, 
1872:278;  Gray,  1873:113;  Bocourt, 
1876:8;  Duges,  1888:106. 

Kinoslernon  oblongum  Gray,  1844:33  (in 
part). 

Cinoslermon  hiriipes  Gray,  1844:33  (in 
synonymy). 

Kinosternum  hiriipes  LeConte,  1854:186; 
LeConte,  1859:5;  MuUer,  1885:716. 

Kinoslernon  hiriipes  Grdiy,  1855:47,  1869: 
183,  1870:67;  Stejneger,  1899:64;  Rust, 
1934:59;  Taylor,  1936:529  (in  part;  Sin- 
aloa records  based  on  K.  integrum); 
Martin  del  Campo,  1937:265  (in  part; 
Hidalgo  record  based  on  A',  integrum); 
Rust,  1938:22;  Caballero  y  Caballero, 
1938a:  103  (in  part;  Hidalgo  record 
based  on  A',  integrum);  Caballero  y 
Caballero,  1940a:225  (in  part;  Urua- 
pan,  Michoacan  locality  based  on  K.  in- 
tegrum );  H.M.  Smith  and  E.H. 
Taylor,  1950a:25  (in  part;  Chihuahua, 
Michoacan,  Guanajuato,  Mexico,  and 
Distrito  Federal;  other  locality  based  on 
K.  sonoriense,  K.  flavescens,  or  K.  inte- 
grum); H.M.  Smith  and  E.H.  Taylor, 
1950b:342  (in  part;  type-locality  re- 
striction to  Mazatlan,  Sinaloa  invalid); 
Glass  and  Hartweg,  1951:50;  Taylor, 
1952:793;  Schmidt,  1953:89;  Mertens 
and  Wermuth,  1955:336;  Cable  in  Blair 
et  al.  1957:281  (in  part;  Arizona  records 
based  on  A.  sonoriense);  Malkin, 
1958:75  (in  part;  Nayarit  records  based 
on  A.  integrum);  Zweifel,  1960:94  re- 
jects Tres  Marias  records;  Wermuth 
and  Mertens,  1961:19;  Croulet,  1963:4 
(in  part;  Nayarit  record  based  on  K.  in- 
tegrum); Liner,  1964:221  (in  part; 
Guanajuato  records  on  A',  integrum); 


No.  1 


Kinosternon  Biosystematics 


45 


Casas  Andreu,  1965:285  (in  part;  Sina- 
loa,  Colima,  and  Hidalgo  records  based 
on  K.   integrum);  Pritchard,   1967:37; 
Casas  Andreu,  1967:44  (in  part;  Sina- 
loa,  Colima,  Hidalgo  and  Nayarit  rec- 
ords based  on  K.  integrum);  Hardy  and 
McDiarmid,   1969:104  (rejects  Sinaloa 
records,    including    H.M.    Smith    and 
Taylor's    1950b:343    type-locality    re- 
striction); Cochran  and  Coin,  1970:135 
(in  part;  Arizona  records  based  on  K. 
sonoriense);  Ernst  and  Barbour,  1972: 
66  (in  part;  Arizona  records  based  on 
K.  sonoriense);  Dixon,  Ketchersid,  and 
Leib,  1972:228  (in  part;  Queretaro  rec- 
ord based  on  K.   integrum);  Greene, 
1972:  2  (in  part;  Puebla  record  based  on 
K.  integrum);  Bravo-HoUis  and  Cabal- 
lero    Deloya,    1973:109;    Conant    and 
Berry,  1978:1;  Iverson,  1978:1,  Iverson 
and     Berry,     1979:318;     Pritchard, 
1979:537;  H.M.  Smith  and  R.B.  Smith, 
1980:137;  Berry  and  Legler,  1980:1. 
Cinosternon  pensylvanicum  Lichtenstein, 
1856:2   (in   part;    Mexico);    Westphal- 
Castelnau,   1872:278  (in  part;  Guana- 
juato); Bocourt,  1876:5  (in  part;  Mex- 
ico);   Herrera,    1890:330,    1891:46   (in 
part;     Valley    of    Mexico);     Strauch, 
1890:88    (in    part;    Mexico);    Herrera, 
1893:339    (in    part;    Mexico);    Duges, 
1898:40  (in  part;  Mexico);  Herrera  and 
Lope,     1899:281     (in    part;    Mexico); 
Herrera,  1904:5  (in  part;  Mexico). 
Thryrosternum  hirt ipes  Agassiz,  1857:429. 
Ozotheca  hirtipes  LeConte,  1859:6;  Tros- 

chel,  1860:270. 
Ozotheca  odorata  Duges,  1869:143  (in 
part;  states  of  Guanajuato  and  Mexico); 
Velasco,  1890b:54  (in  part, 
Guanajuato); 
Velasco,  1891:52  (in  part;  Queretaro 
record  unsubstantiated);  Velasco, 
1892b:40  (in  part;  Tlaxcala  record  in- 
correct); Velasco,  1893b:81  (in  part; 
Sonora  record  incorrect);  Velasco, 
1894:40  (in  part;  Zacatecas  record  un- 
substantiated); Velasco,  1896a:30  (in 
part;  Aguascalientes  record  unsubstan- 
tiated); Velasco,  1898:62  (in  part; 
Chiapas  record  incorrect). 


Ozhoteca  odorata  Garcia  Cubas,  1884: 
179  (in  part;  Mexico);  Velasco,  1890a: 
35  (in  part;  Nuevo  Leon  record  incor- 
rect); Velasco,  1893a:64  (in  part;  Dur- 
ango  record  unsubstantiated);  Velasco 
1897:41  (in  part;  Coahuila  record  incor- 
rect). 

Cinosternum  hirtipes  Gunther,  1885:13 
(in  part;  Sinaloa  records  based  on  K.  in- 
tegrum; Arizona  and  New  Mexico  rec- 
ords based  on  K.  sonoriense);  Cope, 
1885:390;  Cope,  1887:23  (in  part;  Col- 
ima and  Sinaloa  records  based  on  K.  in- 
tegrum); Garman,  1887:16  (in  part;  San 
Luis  Potosf  record  based  on  K.  inte- 
grum); Boulenger,  1889:38;  Gadow, 
1905:209  (in  part;  Arizona  and  New 
Mexico  records  based  on  K.  sonori- 
ense); Siebenrock,  1906:94,  1907: 
551;  Gadow,  1908:5;  Siebenrock,  1909: 
444;  Gadow,  1930:50. 

Cinosternon  pennsylvanicum  Duges, 
1888:10  (in  part;  Valley  of  Mexico  and 
Guanajuato  state);  Velasco,  1890b:291 
(in  part;  Guanajuato);  Duges,  1895:5 
(in  part;  Guanajuato);  Duges,  1896a:  Iv 
(in  part;  Mexico);  Duges,  1896b:329  (in 
part;  Mexico);  Duges,  1896c:479  (in 
part,  Guanajuato). 

Ozothea  odorata  Velasco,  1892a:76  (in 
part;  Guerrero  record  incorrect);  Velas- 
co, 1892c:79  (in  part;  Tamaulipas  rec- 
ord incorrect);  Velasco,  1895:38  (in 
part;  Campeche  record  incorrect);  Vel- 
asco, 1896b:37  (in  part;  Colima  record 
incorrect). 

Cynosternon  pennsylvanicum  Herrera, 
1893:342  (in  part;  Valley  of  Mexico). 

Cinosternum  pennsylvanicum  Gadow, 
1905:209  (in  part;  Valley  of  Mexico). 

Kinosternum  pennsilvanicum  Cope,  1896: 
1021  (in  part;  Austrocentral  district  of 
Mexico). 

Cinosternonus  pensylvanicum  Herrera, 
1899:28  (in  part;  Mexico). 

Cynosternon  pensylvanicum  Herrera  and 
Lope,  1899:131  (in  part;  Valley  of  Mex- 
ico). 

Cinosternum  pennsilvanicum  Cope,  19(X): 
1229  (in  part;  Valleys  of  Mexico  and 
Toluca    northward    through    Guana- 


46 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


juato). 
Cinosternum    pensylvanicum    Lampe, 

1901:184-85  (North  Mexico). 
Cinosternon  sp.  Herrera,  1904:6  (Valley 

of  Mexico). 
Cinosternum  integrum  Gadow,  1908:518 
(in  part;  Laguna  de  Zapotlan,  Jalisco). 
Kinosternon  sonoriense  Dunn,   1936:472 
(in  part;  Chihuahua);  H.M.  Smith  and 
E.H.  Taylor,   1950a:26  (in  part;  Chi- 
huahua  and   Durango);   Mertens   and 
Wermuth,   1955:338  (in  part;  Chihua- 
hua to  Durango);  Casas  Andreu,  1965: 
386,  1967:52  (in  part;  Chihuahua  and 
Durango);  Legler  and  Webb,  1970:163 
(in  part;  western  Chihuahua);  Iverson, 
1976:1  (in  part;  upper  Rfo  Yaqui,  Chi- 
huahua; see  Iverson,  1978). 
Chinosternum  hirtipes  Caballero  y  Cabal- 
lero,  1938b:449  (in  part;  Hidalgo  local- 
ity based  on  K.  integrum). 
Sternotherus  odoratus  Brown,   1950:230 
(in    part;    Presidio    Co.,    Texas;    see 
Conant  and  Berry,  1978). 
Kinosternon  murrayi  Glass  and  Hartweg, 
1951:50  (type-locality,  "Harper  Ranch, 
37    miles    south    of    Marfa,    Presidio 
County,  Texas."  Holotype,  Texas  Co- 
operative  Wildlife   Collection   650,    a 
young  male,  collected  by  S.H.  Wheeler 
on  12  August  1941);  Peters,   1952:54; 
Legler,  1960:139  (Lajitas,  Texas  record 
in  error);  Cochran,  1961:232. 
Kinosternon  flavescens  Stebbins,  1966:82 
(in  part;  Durango;  see  Iverson,  1978); 
Morafka,    1977:70,   Map  25  (in  part; 
some  northern  Mexico  records  based  on 
K.  hirtipes.). 
Kynosternon  hirtipes  Lopez  1975:2  (Val- 
ley of  Mexico). 
Kinosternon    hertipes    Semmler    et    al., 
1977:  18  (Chihuahua). 
Types.  Only  the  holotype  (Fig.  25),  an 
adult  male,  preserved  whole,  is  available, 
contrary  to  Bocourt's  (1876:8)  suggestion 
that  Wagler's  (1830,  1833)  figures  (Plate 
5:fig.  29-30  and  Plate  30:figs.  1-3,  respec- 
tively) of  Cinosternon  (-   Kinosternon) 
hirtipes  were  based  on  two  different  speci- 
mens. 


Content.  Six  subspecies,  four  new,  are 
described:  K.  h.  hirtipes,  K.  h.  chapal- 
aense,  K.  h.  murrayi,  K.  h.  magdalense, 
K.  h.  tarascense,  and  K.  h.  megacephal- 
um. 

Diagnosis.  A  Kinosternon  of  the  hir- 
tipes species  group  with  1)  the  adult  nasal 
scale  reduced  and  crescent-shaped,  or 
larger  and  furcate  behind,  or  still  larger 
and  triangular  or  bell  shaped  (the  latter 
combination  characteristic  only  of  Valley 
of  Mexico  turtles);  2)  usually  three  pairs 
of  relatively  short  chin  barbels  (  <  half 
orbit  diameter);  3)  male  plastron  relative- 
ly narrow  (PWB  36  to  50%  of  CL;  k  = 
43'Vo);  4)  first  neural  rarely  (10.2%)  con- 
tacting nuchal  bone;  5)  the  male  generally 
larger  than  the  female;  and  6)  populations 
confined  to  Central  Mexico  from  Chihua- 
hua (and  adjacent  Texas)  to  Jalisco, 
Michoac^n,  and  Mexico  (state).  Despite 
its  anomalous  absence  on  the  holotype 
(Fig.  25),  a  nuchal  scute  is  typically  pres- 
ent. 

Description.  As  for  K.  sonoriense  ex- 
cept as  stated  above,  and  1)  the  carapacial 
keels  are  almost  never  absent  (i.e.,  the 
median  keel  is  virtually  always  evident  at 
least  posteriorly);  2)  maximum  female  size 
is  157  mm  carapace  length,  male  182  mm; 
3)  carapace  light  to  dark  brown  to  nearly 
black  in  color;  4)  plastron  usually  yellow 
to  brown  with  darker  brown  seams  but 
sometimes  (stained  ?)  nearly  black;  5) 
head  markings  extremely  variable  (coarse- 
ly mottled,  reticulated  or  spotted  to 
almost  unmarked;  see  subspecific  ac- 
counts). 

Remarks.  Most  of  the  literature  is 
synthesized  in  Iverson  (in  press).  A  discus- 
sion of  the  evolutionary  significance  of 
the  geographically  variant  sexual  size  di- 
morphism of  this  species  appears  in  Iver- 
son (MS  2).  Reproductive  parameters  are 
summarized  here  (Table  2)  and  in  Iverson 
(MS  2).  Clutch  size  data  (4-5  and  4-7  eggs) 
in  Moll  and  Legler  (1971)  are  all  referable 
to  the  subspecies  murrayi.  Scaling  of  skel- 
etal components  is  discussed  in  Iverson 
(MS  1)  and  Iverson  and  Weyman  (MS). 


No.  1 


Kinosternon  Biosystematics 


47 


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48 


Tulune  Sludles  in  Zoology  und  Botany 


Vol.  23 


Range.  Primarily  distributed  on  the 
Mexican  Plateau,  Kinosternon  hirdpes 
ranges  from  Alamito  Creek  in  Texas  in 
the  United  States  and  the  Rfos  Santa 
Maria,  Carmen,  and  Conchos  in  northern 
Mexico  south  and  eastward  to  the  Chap- 
ala,  Zapotlan,  San  Juanico,  Pa'tzcuaro, 
and  Valle  de  Me'xico  basins  of  the  Sierra 
Volcanica  Transversal  of  southern  Mex- 
ico. It  is  known  from  between  at  least  800 
and  2600  m  in  elevation. 

Specimens  examined  and  Additional 
Records.  See  locality  list. 

Etymology.  The  specific  name  hirtlpes 
is  from  the  Latin,  hirtus,  meaning  rough, 
and  pes  meaning  foot,  and  refers  to  the 
rough  scales  on  the  feet  of  the  species. 

Kinosternon  hirtlpes  hirtlpes  Wagler 
Valley  of  Mexico  Mud  Turtle 

CInosternon    hirtlpes   Wagler,    1830:187 

(see  species  synonymy). 
Ozotheca   odorata   Duges,    1869:143    (in 

part;  State  of  Mexico). 
CInosternon    pennsylvanlcum    Duges, 

1888:107  (in  part;  Valley  of  Mexico). 
CInosternum  /?//7//7e5  Boulenger,  1889:38; 

Siebenrock,  1906:94,  1907:551  (State  of 

Mexico);  Gadow,  1908:5  (Chalco  lakes, 

Valley  of  Mexico). 
CInosternon    pensvlvanlcum     Herrera, 

1890:330,   1891:46  (in  part;  Valley  of 

Mexico). 
Cynostenum    pennsylvanlcum    Herrera, 

1893:342  (in  part;  Valley  of  Mexico). 
Cynosternon    pensylvanlciim     Herrera 

and  Lope,  1899:131  (in  part;  Valley  of 

Mexico). 
CInosternom  pennsllvanlcum  Cope,  1900: 

1229  (in  part;  Valley  of  Mexico). 
CInosternon  sp.  Herrera,   1904:6  (Mexi- 

calzingo.  Valley  of  Mexico). 
CInosternum    pennsylvanlcum     Gadow, 

1905:209  (in  part;  Valley  of  Mexico). 
Kinosternon  hirtlpes  Martin  del  Campo, 

1938:391  (Valley  of  Mexico);  Caballero 

y  C,   1939:279  (Xochimilco,  Mexico, 

Distrito    Federal);    H.M.    Smith    and 

Taylor,     1950a:25    (Distrito    Federal); 

Glass  and  Hartweg,  1951:50  (Valley  of 


Mexico);  Schmidt,  1953:89;  Beltz, 
1954:124  (Mexico  City,  Mexico); 
Martin  del  Campo,  1955:66  (Valley  of 
Mexico);  Deevey,  1957:240  (Valley  of 
Mexico);  Casas  Andreu,  1965:385  (Dis- 
trito Federal);  Kranz,  Smith,  and 
Smith,  1971:23  (near  City  of  Mexico); 
Greene,  1972:2  (in  part;  Mexico  City, 
Puebla  locality  based  on  A'.  Integrum); 
Perez  Villegas  and  Reyna  Trujillo, 
1978:215  (southern  region  of  Valley  of 
Mexico). 
Kinosternon  hirtlpes  hirtlpes  Mertens  and 
Wermuth,  1955:336  (first  use  of  this 
combination;  in  part;  State  of  Mexico); 
Wermuth  and  Mertens,  1961:20  (in 
part;  State  of  Mexico);  Duellman, 
1961:57,  1965:653  (in  part;  Michoacan 
localities  not  this  subspecies);  H.M. 
Smith,  Williams  and  Moll,  1963:209; 
Liner,  1964:221  (in  part;  Guanajuato 
records  not  this  subspecies);  Pritchard, 
1967:37  (in  part;  State  of  Mexico); 
Casas  Andreu,  1967:44  (in  part;  State 
of  Mexico);  Parsons,  1968:1238;  Legler 
and  Webb,  1970:163  (in  part;  Chihua- 
hua records  based  on  A",  h.  murrayl); 
Mittermeier,  1971:16  (Mexico  City); 
Moll  and  Legler,  1971:92  (in  part;  Chi- 
huahua records  based  on  K.  h. 
murrayl);  Wermuth  and  Mertens,  1977: 
7;  Pritchard,  1979:537  (in  part;  Mexico 
City). 
Kynosternon  hirtlpes  Lopez  1975:2  (Val- 
ley of  Mexico. 
Kinosternon  sp.  Niederberger,  1979:134 
(Valley  of  Mexico  archeological  re- 
mains: 5500  BC). 

Types.  Only  the  holotype  (Fig.  25)  is 
available  (see  Remarks  below). 

Diagnosis.  A  subspecies  oi  Kinosternon 
hirtlpes  with  1)  the  adult  nasal  scale  tri- 
angular, rhomboidal,  or  bell-shaped  (fur- 
cate behind  in  subadults,  but  not  in  large 
adults);  2)  a  mottled  head  pattern,  typical- 
ly organized  into  a  light  streak  extending 
posteriorly  from  the  angle  of  the  jaw, 
with  a  similar  light  postorbital  streak  vari- 
ably evident;  3)  one  or  (typically)  two 
pairs  of  mental  chin  barbels,  the  anterior 
pair  largest;  4)  medium  body  size  (maxi- 


No.  1 


Kinosternon  Biosystematics 


49 


mum  male  size  140  mm  CL;  female,  140 
mm);  5)  relatively  short  bridge  length 
(male  BL/CL,    17.6<7o;    female 

21.7'^o);  6)  relatively  short  interfemoral 
seam    length    (male  IF/CL,    6.9%; 

female  ,  7.1%);  7)  relatively  long  inter- 
anal  seam  length  (male  lAN/CL, 
20.6%;  female  ,25.8%)  and  8)  popula- 
tions confined  to  the  Valley  of  Mexico. 

Remarks.  As  mentioned  under  SYN- 
THESIS (above),  the  allocation  of  the 
holotype  of  Kinosternon  hirtipes  to  the 
Valley  of  Mexico  must  remain  uncertain 
until  additional  material  is  available  from 
the  southern  and  southwestern  margins  of 
the  Mexican  Plateau. 

Specific  natural  history  data  are  un- 
available for  Kinosternon  hirtipes  hir- 
tipes. A  photograph  of  the  plastron  of 
UMMZ  99449,  an  adult  female,  appears 
in  H.M.  Smith  and  R.  B.  Smith  (1980; 
plate  19,  bottom). 

Range.  Kinosternon  hirtipes  hirtipes  is 
known  only  from  the  drainages  of  the 
Valley  of  Mexico. 

Specimens  Examined  and  Additional 
Records.  See  locality  lists. 
Etymology.  See  species  account. 
Moll  and  Legler,  1971:92  (in  part;  Chi- 
huahua   records    based    on    k.     h. 
murrayi);  Wermuth  and  Mertens,  1977: 
7;  Pritchard,  1979:537  (in  part;  Mexico 
City). 
Kynosternon  hirtipes  Lopez  1975:2  (Val 

ley  of  Mexico. 
Kinosternon  sp.  Niederberger,   1979:134 
(Valley    of   Mexico    archeological    re- 
mains: 5500  BC). 

Types.  Only  the  holotypes  (Fig.  27)  is 
available  (see  Remarks  below). 

Diagnosis.  A  subspecies  of  Kinosternon 
hirtipes  with  1)  the  adult  nasal  scale  tri- 
angular, rhomboidal,  or  bell-shaped  (fur- 
cate behind  in  subadults,  but  not  in  large 
adults);  2)  a  mottled  head  pattern,  typical- 
ly organized  into  a  light  streak  extending 
posteriorly  from  the  angle  of  the  jaw, 
with  a  similar  light  postorbital  streak  vari- 
ably evident;  3)  one  or  (typically)  two 
pairs  of  mental  chin  barbels,  the  anterior 
pair  largest;  4)  medium  body  size  (maxi- 


mum male  size  140  mm  CL;  female,  140 
mm);  5)  relatively  short  bridge  length 
(male  x  BL/CL,  17.6%;  female  x  , 
21.7%);  7)  relatively  short  interfemoral 
seam  length  (male  x  IF/CL,  6.9%; 
female  x  ,  7.1%);  8)  relatively  long  inter- 
anal  seam  length  (male  x  lAN/CL, 
20.6%;  female  x,  25.8%)  and  9)  popula- 
tions confined  to  the  Valley  of  Mexico. 

Remarks.  As  mentioned  under  SYN- 
THESIS (above),  the  allocation  of  the 
holotype  of  Kinosternon  hirtipes  to  the 
Valley  of  Mexico  must  remain  uncertain 
until  additional  material  is  available  from 
the  southern  and  southwestern  margins  of 
the  Mexican  Plateau. 

Specific  natural  history  data  are  un- 
available for  Kinosternon  hirtipes  hir- 
tipes. A  photograph  of  the  plastron  of 
UMMZ  99449,  an  adult  female,  appears 
in  H.M.  Smith  and  R.  B.  Smith  (1980; 
plate  19,  bottom). 

Range.  Kinosternon  hirtipes  hirtipes  is 
known  only  from  the  drainages  of  the 
Valley  of  Mexico. 

Specimens  examined  and  Additional 
Records.  See  locality  lists. 

Etymology.  See  species  account. 

Kinosternon  hirtipes  murrayi 
Glass  and  Hartweg 

Murray's  Mud  Turtle 

Ozotheca  odorata  Duges,  1869:143  (in 
part;  Guanajuato);  Velasco  1890b: 54  (in 
part;  Guanajuato). 

Cinosternon  hirtipes  Westphal-Castelnau, 
1872:278  (Guanajuato). 

Cinosternum  hirtipes  Cope,  1887:23  (in 
part;  city  of  Chihuahua,  Guanajuato). 

Cinosternum  pennsylvanicum  Duges, 
1896c:479  (Guanajuato). 

Cinosternon  pennsilvanicum  Cope,  1900: 
1229  (Toluca  Valley  northward  through 
Guanajuato. 

Kinosternon  sonoriense  Dunn,  1936:472 
(in  part;  Rio  Conchos,  Julimes,  Chi- 
huahua); H.M.  Smith  and  Taylor 
1950a:26  (in  part;  Chihuahua  and  Dur- 
ango);  Mertens  and  Wermuth,  1955: 
338  (in  part;  Chihuahua  and  Durango); 


50 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Casas  Andreu,  1965:386  (in  part;  Chi- 
huahua   and    Durango);    Legler    and 
Webb,   1970:163  (in  part;  Rfos  Papi- 
gochic  and  Tomuchic  in  western  Chi- 
huahua);   Iverson,    1976:1    (in    part; 
Upper  Rfo  Yaqui,  Chihuahua;  see  Iver- 
son, 1978). 
Kinosternon    hirtipes    Caballero    y    C, 
1940b: 562   (Rio   Lerdo,   Guanajuato); 
Caballero  y  C.  y  Cerecero,   1943:534 
(Rio    Lerdo    del    Valle    de    Santiago, 
Guanajuato);  H.M.  Smith  and  Taylor, 
1950b:25  (in  part;  Chihuahua,  Guana- 
juato); Williams,  Smith,  and  Chrapliwy, 
1960:36  (Chihuahua,  1  mi.  E  La  Cruz); 
Casas  Andreu,  1965:385  (in  part;  Chi- 
huahua, Guanajuato);  Conant,   1978: 
465  (Texas,  Chihuahua,  Durango  and 
Zacatecas). 
Sternothenis  odoratus  Brown,   1950:230 
(in  part;  Presidio  Co.,  Texas;  based  on 
holotype  oiK.  murrayi;  see  Conant  and 
Berry,  1978:15). 
Kinosternon  murrayi  Glass  and  Hartweg, 
1951:50     (Type-locality,     "Harper 
Ranch,  37  miles  south  of  Marfa,  Pres- 
idio    County,     Texas."     Holotype, 
TCWC  650,  a  young  male,  collected  12 
August  1941  by  S.H.  Wheeler.);  Peters, 
1952:54  (Texas);  Legler,  1960:139  (Jet. 
Rio  San  Pedro  and  Conchos,  and  Ojin- 
aga.  Chihuahua). 
Kinosternon    hirtipes  murrayi   Schmidt, 
1953:89    (first    use    of    combination; 
Texas);  Mertens  and  Wermuth,  1955: 
336  (Texas);  H.M.  Smith,  Williams  and 
Moll,    1963:207    (Chihuahua);    Casas 
Andreu,    1967:45   (Texas,  Chihuahua, 
and    Durango);    Parsons,    1968:1238; 
Cochran  and  Goin,  1970:135  (Texas); 
Moll  and  Legler,  1971:92  (Durango  and 
Chihuahua);  Ernst  and  Barbour,  1972: 
66    (Texas);     Hambrick,     1976:292 
(Texas);  Wermuth  and  Mertens,  1977:7 
(Texas);    Conant    and    Berry,    1978:1 
(Texas  and  Chihuahua);  Iverson,  1978: 
476  (Chihuahua). 
Kinosternon  hirtipes  hirtipes  Mertens  and 
Wermuth,  1955:336  (in  part;  Chihua- 
hua);  Duellman,    1961:57   (in  part  ?; 
Michoacan,  8  km  W  Ciudad  Hidalgo 


and  Lago'de  Cuitzeo);  Casas  Andreu, 
1967:44  (in  part;  Chihuahua,  Michoa- 
can, and  Guanajuato);  Legler  and 
Webb,  1970:163  (in  part;  Rios  Papi- 
gochic  and  Tomuchic,  western  Chihua- 
hua); Moll  and  Legler,  1971:92  (in  part; 
Chihuahua);  Wermuth  and  Mertens, 
1977:7  (middle  and  western  Mexico); 
Pritchard  1979:537  (in  part). 
Kinosternon  flavescens  Stebbins,  1966:82 
(in  part;  Durango;  see  Iverson,  1978). 
Kinosternon  hirtipes  murryi  Ashton  et  al., 

1976:51  (lapsus  pro  murrayi). 
Kinosternon  hertipes  Semmler,  et  al., 
1977:18  (near  Galeana,  Chihuahua). 
Types.  Holotype:  see  subspecies  synon- 
ymy. Paratypes:  USNM  15860,  adult 
male,  preserved  whole,  from  "Marfa, 
Presidio  County,  Texas",  collected  by 
Vernon  Bailey:  UMMZ  101294,  adult 
male,  preserved  whole,  and  UMMZ  S- 
1083,  shell  of  adult  male,  both  topotypic 
and  collected  12  June  1950  by  Herndon 
G.  Dowling. 

Diagnosis.  A  subspecies  oi Kinosternon 
hirtipes  with:  1)  a  large  posteriorly  furcate 
nasal  scale  (typically  exending  posterior  to 
the  orbits);  2)  an  extremely  variable  mot- 
tled to  reticulated  head  pattern;  3)  typical- 
ly two  pair  of  mental  chin  barbels,  the 
anterior  pair  largest;  4)  medium  to  large 
body  size  (maximum  known  male  size, 
182  mm  CL;  female,  157  mm);^)  relative- 
ly long  bridge  length  (male  x  BL/CL, 
20.0<^o;  female  x,  23. 7%);  6)  relatively 
long  gular  length  (male  x  GL/CL, 
14.7<^o;  female  x,  15.8%);  and  7)  popu- 
lations confined  to  the  Big  Bend  region  of 
Texas  and  adjacent  Chihuahua  southward 
across  the  Mexican  Plateau  to  northern 
Jalisco,  northern  Michoacan,  and  eastern 
Mexico  (state). 

Remarks.  As  discussed  in  the  results, 
there  appears  to  be  a  slight  morphometric 
distinction  between  populations  of  K.  h. 
murrayi  in  the  Ri'o  Nazas  northward,  and 
populations  in  the  Rio  Aguanaval  south- 
ward. This  difference  is  not  considered 
significant  enough  to  warrant  subspecific 
designation,  but  has  some  interesting  zoo- 
geographic  impUcations. 


No.  I 


Kinosternon  Biosystematics 


51 


Range.  Kinosternon  hirtipes  murrayi  is 
known  from  the  following  basins  in 
Aguascalientes  ,  Chihuahua,  Coahuila, 
Durango,  Guanajuato,  Jalisco,  Mexico, 
Michoacan,  San  Luis  Potosi,  Texas,  and 
Zacatecas:  Santa  Maria  (Chihuahua), 
Carmen,  El  Sauz,  Conchos,  Bustillos, 
Papigochic,  Nazas,  Viesca,  Aguanaval, 
Santiaguillo,  Mezquital,  El  Salto,  Santa 
Maria  (San  Luis  Potosi;  presumably  in- 
troduced), Aguascalientes,  Verde,  Lerma 
(except  Chapala),  Cuitzeo,  Balsas,  and 
Villa  Victoria  (with  reservation). 

Specimens  examined  and  Additional 
Records.  See  locality  lists. 

Etymology.  The  subspecific  name  mur- 
rayi is  a  patronym,  honoring  Dr.  Leo  T. 
Murray  of  Texas  A  &  M  College. 

Kinosternon  hirtipes  chapalaense  ssp.  nov. 

Lake  Chapala  Mud  turtle 

Cinosternum  integrum  Gadow,  1908:518 

(in  part;  Laguna  de  Zapotlan,  Jalisco). 
Kinosternon  hirtipes  Altini,  1942:153  (in 

part;  Lake  Chapala,  Jalisco). 
Kinosternon  hirtipes  hirtipes  Duellman, 

1961:57,  1965:653  (in  part;  Jiquilpan, 

La  Palma,  Lago  de  Came'cuaro  =   14 

km  E  Zamora,  Michoacan). 
Kinosternon  hirtipes  chapalaense  Pritch- 

ard,    1979:557   (nomen  nudum;  Lake 

Chapala). 

Holotype.  UMMZ  97128,  adult  male, 
preserved  whole,  from  Lake  Chapala, 
0.25  mile  off  Chapala,  Jalisco,  Mexico 
[20°18'N,  103°12'W];  collected  15  July 
1947,  by  Norman  Hartweg. 

Paratypes.  All  preserved  whole:  UMMZ 
97122-23,  topotypic  adult  females;  UMMZ 
97124,  topotypic  subadult  male;  97125-27 
and  97129-30,  topotypic  adult  males;  and 
UU  12126-12128,  adult  male,  subadult  fe- 
male, and  juvenile,  Lago  de  Chapala,  3.2 
km  W  Chapala;  and  UU  12125,  adult  fe- 
male, Lago  de  Chapala,  6.1  km  W  Ajijic, 
all  collected  on  21-22  June  1969  by  Clyde 
Barbour. 

Diagnosis.  A  subspecies  of  Kinosternon 
hirtipes  with  1)  a  reduced  crescent-shaped 
nasal  shield,  which  nearly  always  lies  an- 


terior to  the  orbits  (Figure  23);  2)  a  reduc- 
tion of  dark  pigment  on  the  head  and 
neck,  dark  markings  confined  to  isolated 
spots  or  reticulations  dorsally  (Figures  23 
and  24),  but  laterally  sometimes  organ- 
ized as  two  dark,  nearly  parallel  post- 
orbital  stripes;  3)  the  neck  and  chin  virtu- 
ally unmarked  and  the  mandibular  and 
maxillary  sheaths  bearing  only  a  few  dark 
streaks,  if  any;  4)  one,  two,  or  three  pairs 
of  mental  barbels  present,  the  anterior- 
most  pair  (near  the  mandibular  symphysis) 
usually  the  largest;  5)  medium  body  size 
(maximum  known  size  for  males  is  152 
mm  CL;  females,  149  mm);  6)  relatively 
long  bridge  length  (male  x  BL/CL, 
20.3<^o;  female  x,  25.3%);  7)  relatively 
long  interanal  _seam  (male  x  lAN/CL, 
19.1%;  female  x,  25.2%);  and  8)  popula- 
tions confined  to  the  Chapala  and  Zapot- 
lan (and  possibly  Duero)  basins  of  Jalisco 
and  Michoacan. 

Remarks.  Field  notes  accompanying 
the  topotypes  provide  no  additional  geo- 
graphical and  ecological  information. 
However,  Clyde  Barbour  (pers.  comm.) 
obtained  the  non-topotypic  paratypes 
(during  the  night  21-22  June  1969)  along 
the  shore  of  Lake  Chapala,  on  trot-lines 
baited  with  liver.  These  lines  were  neces- 
sarily buoyed  off  the  lake  bottom  with 
floats  to  avoid  bait  removal  by  crabs. 
Peter  Meylan  found  a  single  rotten  car- 
cass of  this  species  on  the  south  shore  of 
Lake  Chapala  just  east  of  Tuxcueca  dur- 
ing my  field  trip  to  the  area  on  15  June 
1978;  trapping  at  that  locality  produced 
no  turtles.  Trapping  in  isolated  spring-fed 
pools  just  northeast  of  the  town  of  Chap- 
ala on  9  May  1981  produced  only  K.  inte- 
grum. 

Range.  Kinosternon  hirtipes  chapalense 
is  known  only  from  the  Lago  de  Chapala 
and  Laguna  de  Zapotlan  drainage  basins 
in  Jalisco  and  Michoacan,  Mexico.  Speci- 
mens from  the  Rio  Duero  basin  are  tenta- 
tively considered  intergrades  with  K.  h. 
murrayi. 

Specimens  Examined  and  Additional 
Records.  See  locality  list. 

Etymology.  The  subspecific  name  chap- 


52 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


alaense  refers  to  Lake  Chapala  wherein 
the  type  series  was  collected. 

tKinosternon  hirtipes  megacephalum 
ssp.  nov. 

Viesca  Mud  Turtle 

Holotype.  SM(BCB)  1 1466,  adult  male, 
preserved  whole,  from  3.2  km  SE  Viesca 
[25  °2rN,  102  °48'W],  Coahuila;  collected 
4  June  1961  by  Bryce  C.  Brown  and  John 
Wottring  by  seining  a  drying  pond. 

Paratypes.  SM(BCB)  11460-65,  adult 
females,  preserved  whole,  all  topotypic 
(11461  photographed  in  H.M.  Smith  and 
R.B.  Smith,  1980);  and  SM(BCB)  9823, 
adult  male,  preserved  whole,  from  9.7  km 
SW  Viesca,  also  collected  on  4  June  1961 
by  Brown  and  Wottring  (see  Figs.  22  and 
24). 

Diagnosis.  A  subspecies  of  Kinosternon 
hirtipes  with:  1)  enlarged  head,  hyper- 
trophied  head  musculature,  and  broad  al- 
veolar jaw  surfaces  (Fig.  24);  2)  the  nasal 
scale  furcate  posteriorly;  3)  the  head  pat- 
tern mottled  or  reticulated  as  in  K.  h. 
murrayi;  4)  three  to  four  pairs  of  chin 
barbels  are  present,  two  to  three  mental 
pairs  (anterior  usually  the  largest)  and  one 
small  pair  at  level  of  anterior  edge  of 
tympanum;  5)  small  body  size  (maximum 
known  size  for  males  99  mm  CL;  females, 
1 17  mm);  6)  plastron  extremely  reduced  in 
size  (Fig.  22);  7)  relatively  short  bridge 
length  (male  x  BL/CL,  17.3^70;  female  x , 
23.90/0);  8)  relatively  short  gular  length 
(male  x  GL/CL,  ll.Oo/o;  female  x, 
12.8°7o);  9)  relatively  short  interanal  seam 
length  (male  x  lAN/CL,  15.9"7o;  female 
X  ,  20.90/0);  and  10)  populations  confined 
to  southwestern  Coahuila. 

Remarks.  This  subspecies  is  known 
only  from  the  type  series.  Field  work  in 
the  area  of  the  type  locality  (see  MATER- 
IALS AND  METHODS)  suggests  that  K. 
h.  megacephalum  is  now  extinct;  natural 
permanent  water  habitats  apparently  no 
longer  exist  near  Viesca.  Future  field 
work  should  be  concentrated  in  the 
mountains  south  of  the  city  of  Viesca  in 


hope  of  discovering  permanent  water  situ- 
ations where  turtles  (and  fishes?)  might 
still  exist. 

The  distinctive  trophic  apparatus  of 
this  subspecies  is  likely  an  adaptation  to 
stenophagous  molluscivory. 

Range.  Known  only  from  the  two  local- 
ities in  Coahuila  at  which  the  type  series 
was  collected. 

Etymology.  The  subspecific  name 
megacephalum  is  from  the  Greek  mega, 
meaning  large,  and  kephale,  meaning 
head,  and  refers  to  the  enlarged  head,  di- 
agnostic of  the  subspecies. 

Kinosternon  hirtipes  tarascense  ssp.  nov. 
Patzcuaro  Mud  Turtle 

Kinosternon  hirtipes  Altini,  1942:153  (in 

part;  Lake  Patzcuaro,  Michoacan). 
Kinosternon  hirtipes  hirtipes  Duellman, 
1961:57,    1965:653   (in  part;   Lago  de 
Patzcuaro,  Michoacan);  Casas  Andreu, 
1967:45  (in  part;  Patzcuaro,  Canal  de  la 
Tzipecua,  Michoacan). 
Holotype.  UF  43506,  adult  male,  pre- 
served whole,  from  Lago  de  Patzcuaro, 
adjacent  to  city  of  Patzcuaro  [19°32'N, 
101°36'W];     purchased    in    Patzcuaro 
market  13  June  1978  by  John  B.  Iverson. 
Paratypes.  All  topotypic  and  preserved 
whole:     UF    43505    and    43596,    adult 
females;  and  UF  43507  and  43595,  adult 
males. 

Diagnosis.  A  subspecies  of  K.  hirtipes 
with:  1)  a  typically  finely  mottled  to 
spotted  head  (Fig.  24);  2)  variable  red- 
brown  to  brown  staining  on  the  otherwise 
light  yellow  plastral  scutes  [The  dark  plas- 
tral  scutes  are  apparently  a  result  of 
natural  staining;  the  character  is  exhibited 
to  variable  degrees  by  individuals  and  the 
dark  color  is  lost  when  plastral  scutes  are 
shed  (Fig.  22)];  3)  the  large  nasal  scale 
posteriorly  furcate;  4)  two  pairs  of  mental 
chin  barbels  typically  present;  5)  small  to 
medium  body  size  (maximum  known  size 
for  males  136  mm  CL;  females,  132  mm)^: 
6)  relatively  short  bridge  length  (male  x 
BL/CL,  18.0^^0;  female  x,  21. 40/0);  7)  rel- 


No.  1 


Kinosternon  Biosystematics 


53 


atively  short  gular  length  (male  x 
GL/CL,  10.6%;  female  x  ,  12.6%);  8)  rel- 
atively long  interpectoral  seam  length 
(malex  IP/CL,  10. lo/o;  female  x,  8.5%); 
and,  9)  populations  confined  to  the  Lago 
de  Patzcuaro  drainage  basin. 

Remarks.  Despite  considerable  study  of 
other  components  of  the  biota  of  the 
Lago  de  Patzcuaro  (see  review  in  Cole, 
1963  and  Barbour,  1973),  the  mud  turtles 
have  been  ignored.  Reproductive  infor- 
mation resulting  from  my  studies  appears 
in  Table  2. 

Range.  Known  only  from  the  basin  of 
the  Lago  de  Patzcuaro,  Michoacan. 

Specimens  examined  and  Additional 
Records.  See  locality  list. 

Etymology.  The  subspecies  name  tara- 
scense  honors  the  native  tribe  of  Indians, 
the  Tarascas,  inhabiting  the  Patzcuaro 
area. 

Kinosternon  hirtipes  magdalense  ssp.  nov. 
San  Juanico  Mud  Turtle 

Holotype.  UF  45035,  an  adult  male, 
preserved  whole,  from  along  the  face  of 
the  dam  at  Presa  San  Juanico,  Michoacan 
[ca.  19°50'N,  102°40'W]  (Fig.  28).  Holo- 
type collected  15  June  1978  by  John  B. 
Iverson,  Peter  A.  Meylan,  and  Ron 
Magill. 

Paratypes.  UF  45036,  a  subadult  fe- 
male, UF  45038,  female  shell;  UF  45039- 
40,  male  shells,  all  topotypic;  and  TUL 
18677,  aduU  male,  collected  atop  Presa 
San  Juanico  9  August  1963  by  Clyde  D. 
Barbour  and  Salvador  Contreras-Balderas. 

Diagnosis.  A  subspecies  o{  Kinosternon 
hirtipes  with:  1)  a  finely  mottled  to  spot- 
ted head  pattern  with  jaw  streaking  mini- 
mal or  absent;  2)  a  large  nasal  scale,  fur- 
cate behind;  3)  two  pairs  of  mental  chin 
barbels  present;  4)  small  body  size  (max- 
imum known  male  size  94  mm  CL;  fe- 
male, 91  mm);  5)  a  relatively  small  plas- 
tron (male  x  PWB/CL,  41.9%;  female 
X,  43.5%);  6)  relatively  short  bridge 
length  (male  x  BL/CL,  18.5%;  female  x , 
19.7%);  7)  relatively  short  gular  length 


(male  x  GL/CL,  9.9%;  female  x, 
11.0%);  8)  relatively  long  interpectoral 
seam  length  (male  x  IP/CL,  8.7%;  fe- 
male X,  11.0%);  and,  9)  populations  re- 
stricted to  the  Magdalena  Valley,  Micho- 
acan. 

Remarks.  As  mentioned  earlier  (MAT- 
ERIALS AND  METHODS),  the  turtles 
inhabiting  the  Presa  San  Juanico  are 
poorly  known.  Future  field  work  in  the 
area  should  help  delimit  the  subspecific 
range  within  the  Magdalena  Valley  and 
also  provide  basic  natural  history  infor- 
mation. 

Range.  Kinosternon  hirtipes  magdal- 
ense is  known  only  from  the  type  series, 
all  from  the  reservoir  above  Presa  San 
Juanico  in  the  Magdalena  Valley  of 
Michoacan,  Mexico. 

Etymology.  The  subspecific  name  mag- 
dalense refers  to  the  Magdalena  Valley  of 
Michoacan  to  which  the  subspecies  is 
apparently  endemic. 


Figure  28.   Holotype  (UF  45035)  of  Kinosternon 
hirtipes  magdalense. 


54 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Evolution 

Based  on  the  derived  characters  of  the 
turtles  of  the  Kinosternon  hirtipes  species 
group  (Table  3),  I  have  constructed  a  phy- 
logeny  of  the  included  taxa  (Fig. 29).  For 
reasons  discussed  by  Farris  (1966)  and 
Kluge  and  Farris  (1969)  (e.g.,  high  intra- 
familial  variation  and  indiscrete  character 
shifts),  I  have  not  always  assumed  that 
taxa  sharing  derived  morphometric  char- 
acters are  closely  related.  In  fact,  the  dis- 
tribution of  some  character  states  among 
taxa  clearly  indicates  that  those  characters 
are  not  a  result  of  single  origin,  but  rather 
of  convergence.  For  example,  the  length- 
ening of  the  interanal  seam  in  K.  sonor- 
iense  (longifemorale)  and  K.  hirtipes  (hir- 
tipes and  chapalaense)  certainly  illustrates 


multiple  origin  of  a  derived  character 
state.  In  addition,  Viesca  (megacephal- 
um),  Patzcuaro  (tarascense),  and  San 
Juanico  turtles  (magdalense)  all  share  a 
relatively  short  bridge  (with  Valley  of 
Mexico  turtles),  a  short  gular,  and  small 
body  size  (the  latter  two  also  share  a  long 
interpectoral  seam),  yet  geographically 
and  zoogeographically  (Iverson,  in  prep- 
aration) the  three  populations  likely  do 
not  represent  a  monophyletic  divergence 
from  a  pre-murrayi  stock.  Rather,  the 
evolution  of  these  character  states  is  more 
likely  a  response  to  selection  in  similar, 
very  narrow  adaptive  zones  (i.e.,  isolated, 
very  small  basins).  Unfortunately,  the 
functional  significance  of  those  characters 
is  unknown,  as  is  that  of  most  of  the  other 
characters  herein  examined  (but  see  Iver- 
son, MS  2) 


Table  3.  Tally  of  subspecific  taxa  exhibiting  derived  character  states  in  the  Kinosternon 
hirtipes  species  group.  Primitive  states  are  discussed  in  the  text. 


Derived  Character 


1 

small  plastron 

2 

short  bridge 

3 

short  gular 

4 

long  gular 

5 

long  interpectoral 

6 

short  interpectoral 

7 

short  interfemoral 

8 

long  interanal 

9 

short  interanal 

10 

V-shaped  nasal 

11 

reduced  nasal 

12 

multiple,  long  chin 

barbels 

13 

reduced  head 

pigment 

14 

well-developed 

head  stripes 

15 

large  head 

16 

female  >  male 

17 

small  body  size 

18 

tendency  toward 

unicarination 

Taxa  exhibiting  derived  character 


magdalense,  megacephalum 

magdalense,  tarascense,  hirtipes,  megacephalum 

magdalense,  tarascense,  megacephalum 

sonoriense  and  longifemorale 

magdalense,  tarascense 

sonoriense,  longifemorale 

hirtipes 

chapalaense,  hirtipes,  longifemorale 

megacephalum 

murrayi,  magdalense,  tarascense,  megacephalum, 

chapalaense 
chapalaense 
sonoriense,  longifemorale 

chapalaense 

hirtipes 

megacephalum 

sonoriense,  longifemorale  (possibly  tarascense, 

megacephalum) 
magdalense,  tarascense,  megacephalum 
murrayi,  hirtipes,  magdalense,  tarascense, 

megacephalum,  chapalaense 


No.  I 


Kinosternon  Biosystematics 


55 


The  Kinosternon  hirtipes  species  group 
apparently  evolved  on  the  Mexican  Pla- 
teau from  an  ancestor  as  yet  unknown. 
Despite  the  fact  that  several  coastal 
streams  have  come  to  drain  the  Plateau 
due  to  headwater  stream  erosion  (e.g., 
Rios  Yaqui,  Me^quital,  Santiago,  Balsas; 
see  Fig.  4),  K.  hirtipes  has  nowhere  left 
the  Plateau.  This  is  surprising  since  K. 
integrum  has  apparently  moved  both  up 
and  down  several  of  these  basins  (Balsas, 
Santiago-Lerma,  and  Mezquital;  Iverson, 
unpublished).  K.  sonoriense  apparently 
evolved  from  a  K.  hirtipes-\\ke  ancestor 
isolated  in  the  Sonoran  Desert,  possibly 
following  migration  across  the  well- 
documented  Sonora  Desert-Chihuahua 
Desert  filter  barrier  in  southeastern  Ari- 
zona, southwestern  New  Mexico  and  ad- 
jacent Mexico  (see  review  in  Morafka, 
1977).  Because  so  much  geological  infor- 
mation concerning  the  Mexican  Plateau  is 
now  available  (see  reviews  in  Barbour, 
1973  and  Wauer  and  Riskind,  1978),  a 
discussion  of  the  historical  zoogeography 
of  the  K.  hirtipes  species  group  will 
appear  elsewhere  (Iverson,  in  prepara- 
tion). 

The  relationship  between  the  Kino- 
sternon hirtipes  species  group  and  other 
Kinosternon    is    unclear.     Siebenrock 


p*  „.''^/^  ..^' y   ./-  j^"-   y 


Figure  29.  A  theory  of  relationships  among  the 
subspecific  taxa  of  the  Kinosternon  hirtipes 
species  group.  Numbers  refer  to  derived  character 
states  listed  in  Table  3.  Solid  lines  cutting  line- 
ages mark  identical  shifts  (convergence)  in 
character  states. 


(1907:551)  included  K.  hirtipes  and  K. 
sonoriense,  K.  baurii,  K.  subrubrum,  K. 
flavescens,  and  K.  steindachneri  (=  K. 
subrubrum)  in  the  K.  subrubrum  species 
group.  However,  I  believe  that  K.  baurii 
and  K.  subrubrum  (including  steindach- 
neri) represent  a  species  group  distinct 
from  the  K.  hirtipes  group,  and  that  K. 
flavescens  is  similarly  distinct.  Perhaps 
the  closest  relative  of  the  hirtipes  group  is 
K.  herrerai  (found  in  the  Tampico  Em- 
bayment  of  eastern  Mexico;  i.e.,  non- 
Plateau),  which  shares  with  most  K.  hir- 
tipes the  elevated  scale  patches  on  the 
hindlegs  of  males,  the  tendency  toward 
unicarination  in  adults,  the  furcate  nasal 
scale,  the  reduced  plastron,  the  broad 
inguinal-axillary  contact,  and  several 
morphometric  plastral  characters.  Un- 
fortunately, the  determination  of  the 
phylogenetic  relationship  of  the  K.  hir- 
tipes group  to  the  other  species  groups  in 
the  genus  must  await  further  analysis. 

Key  To  Adult  Turtles  Of  The 
Kinosternon  hirtipes  SPECIES  GROUP 

lA.  Nasal  shield  triangular,  rhomboidal, 
or  bell  shaped;  largest  2  pairs  of  chin 
barbels  relatively  long  (at  least  one 
pair  >  half  orbit  diameter,  with  one 
pairmental  and  other  at  mid-tympan- 
um level);  interpectoral  length 
averages  5.0%  of  plastron  length  in 
males  (less  than  8%  in  95%  of  cases) 
and  4.0%  in  females  (less  than  7%  in 
96%  of  cases);  posterior  width  of 
plastral  forelobe  (PWB)  averages 
47.2%  of  carapace  length  in  males 
(more  than  44%  in  95%  of  cases)  and 
49.0%  in  females  (more  than  45%  in 
96%  of  cases);  maximum  gular  width 
averages  19.7%  of  carapace  length  in 
males  (more  than  18%  in  94%  of 
cases)  and  19.1%  in  females  (more 
than  17%  in  94%  of  cases);  first 
neural  bone  often  (38.1%)  in  contact 
with  nuchal  bone;  northwestern 
Chihuahua  and  Sonora,  Mexico  and 
adjacent  New  Mexico,  Arizona  and 
California Kino- 
sternon sonoriense 2 


56 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


IB.  Nasal  shield  large  and  deeply  notched 
posteriorly  (V-shaped),  or  reduced  to 
crescent-shaped    scale    lying    fully 
anterior    to    level    of    orbits,     or 
triangular,     rhomboidal,     or     bell 
shaped   if   from   Valley  of  Mexico; 
largest     2     pairs    of    chin     barbels 
relatively    short    (<     half    orbit 
diameter),    mentally    located,    with 
anterior    pair    larger;     interpectoral 
length    averages    8.2*^0    of   plastron 
length  in  males  (more  than  4.5%  in 
97<^o  of  cases);  and  6.6%  in  females 
(more  than  3.5%  in  94%  of  cases); 
posterior  width  of  plastral  forelobe 
(PWB)  averages  42.8%  of  carapace 
length  in  males  (less  than  48%  in  98% 
of  cases)  and  47.6%  in  females  (less 
than  51%  in  95%  of  cases);  maximum 
gular    width    averages     17.3%     of 
carapace  length  in  males  (less  than 
20%  in  98%  of  cases)  and  17.0%  in 
females  (less  than  20%  in  98%  of 
cases);  first  neural  rarely  (10.2%)  in 
contact    with    nuchal;     Chihuahua, 
Mexico   and  adjacent  Texas   south- 
ward   to    Jalisco,    Michoacan,    and 

Me'xico,  Mexico  (state) 

Kinosternon  hirtipes 3 

2A.  Interanal  seam  length  averages 
19.5%  of  carapace  length  in  males 
(more  than  16.5%  in  97%  of  cases) 
and  23.0%  in  females  (more  than 
21%  in  90%  of  cases);  interfemoral 
seam  length  averages  10.1%  of 
carapace  length  in  males  (less  than 
13%  in  93%  of  cases)  and  10.1%  in 
females  (less  than  12.5%  in  95%  of 
cases);  maximum  first  vertebral  width 
averages  24.4%  of  carapace  length  in 
males  Oess  than  28%  in  97%  of  cases) 
and  25.5%  in  females  (less  than  28% 
in  90%  of  cases);  and  gular  width 
averages  20.0%  in  males  (more  than 
18.5%  in  93%  of  cases)  and  19.4%  in 
females  (more  than  17.5%  in  90%  of 
cases);  Bill  Williams,  lower  Colorado, 
Gila,  Sonora,  Magdalena,  Yaqui, 
southwest  New  Mexico,  and  Casas 
Grandes  basins K.s.  sonoriense 


2B.  Interanal  seam  length  averages 
14.4%  of  carapace  length  in  males 
(less  than  16%  in  90%  of  cases),  and 
18.5%  in  females  (less  than  22%  in 
100%  of  cases);  interfemoral  seam 
length  averages  12.8%  of  carapace 
length  in  males  (more  than  10%  in 
100%  of  cases)  and  13.5%  in  females 
(more  than  11.5%  in  91%  of  cases); 
maximum  first  vertebral  width 
averages  28.9%  of  carapace  length  in 
males  (more  than  28%  in  90%  of 
cases)  and  28.8%  in  females  (more 
than  26%  in  100%  of  cases);  and 
gular  width  averages  17.7%  of 
carapace  length  in  males  (less  than 
19%  in  100%  of  cases)  and  17.8%  in 
females  (less  than  20%  in  100%  of 
cases);  Rio  Sonoyta  basin,  Arizona, 

and  Sonora,  Mexico 

K.  s.  longifemorale 

3A.  Nasal  shield  reduced  to  crescent- 
shaped  scale  lying  anterior  to  level  of 
orbits;  dark  reticulate  head  markings 
reduced  or  nearly  absent;  plastral 
width  at  humero-pectoral  seam 
(PWA)  averages  33.3%  of  carapace 
length  in  males  (less  than  35.5%  in 
100%  of  cases)  and  37.0%  in  females 
(less  than  40%  in  93%  of  cases); 
bridge  length  averages  20.3%  of 
carapace  length  in  males  (over  18%  in 
100%  of  cases)  and  25.3%  in  females 
(more  than  22%  in  100%  of  cases); 
gular  length  averages  11.8%  of 
carapace  length  in  males  (less  than 
13%  in  93%  of  cases)  and  14.5%  in 
females  (less  than  18.5%  in  100%  of 
cases);  forelobe  length  averages 
30.5%  of  carapace  length  in  males 
(less  than  33.5%  in  100%  of  cases) 
and  31.8%  in  females  (less  than  34% 
in  100%  of  cases);  interhumeral  seam 
length  averages  14.0%  of  maximum 
plastron  length  in  males  (more  than 
12%  in  93%  of  cases)  and  12.7%  in 
females  (more  than  10%  in  88%  of 
cases);  interabdominal  seam  length 
averages  28.6%  of  maximum  plastron 
length  in  males  (more  than  26%  in 


No.  I 


Kinosternon  Biosystematics 


57 


93*^0  of  cases)  and  ZQ.S'Vo  in  females 
(more  than  25. 5 "^o  in  100<Vo  of  cases); 
Lake  Chapala  and  Lake  Zapotlan 
basins,    Michoacan    and    Jalisco, 

Mexico 

....  Kinosternon  hirtipes  chapalaense 

3B.  Nasal  shield  large  and  deeply  notched 
posteriorly,  triangular,  rhomboidal  or 
bell  shaped;  head  with  abundant  dark 
head  markings;  plastral  width  at 
humero-pectoral  seam  (PWA)  av- 
erages 36.1%  of  carapace  length  in 
males  (more  than  32.5%  in  98%  of 
cases)  and  38.1%  in  females  (more 
than  35%  in  97%  of  cases);  bridge 
length  averages  19.9%  of  carapace 
length  in  males  (less  than  22%  in  95% 
of  cases)  and  23.4%  in  females  (less 
than  26%  in  93%  of  cases);  gular 
length  averages  14.6%  of  carapace 
length  in  males  (more  than  12%  in 
95%  of  cases;  excluding  turtles  from 
Patzcuaro,  San  Juanico,  Cuitzeo,  and 
Viesca  basins)  and  15.6%  in  females 
(more  than  12.5%  in  97%  of  cases; 
excluding  turtles  from  Patzcuaro,  San 
Juanico,  Cuitzeo,  and  Viesca  basins); 
forelobe  length  average  31.2%  of 
carapace  length  in  males  (more  than 
28.5%  in  100%  of  cases)  and  34.4% 
in  females  (more  than  30%  in  98%  of 
cases);  interhumeral  seam  length  aver- 
ages 11.8%  of  maximum  plastron 
length  in  males  (less  than  15%  in  90% 
of  cases)  and  12.6%  in  females  (less 
than  17%  in  96%  of  cases);  interab- 
dominal  seam  length  averages  28.1% 
of  maximum  plastron  length  in  males 
(less  than  31%  in  96%  of  cases)  and 
28.0%  in  females  (less  than  31%  in 
95%  of  cases);  Chihuahua,  Mexico 
and  adjacent  Texas  southward  to  Ja- 
lisco, Michoacan,  and  Mexico,  Me'x- 
ico,  except  Chapala  and  Zapotlan 
basins 4 

4A.  Gular  length  averages  10.5%  of  cara- 
pace length  in  males  (less  than  13%  in 
100%  of  cases)  and  12.4%  in  females 
(less  than  14%  in  93.0%  of  cases); 
plastron    width    at    humero-pectoral 


seam  (PWA)  averages  34.6%  of  cara- 
pace length  in  males  (less  than  38%  in 
100%  of  cases)  and  35.9%  in  females 
(less  than  38%  in  93%  of  cases);  pos- 
terior width  of  plastral  forelobe 
(PWB)  averages  42.5%  of  carapace 
length  in  males  (less  than  45.6%  in 
100%  of  cases)  and  45.8%  in  females 
(less  than  47.5%  in  88%  of  cases);  an- 
terior width  of  plastral  hindlobe 
(PWC)  averages  39.3%  of  carapace 
length  in  males  (less  than  43%  in 
100%  of  cases)  and  42.8%  in  females 
(less  than  46%  in  93%  of  cases);  max- 
imum carapace  length  140  mm  in 
males,  135  mm  in  females;  Patzcuaro, 

San  Juanico,  and  Viesca  basins 

populations  with  small  plastron ....  5 

4B.  Gular  length  averages  14.8%  of  cara- 
pace length  in  males  (more  than  12% 
in  94%  of  cases)  and  15.9%  in  fe- 
males (more  than  13%  in  94%  of 
cases);  plastron  width  at  humero-pec- 
toral seam  (PWA)  averages  36.2%  of 
carapace  length  in  males  (more  than 
33%  in  97%  of  cases)  and  38.3%  in 
females  (more  than  35.5%  in  96%  of 
cases);  posterior  width  of  plastral 
forelobe  (PWB)  averages  42.9%  of 
carapace  length  in  males  (more  than 
39%  in  94%  of  cases)  and  47.6%  in 
females  (more  than  45%  in  94%  of 
cases);  anterior  width  of  plastral  hind- 
lobe  (PWC)  averages  38.0%  of  cara- 
pace length  in  males  (more  than  34% 
in  96%  of  cases)  and  43.2%  in  fe- 
males (more  than  40%  in  97%  of 
cases);  maximum  carapace  length  185 
mm  in  males,  160  mm  in  females; 
Chihuahua  and  Texas  to  Jalisco, 
Michoacan,  and  Mexico,  except  Cha- 
pala, Zapotlan,  Patzcuaro,  San  Juan- 
ico, and  Viesca  basins popula- 
tions with  large  plastron 7 

5A.  Head  enlarged,  jaws  with  extremely 
broad  alveolar  surfaces;  carapace 
width  averages  61.9%  of  carapace 
length  in  males  (less  than  65%  in 
100%  of  cases)  and  68.1%  in  females 
(less  than  71.5%  in  100%  of  cases); 


58 


Tulanc  Studies  in  Zoology  and  Botany 


Vol.  23 


plastral  torelobe  length  averages 
28.7%  of  carapace  length  in  males 
(less  than  30%  in  lOOo/o  of  cases)  and 
29.0%  in  females  (less  than  30.5%  in 
100%  of  cases);  plastral  width  at  fe- 
moro-anal  seam  (PWD)  averages 
28.2%  in  males  (less  than  29%  in 
100%  of  cases)  and  31.4%  in  females 
(less  than  32.5%  in  100%  of  cases); 
interpectoral  seam  length  averages 
6.8%  of  carapace  length  in  males  (less 
than  8%  in  100%  of  cases)  and  4.8% 
in  females  (less  than  6.5%  in  100%  of 
cases);  bridge  length  averages  17.3% 
of  carapace  length  in  males  (less  than 
17.5%  in  100%  of  cases)  and  23.9% 
in  females  (more  than  23%  in  100% 

of  cases);  Viesca  area,  Coahuila 

K.  h.  tnegacephalum 

5B.  Head  not  enlarged,  jaws  with  narrow 
alveolar  surfaces;  carapace  width  av- 
erages 72.0%  of  carapace  length  in 
males  (more  than  65%  in  100%  of 
cases)  and  72.7%  in  females  (more 
than  66.5%  in  100%  of  cases);  plas- 
tral forelobe  length  averages  31.2% 
of  carapace  length  in  males  (more 
than  29.5%  in  100%  of  cases)  and 
33.3%  in  females  (more  than  30.5% 
in  100%  of  cases);  plastral  width  at 
femoro-anal  seam  (PWD)  averages 
29.1%  in  males  (more  than  28%  in 
100%  of  cases)  and  34.0%  in  females 
(more  than  31%  in  100%  of  cases); 
interpectoral  seam  length  averages 
9.6%  of  carapace  length  in  males 
(more  than  8%  in  100%  of  cases)  and 
8.9%  in  females  (more  than  6%  in 
100%  of  cases);  bridge  length  aver- 
ages 18.2%  of  carapace  length  in 
males  (more  than  16%  in  100%  of 
cases)  and  20.9%  in  females  (less  than 
23.5%  in  100%  of  cases);  Pa'tzcuaro 
and/or  San  Juanico  basins,  Michoa- 
can 6 

6A.  Plastral  scutes  usually  immaculate, 
not  darkly  stained;  maximum  plastral 
hindlobe  length  averages  30.1%  of 
carapace  length  in  males  (less  than 
32%  in  100%  of  cases)  and  31.5%  in 


females  (less  than  33%  in  100%  of 
cases);  plastral  width  at  humero-pec- 
toral  seam  (PWA)  averages  33.7%  of 
carapace  length  in  males  (less  than 
38%  in  100%  of  cases)  and  34.8%  in 
females  (less  than  36%  in  100%  of 
cases);  posterior  width  of  plastral 
forelobe  averages  41.9%  of  carapace 
length  in  males  (less  than  43%  in 
100%  of  cases)  and  43.5%  in  females 
(less  than  45.5%  in  100%  of  cases); 
interpectoral  seam  length  averages 
10.3%  of  maximum  plastron  length  in 
males  (less  than  12%  in  100%  of 
cases)  and  12.3%  in  females  (more 
than  11.5%  in  100%  of  cases);  first 
vertebral  scute  width  averages  22.3% 
of  carapace  length  in  males  (less  than 
23.5%  in  100%  of  cases)  and  21.4% 
in  females  (less  than  22.5%  in  100% 
of    cases);     San     Juanico     basin, 

Michoacan K.  h.  magdalense 

6B.  Plastral  scutes  often  stained  red- 
brown  to  dark  brown;  maximum  plas- 
tral hindlobe  length  averages  31.3% 
of  carapace  length  in  males  (more 
than  29%  in  100%  of  cases)  and  34% 
in  females  (more  than  32%  in  100% 
of  cases);  plastral  width  at  humero- 
pectoral  seam  (PWA)  averages  35.4% 
of  carapace  length  in  males  (more 
than  33.5%  in  100%  of  cases)  and 
36.8%  in  females  (more  than  35%  in 
100%  of  cases);  posterior  width  of 
plastral  forelobe  averages  43.6%  of 
carapace  length  in  males  (more  than 
41%  in  100%  of  cases)  and  46.8%  in 
females  (more  than  43.5%  in  100%  of 
cases);  interpectoral  seam  length  aver- 
ages 11.7%  of  maximum  plastron 
length  in  males  (more  than  9%  in 
100%  of  cases)  and  9.1%  in  females 
(less  than  12%  in  100%  of  cases);  first 
vertebral  scute  width  averages  22.9% 
of  carapace  length  in  males  (more 
than  20.5%  in  100%  of  cases)  and 
24.5%  in  females  (more  than  22%  in 
100%    of    cases);    Lake    Patzcuaro 

basin,  Michoacan 

K.  h.  tarascense 


No.  1 


Kinosternon  Biosystematics 


59 


7A.  Nasal  scale  triangular,  rhomboidal, 
or  bell  shaped;  maximum  plastron 
length  averages  86.4%  of  carapace 
length  in  males  (less  than  90.5%  in 
100%  of  cases)  and  91.6%  in  females 
(less  than  94%  in  100%  of  cases); 
bridge  length  averages  17.6%  of  cara- 
pace length  in  males  (less  than  19.5% 
in  100%  of  cases)  and  21.7%  in  fe- 
males (less  than  23%  in  94%  of 
cases);  interabdominal  seam  length 
averages  22.7%  of  carapace  length  in 
males  (less  than  23.5%  in  100%  of 
cases)  and  24.3%  in  females  (less  than 
26%  in  94%  of  cases);  interfemoral 
seam  length  averages  6.9%  of  cara- 
pace length  in  males  (less  than  8.5% 
in  1(X)%  of  cases)  and  7.1%  in  fe- 
males (less  than  8.5%  in  100%  of 
cases);  inter  anal  seam  length  averages 
20.6%  of  carapace  length  in  males 
(more  than  19%  in  100%  of  cases) 
and  25.8%  in  females  (more  than 
23.5%  in  100%  of  cases);  Valley  of 
Mexico K.  h.  hirtipes 

7B.  Nasal  scale  deeply  notched  posteri- 
orly (V-shaped);  maximum  plastron 
length  averages  86.0%  of  carapace 
length  in  males  (more  than  81%  in 
95%  of  cases)  and  92.6%  in  females 
(more  than  88%  in  96%  of  cases); 
bridge  length  averages  20.0%  of  cara- 
pace length  in  males  (more  than  18% 
in  97%  of  cases)  and  23.7%  in  fe- 
males (more  than  21%  in  95%  of 
cases);  interabdominal  seam  length 
averages  24.2%  of  carapace  length  in 
males  (more  than  21.5%  in  98%  of 
cases)  and  26.0%  in  females  (more 
than  23%  in  96%  of  cases);  interfe- 
moral seam  length  averages  9.0%  of 
carapace  length  in  males  (more  than 
6%  in  98%  of  cases)  and  9.0%  in  fe- 
males (more  than  6%  in  99%  of 
cases);  interanal  seam  length  averages 
18.6%  of  carapace  length  in  males 
(less  than  22%  in  95%  of  cases)  and 
23.4%  in  females  (less  than  26%  in 
93%  of  cases);  Chihuahua  and  Texas 
south  to  Michoacan,  Jalisco,  and 
Mexico K.  h.  murrayi 


Aknowledgments 

I  am  deeply  indebted  to  many  persons 
for  the  loan  or  gift  of  specimens  or  infor- 
mation, including  T.  Alvarez,  Walter 
Auffenberg,  R.  H.  Baker,  CD.  Barbour, 
J.  F.  Berry,  R.  L.  Bezy,  J.  Black,  Bryce 
Brown,  C.  C.  Carpenter,  A.  F.  Carr,  G. 
Casas  Andreu,  A.  H.  Chaney,  J.  Christ- 
iansen, J.  T.  Collins,  R.  Conant,  R. 
Crombie,  J,  Cross,  B.  J.  Davis,  W.  G. 
Degenhardt,  J.  R.  Dixon,  N.  H.  Douglas, 
H.  A.  Dundee,  W.  E.  Duellman,  M.  J. 
Fouquette,  T.  Fritts,  D.  Frost,  A.  L.  Gen- 
naro,  J.  W.  Gibbons,  U.  Gruber,  D. 
Hahn,  L.  M.  Hardy,  M.  M.  Hensley,  H. 
Hidalgo,  D.  Hoffmeister,  J.  F.  Jackson, 
E.  D.  Keiser,  A.  G.  Kluge,  J.  M.  Legler, 
A.  E.  Leviton,  E.  A.  Liner,  D.  W.  Linzey, 

C.  H.  Lowe,  J.  D.  Lynch,  E.  V.  Malnate, 
R.  F.  Malnate,  R.  F.  Martin,  H.  Marx,  T. 
P.  MasHn,  C.  J.  McCoy,  R.  R.  Miller,  E. 
O.  Moll,  O.  Mooser,  D.  J.  Morafka,  R. 
W.  Murphy,  C.  W.  Myers,  M.  A.  Nick- 
erson,  R.  Nussbaum,  A.  H.  Price,  G.  G. 
Raun,  R.  Reynolds,  M.  D.  Robinson,  D. 
A.  Rossman,  J.  F.  Scudday,  M.  E.  Seidel, 

D.  Smith,  H.  M.  Smith,  P.  W.  Smith,  R. 
C.  Stebbins,  W.  Tanner,  E.  H.  Taylor,  D. 
W.  Tinkle,  F.  Truxal,  T.  M.  Uzzell,  R. 
W.  Van  Devender,  T.  R.  Van  Devender, 
R.  Vogt,  R.  G.  Webb,  E.  E.  WiUiams,  V. 
Wilson,  R.  D.  Worthington,  J.  W. 
Wright,  G.  R.  Zug,  and  R.  G.  Zweifel. 
Tom  Van  Devender  generously  provided 
numerous  hve  specimens  from  Sonora. 
Permission  to  collect  in  Mexico  was 
granted  through  Miguel  Angel  Hernandez 
Garcia  and  Ignacio  Ibarrola  Bejar  of  the 
Direccion  General  de  la  Fauna  Silvestre. 
For  unequaled  field  assistance,  Diderot 
Gicca,  Sheila  Iverson,  Ron  Magill,  Peter 
Meylan,  and  C.  R.  Smith  deserve  special 
mention.  Joan  and  Jill  Iverson  assisted  in 
computation  of  preliminary  data.  The 
University  of  Florida,  Florida  State  Mu- 
seum and  Earlham  College  provided  sup- 
port and  study  space.  Portions  of  the  field 
work  were  supported  by  grants  from 
Sigma  Xi,  the  Theodore  Roosevelt  Memo- 
rial Fund,  the  American  Philosophical 
Society,   the   Earlham   College    Faculty 


60 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Development  Fund,  and  the  National 
Science  Foundation  (DEB-8005586).  Jim 
Berry,  Roger  Conant,  Mike  Seidel,  and 
Hobart  Smith  each  offered  helpful  com- 
ments on  the  manuscript.  Sheila  Iverson 
typed  the  manuscript. 

RESUMEN 
Se  analizaron  las  variaciones  geograficas  del 
escudo  y  las  medidas  de  las  conchas  (mediante  ana- 
lisis  estadi'stico  multivariado),  tamaTio  del  cuerpo, 
morfologia  de  las  escamas  de  la  cabeza  y  del 
menton,  tamano  del  primer  hueso  neural,  escama- 
cio'n  irregular,  asi  como  tamano  de  la  cabeza  y  los 
patrones  de  poblacidnes  de  la  tspecit  ^  Kinosternon 
hirtipes.  Los  resultados  sustentan  la  retencidn  de  las 
especies  alopatricas  A^.  sonoriensey  K.  hirtipes cofho 
especies  completas  dentro  del  grupo,  y  el  recono- 
cimiento  de  dos  subespecies  alopatricas  (una  de  ellas 
nueva)  de  K.  sonoriense  y  de  seis  subespecies  (cuatro 
de  ellas  nuevas  y  todas  aparentemente  alopatricas) 
de  K.  hirtipes.  La  descripcion  de  cada  taxon  incluye 
datos  completes  de  sinonimias,  ecologia  y  repro- 
duccidn.  Tambien  estan  incluidas  claves  para 
adultos  y  una  discusion  de  todos  los  taxa. 

Specimen  List 

All  specimens  examined  as  well  as  local- 
ities plotted  in  Figure  1  are  listed  below  by 
drainage  basin  sample  used  in  the  analy- 
sis. Basins  are  Usted  under  the  appropriate 
taxon  in  approximate  geographic  order 
from  northwest  to  southeast.  Localities 
(including  literature  records)  within  each 
basin  are  listed  alphabetically  by  state, 
county,  and  specific  locality.  Specimens 
marked  with  an  asterisk  were  not  exam- 
ined. All  distances  are  in  km.  The  fol- 
lowing abbreviations  are  used  throughout 
the  list:  C  =  city  or  ciudad;  Cn  =  can- 
yon; Cr  =  creek;  Hwy  =  highway; 
Mtn(s)   =   mountain(s);    nr   =   near;    R 

=  river  or  ri'o;  Rd  =  road;  Spg(s)  = 
spring(s);  trib  =  tributary;  and  vie  = 
vicinity. 

K.  sonoriense     sonoriense. 

BILL  WILLIAMS  (BIG  SANDY)  RIVER  (WILL). 
ARIZONA.  Mojave  Co.:  Big  Sandy  Basin,  NW 
Wickenburg,  UAZ  30826*;  Burro  Cr  Camp- 
ground, ASU  13785;  14.5  km  E  Burro  Cr 
Campground,  ASU  13786;  Trout  Cr  (Hulse, 
1974). 


GILA  and  LOWER  COLORADO  RIVERS 
(GILA). 
ARIZONA.  Cochise  Co.:  Babacomari  R,  ca.  4.8 
km  W  Huachuca  C,  UAZ  38861*;  Bear  Cn, 
16.1  km  W  Coronado  International  Memorial, 
ASU  13783*;  Bear  Cn,  Huachuca  Mtns,  Monte- 
zema  Pass  Rd,  UAZ  27982*;  Fort  Huachuca, 
first  cienega  above  post,   USNM   17780-81*, 
19680*,  21718-19*,  45305*  (Stejneger,  1902);  nr 
Hereford,  San  Pedro  R,  KU  15927*,  CAS-SU 
48886-87*;    Huachuca   Mtns,    AMNH    19450, 
USNM  20975-77*,  20979-80*  (Van  Denburgh 
and  Slevin,  1913;  Van  Denburgh,  1922);  Lewis 
Spgs,     AMNH     15165-69,     18103,     18656-57, 
UMMZ  118269;  3.2  km  S  Miller's  Peak,  Hua- 
chuca  Mtns,    Cochise   Cn,    CAS-SU    13888*; 
Pyeatt  Ranch  nr  West  Gate  Fort  Huachuca,  JBI 
410-14;  San  Pedro  R,  USNM  20547-55*;  San 
Rafael  Valley,  UMMZ  88476*;  Hwy  80  at  St. 
David  (Kauffeld,   1943);  Vasquez  Ranch,  St. 
David,  UAZ  32960*.  Gila  Co.:  Cibecue  Cr  nr 
Salt  R,  ASU  10530*  (Hulse,  1974);  Coyote  Cn, 
ASU    10903-04*   (Hulse,    1974);   66  km   NNE 
Globe,    Salt    R,    UMMZ    105791    (Duellman, 
1955);  Mezquite  Flat  at  SaU  R,  ASU  10527-29* 
(Hulse,   1974);  Natural  Spgs,  just  N  Payson 
(J.    F.    Berry,    pers.    comm.);    Payson,    ASU 
4142*;  4.8  km  N  Punkin  Center  on  Tonto 
Creek,  ASU  12061-68*;  Rice,  San  Carlos  Indian 
Reservation,  USNM  59738*;  Roosevelt  Reser- 
voir (Little,   1940);  San  Carlos  River,  N  San 
Carlos,    UMMZ    105821    (Duellman,     1955); 
Spring  Cr,  16.1  km  W  Young,  UMMZ  105756 
(Duellman,   1955);  Tonto  Cr  nr  Gisela,  ASU 
2372*  (Hulse,  1974,  1976).  Graham  Co.:  Bonita 
Cr,  NE  of  Safford,  UMMZ  105792  (Duellman, 
1955);    Marijilda    Cr    (Nickerson    and    Mays, 
1971);  8.0  km  S  Safford  (Nickerson  and  Mays, 
1971);    9.7    km    S    Safford,    UMMZ    105765, 
105293    (Duellman,    1955);    no    further   data, 
USNM  55627-28  (Van  Denburgh,   1922  as  K. 
flavescens;    Iverson,     1978).     Greenlee    Co.: 
Virden,  1 .6  km  W  New  Mexico  State  line,  UNM 
15561.  Maricopa  Co.:  Agua  Caliente,  CAS-SU 
39102*;  Box  Cn,  8.0  km  N  Wickenburg  (Gates, 
1957);  Cave  Cr,  CAS-SU  17282*,  KU  15926*. 
UAZ   35948*;   Cave   Cr,   Fairbank,   CAS-SU 
20643*,   35157*   (Van   Denburgh  and  Slevin, 
1913;  Van  Denburgh,  1922);  Granite  Reef  Dam, 
ASU  4549*;  Guadalupe,  ASU  1972*;  Hassay- 
ampa  R,  8.0  km  S  Wickenburg,  CHAS  16177 
(Gates,  1957);  Hassayampa  R,  8  km  SE  Wick- 
enburg,  UIMNH   85839,   85842;   Mesa,   ASU 
336*;  Phoenix,  AMNH  73821-22*.  ASU  4268*. 
UMMZ   69417-20,    72497.    USNM    55625-26* 
(Van  Denburgh.  1922);  Phoenix,  Salt  R.  KU 


No. 


Kinosternon  Biosystematics 


61 


2908,  UMMZ  15755*.  USNM  15755*  (Iverson, 
1978);   48.3    km   SW    Phoenix,    Gila    R,    KU 
15928*.    Sycamore    Cr    at    Sunflower.    ASU 
13801-03*.  CM  57121.  57113-14  (Hulse.  1974, 
1976);  Sycamore  Cr  at  Hwy  87,  ASU  12105*; 
Sycamore  Cr,  1.6  km  S.  Sunflower.  UU  11537- 
39*;  Tempe.  ASU   1004*.   Navajo  Co.:   Fort 
Apache  (Hulse,  1974);  Rock  Cr  Cn,  S.  Camp 
Apache.  USNM  1103*  (Yarrow  1875  as  K.  hen- 
rici;  Van  Denburgh.  1922).  Pima  Co.:  Annilo 
Tank,  R17E,  T14S.  Sec  3,  NE  %,  UAZ  36510*; 
Arivaca,  0.8   km  SW  of  Post  Office,   UAZ 
30821,  30823;  0.8  km  E  Arivaca,  UAZ  30824; 
Madrona    Cn.    Rincon    Mtns.    UAZ    27985*, 
36512*,  FB  1551;  Molina  Basin,  Santa  Catalina 
Mtns,  UAZ  27998*;  Posta  Quemada  Cn,  SE 
side  Rincon  Mtns,  UAZ  24753*;  Rincon  Mtns, 
end    of    Kennedy    Rd    via    Speedway,    UAZ 
30825*;     Rincon    Stock    Farm,    nr    Tucson, 
UMMZ  89871-73;  Sabino  Cn,  Santa  Catalina 
Mtns,    CAS-SU    8637-38*,     FMNH    74777, 
SDNHM  14225,  UAZ  27997*  (Van  Denburgh 
and  Slevin,  1913;  Van  Denburgh,  1922);  Santa 
Catalina  Mtns,  AMNH  4520;  Tanque  Verde 
Ranch,  SDNHM  16232-37;  Tucson,  Santa  Cruz 
R,  AMNH  2565,  20538,  CAS-SU  33850-66*, 
MCZ  1920,  USNM  67*,  17018-21*,  16835-36* 
(LeConte,   1854;  Agassiz.   1857;  Baird.   1859; 
Yarrow.  1883;  Gunther.  1885;  Van  Denburgh 
and  Slevin.  1913;  Van  Denburgh,  1922);  Tucson 
Sewage  Disposal  Area,  UAZ  28002*;  nr  Xavier, 
16.1  km  S  Tucson,  CM  19287.  Pinal  Co.:  Boyce 
Thompson  SW  Arboretum,  6.4  km  W  Superior, 
AMNH   66336,   CHAS   9494-97.   9644.   9648. 
10324.  UMMZ  85076  (14  specs);  Queen  Cr, 
Arboretum,  CHAS  9879-80,  13634-44;  Superi- 
or, CHAS  10325.  UAZ  27994-95*.  Santa  Cruz 
Co.:    Alamo   Cn.   4.0  km   SW   Pena   Blanca 
Camp.  Pajarito  Mtns,  MVZ  50903-06.  UAZ 
15104*;  Babacomari  R  at  Babacomari  Ranch. 
ASU  12107-113*;  G.  A.  Jones  Ranch  at  Parker 
Cn.  UAZ  27986*;  Lochiel,  ASU  13804*;  Mon- 
key Spg.  ASU  12077*;  Nogales.  USNM  17127- 
36*.  ASU  13787*  (Van  Denburgh.  1922);  19.3 
km  W  Nogales.  CM  25209;  6.8  km  S  Patagonia 
on  Hwy  82.  LACM  64223;  Pena  Blanca  Spg. 
TUL  15040-41,  UMMZ  75814,  75855  (Camp- 
bell, 1934);  Santa  Rita  Mtns.  CAS-SU  48885* 
(Van  Denburgh.  1922);  SW  of  Tucson.  AMNH 
2559-62.   UMMZ   118268;   Tumacacori   Mtns. 
SDNHM  5720,  CAS-SU  81457-58*;  Turkey  Cr 
at  Canelo,  UAZ  27988*.  Yavapai  Co.:  Bard, 
SDNHM    33866;    12.9    km    S    Camp    Verde, 
SDNHM  17889;  4.8  km  N  Clarksdale,  Verde  R, 
UU    15078-84*;   Ft.   Verde,   USNM   14807-09, 
15708  (Van  Denburgh,  1922  as  K.  flavescens 


and  K.  sonoriense;  Iverson,  1978);  Fossil  Cr, 
9.7  km  N  Verde  R,  ASU   12151-56*  (Hulse, 
1974);    Hassayampa    R    at    Wagoner,    CHAS 
16631;   Hassayampa   R.   3.2  km  S  Wagoner. 
CHAS  15834;  Montezuma's  Well.  ASU  4573*. 
UU  13031*;  Peck's  Lake,  NE  Clarksdale,  JBl 
386-88;   Rock  Spgs,  CM  47751.   MSU   3578; 
Stehr  Lake,  ASU  13790*;  Sycamore  Cr.  E  of 
Dugas,    UMMZ    105822    (Duellman.     1955); 
Sycamore  Cr  at  Verde  R,  ASU  12074-76*;  Tule 
Stream,   ASU    10962-67*,   CM   57115.    57122 
(Hulse,    1974,    1976);    Entrance    to    Tuzigoot 
National  Monument,  ASU  13789*;  Verde  R. 
above     Camp     Verde.     UMMZ     105823* 
(Duellman.  1955);  Verde  R  in  Cottonwood.  JBl 
524.    Yuma    Co.:    Gila    C.    Gila    R,    USNM 
21716-17*,   21817*;   Gila  R.   Adonde  Siding, 
USNM  21715*  (Van  Denburgh.  1922);  North 
Gila  East  Main  Canal,    1.6  km  SW  Laguna 
Dam,  RSF  468*  (Funk,  1974);  Warshaw,  Mex. 
Boundary         line,         USNM         21712- 
14*;  Yuma  (Van  Denburgh  and  Slevin,  1913; 
Van  Denburgh,  1922). 
CALIFORNIA.  Imperial  Co.:  Palo  Verde,  MVZ 
6282  (Van  Denburgh,  1922);  No  further  data, 
CAS-SU  33408  (Van  Denburgh  and     Slevin, 
1913;  Van  Denburgh,  1922). 
NEVADA.  Clark  Co.:   Pyramid  Cn  (LaRivers, 
1942,  as  K.  flavescens,  but  see  Iverson,  1978). 
NEW   MEXICO.   Catron  Co.:   Glenwood,   San 
Francisco  R,  CM  18310;  Taylor  Cr,  2.4  km  NE 
Wall    Lake,    UMMZ    134282-84,    UNM    2568 
(Niles,    1962);    Wall   Lake,    13.7   rd   km   SSE 
Beaverhead,  UMMZ  134281,  UNM  20552,  20609-10 
(Niles,     1962).    Grant    Co.:    Bennett    Ranch.    W 
Cliff.  UNM  8157-69;  3.2  km  ENE  Cliff  (Niles, 
1962);  1.6  km  E  Bedrock  Post  Office,  S  side 
Gila  R,  UNM  20611.  Undetermined  Co.:  Gila 
R,  ANSP  83  (holotype  of  Kinosternum  henrici). 
SONORA.  R  Nutrias,  above  Nutrias  Dam,  UM- 
MZ 105817;  R  San  Pedro,  above  Elias  Dam, 
UMMZ  105816,  105818-20;  R  Santa  Cruz.  6.4 
km  S  Arizona  border.  UMMZ  105814-15;  San 
Pedro  R,  USNM  20968  (Van  Denburgh.  1922); 
Sierra  Magallones.  UAZ  36497*. 
SW  NEW  MEXICO  INTERIOR  DRAINAGES 
(SWNM). 
ARIZONA.  Cochise  Co.:  N  of  Rodeo,  nr  New 
Mexico  border,  UMMZ  86081-86  (Niles,  1962). 
NEW  MEXICO.  Hidalgo  Co.:  Clanton  Cn,  16.1 
km  N  Cloverdale.  LACM  7967-70,  7994;  8  -  9.7 
km  W  Cloverdale  Store,  UNM  20558;  Guada- 
lupe Cn,  3.1  km  E,  2.3  km  N  Arizona-New 
Mexico  border,  UNM  14061;  W  slope  Pelon- 
cillo  Mtns,  T32S,  R21W,  Sec  16,  NE  '/a,  UNM 
15618;  24.1  km  N  Rodeo,  San  Simon  Marsh. 


62 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


NMSU   3050*;    San   Simon   Cienega,    UMMZ 
105800  (Niles,   1962);  Skeleton  Cn,  Peloncillo 
Mtns,  AMNH  109056,  MVZ  70350. 
RIO  MAGDALEN  A  (MAGD). 

ARIZONA.  Santa  Cruz  Co.:  California  Gulch, 
ASU  13633-37*.  CM  57116-20;  Ruby,  UIMNH 
4129,  UMMZ  107480  (Dueliman,  1955);  Syca- 
more Cn,  UAZ  28000*,  30822*,  33582*. 

SONORA.  Imuris.  UIMNH  85832;  14.5  km  N. 
Imuris,  KU  44503-25;  14.5  km  NNE  Imuris, 
KU  48562-63,  50734*,  51429;  1.1  km  S  Magda- 
lena,  UAZ  28010;  nr  Magdalena,  MCZ  46649*; 

25.1  km  NNE  Magdalena,  UMMZ  126442;  42 
km  S  Nogales,  Rancho  de  Tascara,  AMNH 
73004;  69.5  km  S  Nogales  on  Hwy,  2,  LACM 
61107;  R  Arizona,  vie.  Rancho  de  la  Arizona, 
UAZ  28010-11;  R  Magdalena,  1.6  km  SE  Cab- 
orca,  MVZ  51355  (Zweifel  and  Norris,  1955). 

RIO  SONORA  (SNRA}. 

SONORA.  Arispe,  UAZ  27976,  28003-07,  28012- 
14,  28016-18,  28020-21;  24.1  km  W  Cananea, 
AMNH  67503-05,  67507;  4.8  km  downstream 
from  Cucurpe,  UAZ  36509;  Hermosillo,  AM- 
NH 74945;  24.1  km  N  Hermosillo  (Taylor, 
1936);    Cjenega    nr    Rancho    Agua    Fria,    E 

Cucurpe,  'jBI    799-803,   866-870;    16.1    km   E 

Ures,  R  Son         ora,  NMSU  4101*. 

RIO  YAQUI.  (YAQ). 
ARIZONA.  Cochise  Co.:  Ashton  Spg,  nr  San 
Bernardino  Ranch,  UAZ  28001*;  Black  Dam, 
San  Bernardino  Ranch,  UAZ  27999*;  Chirica- 
hua  Mtns,  USNM  33929-30*  (Van  Denburgh, 
1922);  8.0  km  S  McNeal  on  Hwy  666,  LSU 
9861;  San  Bernardino  Ranch,  27.4  km  E  Doug- 
las, CM  40407,  ASU  13784*;  San  Bernardino 
Ranch,  Mex.  boundary,  USNM  21104*;  nr 
Turkey  Cr  Ranger  Station,  UMMZ  105675 
(Dueliman,  1955). 
CHIHUAHUA.  Bavispe  R,  below  3  Rivers, 
Chihuahua-Sonora  border,  BYU  14629;  R 
Gaviian,  11.3  km  SW  Pacheco,  MVZ  46646. 
SONORA.  Guadalupe  Cn,  nr  Monument  72, 
Mex.  boundary  line,  USNM  20970  (Agassiz, 
1857;  Baird,  1859;  Yarrow,  1883;  Van  Den- 
burgh and  Slevin,  1913;  Van  Denburgh,  1922); 

14.2  km  W  Maicova,  UAZ  39968;  Ranchito 
Finos  Altos,  Sierra  Nacori,  UAZ  31613-14; 
San  Bernardino  Ranch,  USNM  20981-88  (Van 
Denburgh,  1922).  Yecora,  UAZ  28211,  35209- 
11*;  18.0  km  E  Yecora,  UAZ  40105. 

Rio  CASA  GRANDES  INTERIOR  BASIN  (CSGR). 
CHIHUAHUA.  3.2  km  N  Old  Casas  Grandes, 
BYU  14132-33;  Colonia  Juarez,  R  Piedras 
Verdes,  FMNH  1873  (2),  UNM  30393-99,  UU 
11522-36;  2.6  km  NW  Colonia  Juarez,  UF 
47642-43,  JBI  946-47;    10.5  km   NW  Cohania 


Juarez,  ASU  5207-08*;  Ramos,  MVZ  46647-50. 
RIO  FUERTE  (FRTE). 
CHIHUAHUA.  Cerocahui,  BYU  14625,   14627, 
14628  (see  text). 
QUESTIONABLE  DATA. 

JALISCO.   12.1   km  N  Magdalena,   BYU   14630 
(Tanner  and  Robison,  1960). 

Kinosternon  sonoriense  longifemorale 

RIO  SONOYTA  (SNTA). 
ARIZONA.  Pima  Co.:  Quitobaquito  Pond,  JBI 
391,  696-699,  701-706,  UF  47719-20  (para- 
types);  Organ  Pipe  National  Monument  Col- 
lection (4  uncatalogued  specimens),  LACM 
105399,  SDNHM  47316,  UAZ  27987  (para- 
types,)  27993,  27996  (paratypes)  (Stebbins, 
1966). 
SONORA.  Sonoyta,  USNM  21709-11  (paratype, 
holotype,  and  paratype,  respectively);  Sonoyta 
R,  USNM  21725;  Sonoyta  R,  4.8  km  from  Son- 
oyta, USNM  21708  (Van  Denburgh,  1922) 
(paratype);  29.0  km  W  Sonoyta  on  Hwy  2, 
LACM  105400. 

Kinosternon  hirtipes  murrayi 

RIO  SANTA  MARIA  INTERIOR  BASIN  (STMR). 
CHIHUAHUA.  Galeana,  R  Santa  Marfa,  BYU 
15266-76;  nr  Galeana,  R  Santa  Maria,  BYU 
16846-47,  UMMZ  117783-84  (Semmler  et  al, 
1977);  4.8  km  N  and  3.2  km  W  Galeana,  R 
Santa  Mari'a,  UU  4457-80,  1251 1;  ca.  4.8  km 
SE  Galeana,  UAZ  36349*;  9.7  km  NW  Gale- 
ana, R  Santa  Maria,  MCZ  62516-22;  Ojo  de 
Galeana,  7.2  km  SE  Galeana,  ASU  5169-82*, 
5185-95*,  FB  1695*,  1844*,  JBI  808-09,  815-20, 
838-43,  850,  958-61,  UAZ  27965-70*,  34766*, 
UF  40536-49,  UNM  32600-12;  outHow  of  Ojo 
de  Galeana,  3.4  km  S  Galeana,  ASU  5196-205*; 
nr  Progreso,  R  Santa  Maria,  UMMZ  118284- 
89,  USNM  105026-28,  105031-34;  R  Santa 
Maria,  USNM  30841-43;  San  Buenaventura, 
below  Presa  El  Tintero,  R  Santa  Man'a  (Casas 
Andreu,  1967). 

RIO  CARMEN  (  =  SANTA  CLARA)  INTERIOR 

BASIN  (CRMN). 

CHIHUAHUA.  3.2  km  W  Carmen,  R  Carmen, 
UU  8539-43;  1.6  km  S  and  0.8  km  E  Santa 
'  Clara,  R  Santa  Clara,  MVZ  72819-43, 
89676-77;  3.2  km  S  Santa  Clara,  MVZ 
70688-95;  R  Carmen  at  Ricardo  Flores  Magon, 
UMMZ  125362. 

RIO  SAUZ  INTERIOR  BASIN  (SAUZ). 
CHIHUAHUA.  Arroyo  El  Sauz,  El  Sauz,  UU 
8549-53;  5  mi  N  Cerro  Campana,  MVZ  68915; 


No.  1 


Kinosternon  Biosystematics 


63 


nr  Encinillas,  UMMZ  117781-82,  117785;  Ojo 
Laguna,  MVZ  70696-98;  Sauz,  FMNH  1405  (5); 
UMMZ  117426-29. 

ALAMITO  CREEK  DRAINAGE.  (TEX). 
TEXAS.  (See  discussion  in  Conant  and  Berry, 
1978).  Presidio  Co.:  Casa  Piedra,  Willie  Russell 
Ranch,  DMNH  985,  1095-96;  Marfa,  USNM 
15860  (paratype)  (Glass  and  Hartweg,  1951) 
(data  obviously  in  error;  see  Conant  and  Berry, 
1978);  48.3  km  S  Marfa,  Harper  Ranch,  USNM 
198055;  59.5  km  S  Marfa,  Harper  Ranch, 
TCWC  650  (holotype)  (Glass  and  Hartweg, 
1951);  60.3  km  SSE  Marfa,  UMMZ  S1083, 
101294  (paratypes)  (Glass  and  Hartweg,  1951; 
Peters,  1952). 

Rib  CONCHOS  (CNCH). 
CHIHUAHUA.  Boquilla  Culebra,  UIMNH 
52198  (Smith  et  al,  1963);  1.6  km  N  Camargo, 
UU  8548;  8  km  N  Camargo,  UMMZ  118075; 
20  km  W  Camargo,  Arroyo  del  Vado  o  La  Pal- 
oma,  Presa  La  Boquilla  (Casas  Andreu,  1967); 

27.4  km  SW  Camargo,  UU  8469-89,  8490-98; 

27.5  km  SW  Camargo,  UIMNH  43528;  R  Cata- 
lina,  24.1  km  N  Villa  Ocampo,  Durango,  UU 
12758-59;  8  km  N  Chihuahua,  MVZ  66121*; 
8  km  N  Falomir,  UIMNH  52199-201  (Smith  et 
al,  1963);  0.8  km  N  Guadalupe  Victoria,  KU 
51237-38*,  51259-60;  Guardiola,  UIMNH 
52194-97  (H.M.  Smith  et  al,  1963);  4.8  km  S 
Hidalgo  del  Parral,  UU  8468;  12.9  km  SW 
Hidalgo  del  Parral,  TCWC  20812;  4.8  km  SW 
Jimenez,  KU  53758-84;  Julimes,  ANSP  20106- 
08,  UIMNH  52190-93,  UU  8546-47  (Smith  et 
al,  1963);  9.7  km  NE  La  Boquilla,  UNM  467; 
0.8  km  E  La  Cruz,  KU  48259-62;  0.4  -  1 .6  km  E 
La  Cruz,  UIMNH  43511-27  (Williams  et  al, 
1963);  cited  erroneously  as  Lago  Toronto  by 

Casas  Andreu,  1967);  Meoqui,  R  San  Pedro, 
MVZ  52256;  8  km  N  and  8  km  E  Meoqui,  KU 
33903*;  nr  Ojinaga,  AMNH  113858-59*  (Con- 
ant and  Berry,  1978);  1.6  km  NW  Ojinaga,  KU 
52159,  69849  (Legler,  1960);  R  San  Pedro,  78.8 
km  SE  Chihuahua,  MVZ  57467;  Mouth  of  R 
San   Pedro,    KU   51221-33,    51239-56,    51276, 
51291-98,     51316-20,     52147-57,     56163-64, 
9136572  (Legler,  1960);  1.6  km  upstream  from 
mouth  of  R  San  Pedro,  KU  51234-36,  51257-58; 
12.9  km  SE  Santa  Barbara  at  Rafael,  AMNH 
6792325;  Santa  Rosalia,  FMNH  5930  (2). 
DURANGO.  4.8  km  E  Las  Nieves,  R  Florido, 
MSU  3180-89. 
LAGUNA  BUSTILLOS  INTERIOR  BASIN  (BUST). 
CHIHUAHUA.  27.4  km  N  Cuauhtemoc,  trib  to 
Laguna  Bustillos,  UMMZ  125358-61. 
RIO  PAPIGOCHIC  DRAINAGE  (PAP). 
CHIHUAHUA.  8  km  N,  1.6  km  W  Cd  Guerrero, 


R  Papigochic,  KU  45020-25,  51425-26,  87854; 
El  Riyito,  17.7  km  WNW  Cocomorachic,  KU 
51311,  51313-14;  Minaca,  FMNH  1102,  MVZ 
58967-70;  3.2  km  W  Minaca,  KU  51261-309, 
52142-43,  87853,  91364,  91373-78;  5.5  km 
NE  Minaca,  BYU  16848;  Ri'os  Papigochic  and 
Tomochic  (Legler  and  Webb,  1970;  erroneous- 
ly recorded  as  K.  sonoriense  and  K.  hirtipes  hir- 
tipes);  Yepbmera,  FB  1545-46,  1595-97,  JBI 
403-404,  MSU  3579,  UAZ  34168*;  1.6  km  N 
Yepbmera,  JBI  821-23,  835-37,  UF  40389-400, 
UNM  32588-599;  3  km  N  Yepbmera,  UAZ 
34169-70*;  3  km  W  Yepdmera,  MCZ  79029-38, 
79039-46;  4  -  5  km  N  Yepomera,  Arroyo  de  la 
Huachin,  UAZ  34171-72*. 

RIO  NAZAS  INTERIOR  DRAINAGE  (NAZ). 
DURANGO.  Lerdo,  USNM  61687-88;  24.1  km 
SW  Lerdo,  AMNH  67496-500,  UMMZ  118267; 
between  Lerdo  and  La  Goma,  USNM  105262- 
64;  R  Nazas,  at  Cardenas  Dam,  nr  El  Palmito, 
JBI  826-31,  UU  8461-66;  22.5  km  NE  Pedriceffa 
UIMNH  19339;  La  Concha,  nr  Penon  Blanco, 
AMNH  88883;  Presa  Francisco  Zarco  on  R 
Nazas  nr  Graseros,  ENCB  10893-94,  JBI  948- 
50,  UF  47602;  Trib  to  R  San  Juan  at  Hwy  45, 
5.6  km  N  turnoff  to  Primo  Verdad,  UU  12075- 
77;  Rodeo,  AMNH  87654-57,  96589;  13.5  km 
S  San  Jacinto,  R  Nazas,  UF  40425-27;  16.1  km 
W  Torreon,  R  Nazas,  USNM  105270-71. 

RIO  AGUANAVAL  INTERIOR  DRAINAGE 

(AG  UN). 
ZACATECAS.  24.1  km  NW  Fresnillo,  R  Florido 
AMNH  85285-91;  25.7  km  N  Fresnillo,  UMMZ 
118056-057,  and  118060*;  La  Florida,  R  Flor- 
ido, UU  12078-80;  Rancho  Grande,  R  Medina, 
AMNH  85296;  1.6  km  N  Rancho  Grande,  R 
Nieves,  UU  8499-538,  8544-45;  17.7  km  E  Som- 
brerete,  UIMNH  28155;  46.7  km  E  Sombrerete, 
UMMZ  126284. 

LAGO  SANTIAGUILLO  INTERIOR  DRAINAGE 

(STGO). 
DURANGO.   22.5   km  SE  Chinacates,  AMNH 
88882;  trib  to  Lago  Santiaguillo,  at  bridge  in 
Guatimape,  UF  40428-30. 

RIO  MEZQUITAL  DRAINAGE  (MEZ). 
DURANGO.  ca.  5  km  from  Colonia  Hidalgo,  km 
937,  Torrebn-Durango  Hwy  (Casas  Andreu, 
1967);  4.8  km  E  Durango,  AMNH  85294;  9.7 
km  E  Durango,  R  Tunal,  AMNH  85292-93; 
10.5  km  E  Durango,  R  Tunal,  UU  4481-520, 
12512-15;  15.8  km  N  Durango,  UIMNH  7051, 
23844;  16.1  km  N  Durango,  R  Canatlan,  MVZ 
57333-35;  17.7  km  E  Durango  on  Hwy  45,  TUL 
18680;  17.7  km  E  Durango,  R  Santiago,  MVZ 
58222;  6.4  km  E  and  11.3  km  S  Durango,  R 
Santiago,  MSU  4245-56;  25.4  km  SW  Durango, 


64 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


R  Chico  on  Hwy  40,  LSU  34319,  JBI  954-55, 
UF  47603-04;  ENCB  10904-08;  27.4  km  N  Dur- 
ango,  CU  46115-16;  37  km  N  Durango,  MSU 
7869;  kilometer  48.5,  N  of  Durango,  Hwy  45, 
UF  40424;  nr  Durango,  6.4  km  E  and  3.2  km 
NE  jet.  hwys  to  Torreon  and  Fresnillo,  UMMZ 
122245-54;  0.8  km  N  Graceros,  KU  68733-36, 
68738-45  (KU  68737  is  K.  integruml);  6.4  km 
SW  La  Pila,  KU  51083-84,  MSU  2680-82,  2684, 
2686-89,   10197-98;  9.7  km  NW  La  Pila,  KU 
51085-86;  R  Mezquital,  at  Mezquital,  86.7  km 
SSE  Durango,  TUL  18670;  6.4  km  S  Morcillo, 
MSU  4243-44  (basis  of  Stebbins'  1966  southern 
Durango  /lavescens  record;  see  Iverson,  1978); 
Ojo  de  Agua  de  San  Juan,  1 .6  km  N  Los  Berros, 
UMMZ  129824-28;  Otinapa,  AMNH  68382;  R 
La  Sauceda  at  Hwy  40,  ENCB  10894-903,  JBI 
825,  832-34,  951-53,  UF  40401-23,  UNM  32588- 
99;  R  Soledad,  La  Soledad,  MSU  2683,  2685; 
6.4  km  S  Villa  Union  jet.  Hwy  45,  CM  53987. 
EL  SALTO  (ACAPONETA)  BASIN  (SALT). 
DURANGO.  9.7  km  ENE  El  Salto,  Hwy  40. 
ENCB  10909-14,  JBI  956-57,  LSU  34320,  UF 
47605-06. 
RIO  SANTA  MARIA  BASIN  (SLP). 
SAN  LUIS  POTOSI.  Laguna  de  las  Rusias,  LSU 
7873-75  (Williams  and  Wilson,  1966);  Arroyo 
la  Hilada,  ca.   1  km  N  Presa  El  Refugio  (  = 
Laguna  de  las  Rusias),  UF  42803-815. 
Rfo  AGUASCALIENTES  DRAINAGE  (AGUAS). 
AGUASCALIENTES.    Aguascalientes,    MCZ 
79047;  Aguascalientes,  R  Morcinique,  MU  793; 
2.1   km  E  Aguascalientes,  UIMNH  43582;  R 
Penueia  nr  Aquido,  CAS-SU  19702-03;  R  Jo- 
coque  Dam,  SE  end  Presa  Jocoque,  CAS-SU 
19692-95;    1.2   km   W  Santiago,    R  Jocoque, 
CAS-SU  19696-701. 
RIO  VERDE  DRAINAGE  (VERDJ. 
JALISCO.  El  Olivo,  19.3  km  W  Lagos  de  Mo- 
reno, AMNH    117953;   Presa  el  Cuarenta  nr 
Paso  de  Cuarenta,  JBI  896-900,  UF  44064-65, 
44078;  3.2  krn  NE  Valle  de  Guadalupe,  trib  to  R 
Verde,  Hwy  80,  JBI  893-95,  TUL  18671,  UF 
44077. 
MARAVATIO  BASIN  (MAR/. 
GUANAJUATO.  1.6  km  SE  Inchamacuaro,  KU 
43637. 
BAJIO  BASIN  (BAJ). 
GUANAJUATO.    No    further   data   (Westphai- 
Castelnau,  1872);  R  Turbio,  12.9  km  E  Pen- 
jamo,  UU  12081-82;  R  Urdo,  Valle  de  Santiago 
(Caballero  y  C.  y  Cerecero,  1943;  Caballero  y 
C,  1940a);  16.1  km  N  San  Miguel  de  Allende, 
AMNH  93363;  22.5  km  N  San  Miguel  de  Allen- 
de, AMNH  85295;  Arroyo  el  Sauz,  ca.  10.5  km 


N  Yuriria-Salvatierra  Hwy  (Casas  Andreu, 
1967);  Taboado,  9.7  km  NW  San  Miguel  Al- 
lende, AMNH  71033,  FMNH  71029;  Hwy  51, 
6.0  km  S  jet.  Hwys  51  and  110,  UF  43613-15; 
11.9  km  S  jet.  Hwys  51  and  110  at  Sebastian, 
UF  44074,  JBI  908. 

JALISCO.  R  Lerma,  0.8  km  NW  jet.  Hwys  90 
and  110,  UU  12120. 
LAKE  CUITZEO  INTERIOR  BASIN  (CUIT). 

MICHOACAN.   Lago  Cuitzeo  (Casas  Andreu, 
1967);    Lake  Cuitzeo,   San   Agustin,    UMMZ 
97136  (Duellman,  1961). 
VILLA  VICTORIA  BASIN  (VILLA). 

MEXICO.    11.3   km  W  Villa  Victoria,   USNM 
108719-26,  UMMZ  118295-296;  3.7  km  S  La 
Presa,  JBI  928;  8.9  km  S  La  Presa,  JBI  927. 
RIO  BALSAS  DRAINAGES  (BALS). 

MICHOACAN.  8  km  W  C  Hidalgo,  AMNH 
62257  (UIMNH  24707  from  the  same  locality 
is  K.  integrum,  not  K.  hirtipes,  as  listed  in 
Duellman,  1961). 

PUEBLA.  Trib  to  R  Atoyae,  4.5  km  S  Molcaxae, 
UU  2096  (Data  questionable). 


Kinosternon  hirtipes  megacephalum 

VIESCA  INTERIOR  BASIN  (VCSA). 
COAHUILA.  3.2  km  SE  Viesea,  SM  11460-66 
(paratypes  and  holotype);  9.7  km  SW  Viesea, 
SM  9823  (paratype). 

Kinosternon  hirtipes  tarascense 

LAGO  PATZCUARO  INTERIOR  BASIN  (PATZ). 
MICHOACAN.  Lago  Patzeuaro,  FMNH  1397, 
2036,  JBI  880-84,  UF  43505-07  (paratype,  holo- 
type, and  paratype),  43595-96  (paratypes), 
UMMZ  96988-91,  97131,  99762,  1 17798  (Duell- 
man, 1961);  Lago  Patzeuaro,  nr  E  end,  UF 
7075;  Isla  Janitzio,  Lago  Patzeuaro,  CU  16142; 
Canal  de  la  Tzipecua,  SW  margin  Lago  Patz- 
euaro (Casas  Andreu,  1967);  Tzintzuntzan, 
AMNH  82128. 

Kinosternon  hirtipes  magdalense 

SAN  JUANICO  (  =  MAGDALENA  or  TOCUMBO) 

VALLEY  INTERIOR  BASIN  (SNJ) 
MICHOACAN.  Atop  Presa  San  Juanico  (road  to 
dam  meets  Hwy  15  ca.  56.3  km  W  of  Zamora), 
TUL  18677  (paratype);  Presa  San  Juanico,  at 
dam,  UF  45035-36  (holotype  and  paratype), 
45038-40  (paratypes),  and  45041. 


No. 


Kinosternon  Biosystematics 


65 


Kinosternon  hirtipes  hirtipes 

VALLEY  OF  MEXICO  (VALLE) 

DISTRITO  FEDERAL.  Mexico  C.  Senck  47875* 
(Greene,  1972);  vie  Mexico  C  (Beltz,  1954); 
San  Juan  Tezompa,  19.3  km  E  Xochimilco, 
UMMZ  99446-60;  Valley  of  Mexico,  Xochimil- 
co, USNM  61247;  Xochimilco,  UMMZ  69264 
(Caballero  y  C,  1939);  Lake  Xochimilco,  nr 
Mexico  C,  MCZ  7866,  UMMZ  80356-57. 

MEXICO.  Chalco,  FMNH  1406  (Gadow,  1908); 
Teotihuacan,  San  Juan,  AMNH  17859-62; 
Lake  Texcoco,  nr  Mexico  C,  AMNH  68699; 
Valle  de  Mexico,  CAS-SU  5849-50  (Martin 
del  Campo,  1938;  Hartweg  and  Glass,  1951; 
Deevey,  1957;  Kranz  et  al,  1970). 

STATE  UNCERTAIN.  "Mexico",  ZSM  1374/0 
(Holotype  of  Cinosternon  hirtipes;  Wagler, 
1830). 

Kinosternon  hirtipes  chapalaense 

LAGO  DE  CHAP  ALA  BASIN  (CHAP). 
JALISCO.  Lago  de  Chapala,  Beach  at  Chapala, 
UMMZ  97190;  Lago  de  Chapala,  0.4  km  off 
Chapala,  UMMZ  97121-130  (includes  holo- 
type and  paratypes);  Lago  de  Chapala,  3.2  km 
W  Chapala,  UU  12126-28;  (paratypes)  Lago  de 
Chapala,  0.8  km  E  Tuxcueca,  JBI  890;  Lago 
de  Chapala,  6.1  km  W  Ajijic,  UU  12125;  para- 
type  3.2  km  S  Jamay,  AMNH  17856;  3.2  km 
SE  Ocotlan  (El  Fuerte),  UMBM  2403;  Ocotlan, 
UMMZ  76129,  1 17796-97  (UMMZ  1 17801  from 
this  locality  is  K.  integrum.) 
MICHOACXn.  Jiquilpan  (Duellman,  1961);  La 
Palma,  USNM  108718  (Duellman,  1961). 

LAGO  DE  ZAPOTLAn  INTERIOR  BASIN 

(ZAPO). 

JALISCO.  1.6  km  NW  C  Guzman,  Lago  de 
Zapotlin,  UMMZ  1 17259-66;  3.2  km  N  C  Guz- 
man, UMMZ  102154;  Laguna  Zapotlan,  BM- 
NH  1906.6.1.253-5*  (Gadow,  1908  as  K.  inte- 
grum). 

Kinosternon  hirtipes  chapalaense  x  murrayi 

RIO  DUERO  DRAINAGE  (DUER). 
MICHOACAN.    Lake  Camecuaro,    14.5   km  E 
Zamora,  JBI  885-889,  UF  43603-610,  44062-63, 
44075-76,  UMMZ  97132-35,  102150-53  (Duell- 
man, 1961). 

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LIFE  HISTORY  OF  ETHEOSTOMA  COOSAE  (PISCES:  PERCIDAE) 
IN  BARBAREE  CREEK,  ALABAMA 

PATRICK  E.  O'NEIL 

Geological  Survey  of  Alabama, 

P.O.  Drawer  O, 

University,  Alabama  35486 


Abstract 

Collections  of  Eiheostoma  coosae  (Coosa  darter) 
were  made  from  April  1977  to  April  1978  in  Bar- 
baree  Creek  (Coosa  River  system),  Clay  County, 
Alabama.  The  principal  habitat  of  this  species  was 
cobble  and/or  gravel  raceways,  riffles  and  pools. 
Spawning  occurred  on  the  surface  of  rocks  and  small 
boulders  from  mid  March  to  late  May  with  peak 
activity  in  mid  April.  The  spawning  position  was 
either  inclined  or  semi-inverted  horizontally.  Indi- 
viduals reached  sexual  maturity  and  spawned  by  the 
first  year.  Maximum  age  was  three  years.  By  the  end 
of  the  second  year,  the  average  size  of  males  and 
females -was  41 .0  mm  and  36.5  mm  SL,  respectively. 
The  sex  ratio,  1:1.3,  was  significantly  different  from 
1:1.  The  principal  diet  consisted  of  Copepoda, 
Cladocera,  Ephemeroptera  (Isonychia)  and  Diptera 
(Chironomidae,  Simuliidae). 

INTRODUCTION 
Eiheostoma  (Vlocentra)  coosae  is 
endemic  to  the  Coosa  River  system  of 
Alabama,  Georgia,  and  Tennessee.  The 
biology  of  the  subgenus  as  a  whole  is 
largely  unknown.  Published  papers  in- 
clude studies  by  Winn  (1958a,  1958b)  on 
the  reproduction  of  two  undescribed 
forms.  Stiles  (1975)  on  the  reproductive 
behavior  of  Etheostoma  simoterum,  and 
Ultsch  et  al.  (1978)  on  habitat  selection  by 
Etheostoma  duryi.  This  study  reports  on 
the  life  history  of  E.  coosae  in  Barbaree 
Creek,  an  eastern  Alabama  stream. 

STUDY  AREA 
A     section     along     Barbaree    Creek, 
T.18S.,    R.7E,    Sec.    22,    Clay    County, 
Alabama  (Coosa  River  system)  was  cho- 


sen as  the  study  site.  Barbaree  Creek  is  a 
perennial  stream  flowing  through  north- 
ern Piedmont  physiography.  Its  head- 
waters originate  in  the  Talladega  Moun- 
tains. 

The  substrate  consisted  of  gravel  and 
sand  shoals  interspersed  with  patches  of 
cobble  and  boulders  that  were  regularly 
broken  by  cobble  or  slab  riffles.  Bedrock 
was  usually  exposed  below  riffles  whereas 
the  pools  contained  unconsolidated  mat- 
erial. The  ranges  of  measured  water 
quality  values  were:  disssolved  oxygen, 
7.9-13.0  ppm;  pH,  6.8-7.3;  turbidity, 
0.7-2.8  JTU;  conductivity,  17-45  umbos; 
and  stream  temperature  3.7-25  °C 
(Boschung  and  O'Neil,  1980). 

METHODS 

Specimens  of  E.  coosae  were  collected 
monthly  from  April  1977  through  April 
1978.  Small  mesh  minnow  seines,  3/16 
inch  delta  weave,  and  a  backpack  shocker 
were  each  operated  approximately  1.5 
hours  during  each  monthly  collection, 
sampling  a  variety  of  habitats.  Upon 
capture  the  fishes  were  preserved  in  a 
20-percent  formalin  solution. 

In  the  lab,  fishes  were  blotted  dry  and 
then  weighed  to  the  nearest  .01  g  on  a 
Mettler  electronic  balance.  Standard 
length  (SL)  and  sex  were  determined.  The 
specimens  are  deposited  in  the  University 
of  Alabama  Ichthyological  Collection. 

Aging  to  year  class  was  determined  by 


EDITORIAL  COMMITTEE  FOR  THIS  PAPER: 

Dr.  David  C.  HEINS,  Assistant  Professor  of  Biology,  Millsaps  College, 
Jackson,  Mississippi  39210 

Dr.  Royal  D.  SUTTKUS,  Professor  of  Biology  and  Director  of  Museum  of 
Natural  History,  Tulane  University,  New  Orleans,  Louisiana  70118 


75 


76 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


scale  annuli.  From  5  to  10  scales  per  fish 
were  analyzed  to  reduce  the  chance  of 
aging  error  due  to  the  presence  of  false 
annuli  and  regenerated  scales.  Aging  to 
month  was  accomplished  by  the  technique 
outlined  by  Page  (1974).  The  following 
symbols  were  used  to  indicate  year 
classes:  -1,  0  to  12  months;  1  + ,  13  to  24 
months;  2  +  ,  >24  months. 

Reproductive  condition  and  length  of 
the  spawning  season  were  determined  by 
field  observations  and  by  determination 
of  female  gonosomatic  indices  (GSI).  The 
GSI  is  the  ratio  of  gonad  weight  to 
corrected  body  weight.  Corrected  body 
weight  is  total  weight  minus  the  viscera 
and  gonad  weight  (Mathur  and  Ramsey, 
1974). 

Fecundity  is  defined  as  the  number  of 
ova  equal  to  or  exceeding  0.2  mm 
diameter.  A  large  number  of  ova  less  than 
0.2  mm  diameter  were  present  in  each 
ovary,  but  past  studies  of  darters  with 
retracted  spawning  seasons  (Fahy,  1954; 
Winn,  1958a;  Scalet,  1973)  have  suggested 
that  these  minute  oocytes  never  differen- 
tiate into  fully  yolked,  enlarged  ova  and 
were,  therefore,  not  spawned  that  year. 
The  smallest  differentiating  ova  of  the 
larger  egg  group  was  0.2  mm  diameter,  so 
this  size  was  used  as  the  lower  Hmit. 

For  food  and  feeding  studies,  whole 
stomachs  were  removed,  and  the  contents 
were  identified  to  family  and  enumerated. 

RESULTS  AND  DISCUSSION 
Habitat 

Etheostoma  coosae  adults  and  juveniles 
were  consistently  collected  over  rubble  in 
raceways  and  around  boulders,  near  sand 
bars  and  occasionally  in  the  foot  of  rif- 
fles. This  habitat  preference  was  main- 
tained seasonally  with  no  indication  of 
age  or  size  specific  habitat  utilization  for 
foraging  or  reproduction. 

The  basis  of  habitat  selection  by  darters 
is  influenced,  if  not  determined,  by  phys- 
iological and/or  ecological  requirements 
of  the  species.  Ultsch  et  al.  (1978)  con- 
ducted a  series  of  critical  O2  experiments 
with  six  species  of  Etheostoma  and  ob- 


served that  four  ecological  groups  exist 
with  respect  to  oxygen  requirements  ver- 
sus habitat  selection.  They  suggested  that 
one  such  group,  typified  by  E.  (Ulocentra) 
duryi  and  E.  (Catonotus)  flabellare, 
preferred  relatively  fast  water  but  main- 
tained its  ability  to  tolerate  periods  of 
hypoxia.  This  group  was  the  most  diverse 
physiologically  in  terms  of  oxygen  use 
strategies.  As  a  result  of  this,  these  darters 
maintained  a  diverse  array  of  habitat 
types.  The  applicability  of  this  expla- 
nation to  habit  selection  by  E.  coosae  Ues 
in  the  close  phylogenetic  and  ecological 
relationships  between  it  and  E.  duryi. 
Demography 

Etheostoma  coosae  was  the  dominant 
percid  species  in  Barbaree  Creek.  It  com- 
prised 5.9  percent  of  the  total  number  of 
fish  specimens  collected  (Table  1).  The 
overall  age  class  distribution  of  E.  coosae 
for  the  year  studied  is  seen  in  Table  2. 
Approximately  64  percent  of  the  popula- 
tion occupied  the  -  1  age  class,  29  percent 
the  1  +  age  class,  and  7  percent  the  2  + 
age  class. 

Seasonal  changes  in  age  class  compo- 
sition (Figure  1)  indicate  that  maximum 
contribution  to  population  size  occurred 
during  winter  in  the  -  1  age  class  as  it 
approached  12  months  of  age.  From  this 
point,  percent  contribution  to  population 
size  declined  throughout  the  older  age 
classes. 

Of  the  750  specimens  examined,  32.9 
percent  and  55.6  percent  of  the  males  and 
females,  respectively,  survived  from  the 
- 1  to  the  1-1-  age  class,  whereas  10.8 
percent  and  10.3  percent  of  the  males  and 
females  respectively,  survived  from  the 
1  +  to  the  2-1-  age  class  (Table  2). 

The  overall  sex  ratio,  1:1.3,  was  signifi- 
cantly different  (X  '  =  9.86;  p  <  .01)  from 
the  expected  1:1.  This  skewed  sex  ratio 
was  most  evident  in  the  1  +  age  class, 
1:1.8  (X'  =  18.96;  p  <.01),  whereas  the 
-  1  and  2  -^  age  class  sex  ratios  were  not 
statistically  different  from  1:1. 
Age  and  Growth 

The  oldest  individuals  collected,  two 
females  and  one  male  (Figure  2),  were  36 


No.  1  Etheostoma  Life  History  77 


Table  1 .  Percent  relative  abundance  and  frequency  of  occurrence  of  fishes  collected  in 
Barbaree  Creek  from  April  1977  through  April  1978. 


Species  Abundance     Occurrence 

Family  Cyprinidae 

Campostoma  anomalum 
Notropis  asperifrons 
Notropis  callistius 
Notropis  lirus 
Notropis  stilbius 
Notropis  trichroistius 
Notropis  venustus 
Notropis  xaenocephalus 
Phenacobius  catostomus 
Semotilus  atromaculatus 

Family  Catostomidae 

Hypentelium  etowanum 
Moxostoma  duquesnei 

Family  Ictaluridae 

Ictalurus  natal  is  0.04  8.3 

Family  Centrarchidae 
Ambloplites  rupestris 
Lepomis  cyanellus 
Lepomis  gulosus 
Lepomis  macrochirus 
Lepomis  megalotis 
Micropterus  coosae 
Micropterus  punctulatus 

Family  Cyprinodontidae 

Fundulus  stellifer  0.02  25.0 

Family  Percidae 

Etheostoma  coosae 
Etheostoma  jordani 
Etheostoma  stigmaeum 
Percina  caprodes 
Percina  nigrofasciata 

Family  Cottidae 

Cottus  carolinae 

Total 


1.69 

91.7 

5.87 

100.0 

8.81 

100.0 

0.19 

25.0 

2.97 

75.0 

45.62 

100.0 

0.34 

41.7 

16.90 

100.0 

0.01 

8.3 

0.09 

25.0 

2.00 

100.0 

0.39 

66.7 

0.07 

41.7 

0.01 

8.3 

0.07 

33.3 

0.04 

8.3 

0.76 

66.7 

1.11 

91.7 

0.06 

25.0 

5.93 

100.0 

1.18 

100.0 

0.97 

100.0 

0.31 

66.7 

0.96 

91.7 

2.94 

100.0 

99.35 

78 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Table  2.  Age-class  distributions  and  survival  of  Etheostoma  coosae  collected  in 
Barbaree  Creek  from  April  1977  through  April  1978.  S,  and  S2  equal  survival 
calculated  from  the  -  1  and  1  +  age  classes,  respectively. 


Sex 


Year 

Number  of 
specimens 

Surviva 

1 

class 

S, 

S, 

-1 

231 

1.000 

_ 

1  + 

76 

0.329 

1.000 

2  + 

25 

0.108 

0.329 

-1 

252 

1.000 

— 

1  + 

140 

0.556 

1.000 

2  + 

26 

0.103 

0.185 

-1 

483 

1.000 

— 

1  + 

216 

0.447 

1.000 

2  + 

51 

0.106 

0.236 

Males 


Females 


Combined 
Sexes 


months  of  age  assuming  a  May  hatching. 
Each  specimen  had  two  annuli  and  the 
third  was  in  the  process  of  being  estab- 
lished. Scale  studies  of  E.  coosae  in 
Barbaree  Creek  indicated  that  annuli  are 
established  in  early  to  middle  spring. 

Males  grew  faster  than  females,  and 
were  on  the  average  significantly  longer  (p 
<.05)  by  the  first  spawning  season 
(Figure  3).  Females  attained  70.3  percent 
and  90.0  percent  of  their  average  maxi- 
mum standard  length  (40.5  mm)  in  12  and 
24  months,   respectively,   whereas  males 


Eq 


_^a_^B_ 


M 


SUMMER 


Figure  1 .  Seasonal  changes  in  age-class  composition 
of  Eiheosloma  coosae  collected  in  Barbaree  Creek. 
Age-class  designations  are:  -  1 ,  1  to  1 2  months;  1  -i- , 
13  to  24  months;  and  2  +  ,  >24  months. 


attained  69.5  percent  and  94.5  percent  of 
their  average  maximum  standard  length 
(43.1  mm)  in  12  and  24  months.  The 
longest  male  and  female  were  47.0  mm 
and  44.1  mm,  respectively  (Figure  2). 
There  were  no  significant  differences  (p 
>.05)  between  male  and  female  length- 
weight  equations  as  tested  by  analysis  of 
covariance  (Fg  =  0.867,  df  =  1,136).  The 
relationship  for  combined  sexes  was  log 
wt.  (grams)  =  3.1657  log  SL  -5.1200,  N 
=  140,  r  =  .978. 

Page  and  Schemske  (1978)  stated  that 
one  possible  factor  affecting  size  in 
darters  of  the  subgenus  Catonotus  was 
interspecific  competition.  They  concluded 
that  maximum  size  in  sympatric  popula- 
tions of  Catonotus  was  determined  by 
the  presence  or  absence  of  Etheostoma 
squamiceps.  In  the  presence  of  E.  squaini- 
ceps,  other  Catonotus  species  were  re- 
duced in  size,  whereas  in  the  absence  of  £. 
sqamiceps  their  maximum  size  was 
greater.  They  suggested  that  the  major 
function  of  the  size  differences  was  for 
size-specific  utilization  of  potential  repro- 
ductive habitats. 

The  overall  growth  trend  of  E.  coosae 
was  for  rapid  growth,  both  sexes  reach- 


No.  1 


Etheostoma  Life  History 


79 


ing  approximately  70  percent  of  the  aver- 
age maximum  standard  length,  the  first 
year  of  life,  with  a  subsequent  reduction 
of  this  rate  in  later  years  (Figure  3).  This 
pattern  is  quite  common  in  darters  (Page, 
1974  and  1975;  Page  and  Burr,  1976; 
Starnes,  1977)  and  in  fishes  generally.  As 
Ricker  (1971)  has  pointed  out,  this  phe- 
nomenon is  usually  attributed  to  physio- 
logical size  limitations  primarily  influ- 
enced by  the  heavier  reproductive  effort 
by  older  individuals. 
Reproduction 

Female  gonosomatic  indices  (Figure  4) 
and  field  observations  indicate  that  E. 
coosae  spawned  from  mid  March  through 
early  to  mid  May  with  peak  spawning  in 
April.  The  spawning  periods  for  species 
of  the  subgenus  Vlocentra  are  similar. 
Winn  (1958a,  1958b)  reported  that  males 
of  Etheostoma  sp.  (Barren  River  form)  in 
Tennessee  established  territories  near  the 
beginning  of  April,  and  spawning  began 
in  about  one  week.  Stiles  (1975)  reported 
that  E.  simoterum  spawned  from  early 


E 

E. 

I 

»- 

O 

z 

lU 

-* 

O 

et 

< 

O 

z 

< 

^ 

A-<^ 

A 

^ 

d^ 

A 

A 

A 

o 

A 

o 

OV 

A 

A 

A 

A 

A 

O 

OA 

A 

O 

A 

o 

o 

Oft 

A 

A 

A 
OA 

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o 

o 
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*-? 

o 

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c* 

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() 

o 

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(> 

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o 

A-? 

A 

G* 

O 

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o 

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» 

o 

o 

o 

o 

O 

O 

o 

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•o 

K) 

o 

o 

o 

O* 

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o 

•o 

•o 

o 

o 

o 

O 

o 

•o 

• 

o 

QA 

O 

•u 

• 

•o 

o 

O 

• 

• 

•u 

• 

o 

O 

«> 

• 

• 

o 

m> 

• 

• 

o 

O 
O      1 

•o 
• 
• 
• 

• 

•o 

• 

• 
• 

• 

• 

• 
• 

o 

• 
• 

• 

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• 

? 

MALE 
FEMALE 

m'j'j'a's'o'n'd'j'f'm'a 


1   •  — 

1+   o  

2+    A  — 


19      20       21 
31       32       33 


MONTHS   OF    AGE 


22       23 
34       35 


Figure  2.   Length-age  relationship  of  Etheostoma 
coosae  collected  in  Barbaree  Creek. 


n</ 


</ 


c/ 


Tzr 


STANDARD 


40 

LENGTH   (p 


Figure  3.  Annual  growth  of  Etheostoma  coosae 
males  and  females  collected  in  Barbaree  Creek.  N 
equals  sample  size;  NS  equals  not  significant;  and  P 
equals  probability  level.  Solid  line  equals  range; 
triangle  equals  mean;  and  rectangle  equals  95% 
confidence  interval. 


April  to  the  end  of  May  throughout  its 
entire  range.  Etheostoma  duryi  females 
had  ripe  ova  and  males  were  in  extreme 
breeding  color  in  late  April  in  Cypress 
Creek,  Lauderdale  County,  Alabama 
(pers.  obs.). 

One  month  prior  to  spawning,  male 
testicular  tissue  was  swollen  with  a  white 
granular  appearance.  During  spawning, 
the  tissue  was  extremely  swollen  with  a 
subdued  beige  color.  Approximately  two 
months  after  spawning  the  testes  were 
reduced  to  transparent,  filamentous 
structures.  The  reproductive  cycle  of 
females  was  shorter  in  length  when  com- 
pared to  males.  Fully  yolked  mature  ova 
were  present  for  about  6  to  8  weeks. 
Mature  testes  and  fully  yolked  ova  were 
present  in  yearling  darters  indicating  the 
possibility  of  spawning  within  the  first 
year  of  Hfe. 

The  spawning  site  of  E.  coosae  con- 
sisted of  almost  any  inclined,  creviced 
surface  within  pools  or  raceways.  Aquar- 
ium observations  revealed  that  E.  coosae 
spawned  at  angles  from  0°  to  90°,  or  in  a 
horizontally  inverted  position  while 
wedged  in  rock  crevices  or  under  ledges. 


80 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


|li«ONTH 


Figure  4.  Monthly  average  gonosomatic  index  of 
Eiheostoma  coosae  females  collected  in  Barbaree 
Creek.  N  equals  sample  size. 


In  typical  percid  fashion,  a  male  would 
display  to  a  female  as  well  as  physically 
stimulate  her  nape  with  his  head,  attempt- 
ing to  elicite  a  spawning  response.  On 
occasion,  a  female  would  assume  the 
dominate  courtship  role  nudging  and  dis- 
playing her  dorsal  fins  to  a  male,  attempt- 
ing to  incite  responses  from  him.  Once  an 
egg  site,  usually  a  small  protective  nook 
or  crack,  was  selected,  the  female  arched 
her  body,  placed  the  genital  papilla  over 
the  site,  and  deposited  one  egg.  The  male 
quickly  arched  his  body,  placed  the  gen- 
ital papilla  over  the  egg  and  fertilized  it. 
The  eggs  were  deposited  with  no  defin- 
able pattern  or  arrangement. 

At  the  approach  of  spawning  season  E. 
coosae  males  acquired  a  light  aqua-green 
tint  around  the  gular  region,  along  the 
tips  of  the  spinous  dorsal  fin,  on  the  first 
three  membranes  of  the  anal  fin,  and  on 


Table  3.  Standard  length,  egg-size  distribution,  total  egg  complement  (TEC),  and  age 
in  years  of  selected  Etheostoma  coosae  females  collected  in  Barbaree  Creek  during 
March,  April,  and  May,  1977  and  1978. 


Standard 

Egg 

diameter  in  mm 

TEC 

length 

0.2-0.4 

0.5-1.0 

>1.0 

Age 

40.1 

194 

156 

72 

422 

2 

38.5 

300 

116 

80 

496 

2 

36.0 

278 

56 

4 

330 

2 

35.9 

320 

118 

54 

492 

2 

34.5 

284 

92 

34 

410 

33,7 

220 

92 

42 

354 

33.5 

172 

78 

38 

288 

31.5 

280 

46 

22 

348 

30.5 

260 

39 

0 

299 

Mean 

256.4 

88.0 

38.4 

382.8 

Percent 

67.0 

23.0 

10.0 

100.0 

No.  1 


Etheostoma  I  ife  History 


81 


the  anterior  dorsal  and  ventral  interradial 
membranes  of  the  caudal  fin.  The  inter- 
radial membranes  throughout  the  length 
of  the  spinous  and  soft  dorsal  fins  had 
rusty-red  quadrate  spots.  Females  main- 
tained a  ground  color  of  light  tan  overlaid 
by  brown  to  black  lateral  blotches  and 
mottling  above  the  lateral  line. 

The  total  egg  complement  increased 
proportionally  with  length,  r  =  .682  (Table 
3). Fecundity  studies  of  other  darters  have 
substantiated  positive  size-fecundity  rela- 
tionships: E.  squamiceps,  r  =  .692  (Page, 
1974);  E.  barbouri,  r  =  530  (Flynn  and 
Hoyt,  1979);  E.  kennicotti,  r  =  .631 
(Page,  1975);  and  Percina  nigrofasciata,  r 
=  .721  (Mathur,  1973).  An  exception  to 
this  general  relationship  was  reported  for 
E.  proeliare  (r  <.l)  by  Burr  and  Page 
(1978).  This  vagary  was  attributed  to  the 
short  life  span  (one  year)  of  the  popula- 


tion studied,  which  yielded  females  of  a 

similar  size. 

Feeding 

The  overall  diet  of  E.  coosae,  consisted 
of  78  percent  Diptera  (Chironomidae  and 
Simuliidae),  12  percent  Crustacea  (Cope- 
poda  and  Cladocera),  3  percent  Ephemer- 
optera  (Baetidae  and  Siphlonuridae)  and 
5  percent  miscellaneous  items  (Acarina, 
Mollusca,  Nematoda,  Trichoptera,  and 
sand).  The  diet  of  various  size  classes  as 
well  as  the  seasonal  diet  of  combined  age 
and  size  classes  is  seen  in  Figure  5. 

Etheostoma  coosae  consumed  midge 
larvae  as  juveniles  and  expanded  their  diet 
as  adults  to  include  mayflies  and  caddis- 
flies.  Midge  larvae  decreased  whereas 
crusaceans  increased  in  importance  from 
spring  to  winter.  Mayflies,  caddisflies, 
and  molluscs  were  important  items  during 
summer  months. 


11-20mm 

N  =  6 


21-30nnm 
N  =  78 


31-40mi 
N  =  143 


41-50mi 
N  =  25 


jf,   Misc 


E|)h 


SPRING 
N=25 


SUMMER 
N=97 


WINTER 
N=34 


Figure  5.  Stomach  contents  of  Etheostoma  coosae  collected  in  Barbaree  Creek  by  size  class  of  darter  and 
season  collected.  Seasonal  analyses  include  all  size  and  age  classes.  Food  items  are  abbreviated  as  follows: 
(Crus)tacea,  (Dip)tera,  (Eph)emeroptera,  (Mis)ellaneous,  (Mol)lusca,  (Plec)optera,  and  (Tri)choptera.  N 
equals  sample  size. 


82 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


The  feeding  mode  of  darters  has  been 
reported  to  be  largely  selective  in  some 
species:  E.  fonticola  (Schenck  and 
Whiteside,  1977),  E.  nigrum  (Roberts  and 
Winn,  1962),  and  E.  radiosum  cyanorum 
(Scalel,  1972);  and  largely  opportunistic 
in  others:  E.  acuticeps  (Bryant,  1979),  E. 
blennioides  (Fahy,  1954),  and  E.  gracile 
(Braasch  and  Smith,  1967).  These  papers 
have  illustrated  that  within  the  genus 
Etheostoma  feeding  behaviors  are  quite 
variable  and  complex.  Based  on  the  liter- 
ature and  my  own  studies,  I  believe  that 
feeding  behavior  is  not  so  restrictive  but 
rather  lies  along  a  dynamic  continuum 
between  selectivity  and  opportunism. 
Species  will  adapt  to  prey  abundance  and 
type  assuming  the  most  energetically  re- 
warding feeding  response.  Prey  switching 
as  a  possible  behavioral  mechanism 
involved  in  feeding  is  supported  by  Ihe 
studies  of  Murdoch  et  al.  (1975)  on 
Poecilia  reticulatua  and  Roberts  and 
Winn  (1962)  on  the  role  of  visual  cues  in 
the  feeding  of  £.  nigrum. 

ACKNOWLEDGEMENTS 
1  wish  to  thank  Dr.  Herbert  Boschung, 
Dr.  Maurice  F.  Mettee,  and  John  Williams 
who  read  and  discussed  various  parts  of 
this  paper;  the  USDA  Forest  Service  for  a 
grant  to  Boschung  from  which  this  study 
was  funded;  and  finally  Irene  Thompson 
who  performed  all  typing  services  on  the 
manuscript. 


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BURR,  B.  and  L.  PAGE.  1978.  The  life  history  of 
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PAGE,  L.  1974.  The  life  history  of  the  spottail 
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.  1975.  The  life  history  of  the  stripetail  darter, 

Etheostoma  kennicoiti,  in  Big  Creek,  Illinois. 
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the  slabrock  darter,  Etheostoma  smithi,  in 
Ferguson  Creek,  Kentucky.  111.  Nat.  Hist.  Surv. 
Biol.  Notes  No.  99.  12  pp. 

PAGE,  L.  and  D.  SCHEMSKE.  1978.  The  effects  of 
interspecific  competition  on  the  distribution 
and  size  of  darters  of  the  subgenus  Catonotus 
(Percidae:  Etheostoma).  Copeia  1978:406-412. 

RICKER,  W.  1971.  Methods  for  assessment  of  fish 
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ROBERTS,  N.  and  H.  WINN.  1962.  Utilization  of 
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570. 

SCALET,  C.G.  1972.  Food  habits  of  the  orange- 
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No.  1  Etheostoma  Life  History  83 

ULTSCH,  C,   H.   BOSCHUNG,  and  M.   ROSS. 

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December  30,  1981 


84  Tulane  Studies  in  Zoology  and  Botany  Vol.  23 


THE  TAXONOMIC  RELATIONSHIP  BETWEEN  MALACLEMYSGKM ,  1844 
AND  GRAPTEMYS  AGASSIZ,  1857  (TESTUDINES:  EMYDIDAE) 


JAMES  L.  DOBIE 

Department  of  Zoology-Entomology 
Auburn  University,  Alabama  36849 


Abstract 

The  turtle  genus  Graptemys  is  a  distinctive  group 
clearly  separable  from  Malaclemys  on  the  basis  of 
external  and  osteological  features.  The  difference 
between  the  groups  indicate  that  the  degree  of 
genetic  relationship  is  no  closer  than  that  resulting 
from  their  both  having  presumably  arisen  from  a 
Pseudemys  -  like  stock  or  Malaclemys  from  a 
Graptemys  stock. 

INTRODUCTION 

Investigators  of  Malaclemys  and  Grapt- 
emys have  based  their  taxonomic  alloca- 
tions on  penial,  skull,  shell,  hind  limb  and 
pelvic  girdle  morphology  and  on  head 
patterns.  Osteological  comparisons,  when 
indicated,  were  usually  limited  to  the 
skull,  and  in  most  cases,  head  patterns 
were  used  to  distinguish  taxa.  The  degree 
of  evolutionary  conservatism  and  paral- 
leUsm  exhibited  by  turtles  argues  against 
the  use  of  external  characters  (e.g.,  head 
striping),  alone  in  determining  taxonomic 
and  phylogenetic  relationships.  Thus, 
both  osteological  and  surficial  features 
have  been  examined  in  this  study. 

HISTORICAL  REVIEW 

The  controversy  about  the  relationship 
between  Malaclemys  and  Graptemys  be- 
gan as  a  resuh  of  the  lumping  of  Grapt- 
emys with  Malaclemys  by  Boulenger 
(1889)  and  the  re-establishment  of  the 
genus  Graptemys  by  Baur  in  1890.  Since 


that  time,  W.P.  Hay  (1904)  and  O.P.  Hay 
(1908)  followed  Baur  in  recognizing  the 
two  genera,  as  did  Carr  in  1949.  Later, 
however,  Carr  (1952)  questioned  the 
validity  of  separating  the  two  genera  and 
McDowell  (1964),  without  presenting  sup- 
porting data,  lumped  Graptemys  with 
Malaclemys.  Zug  (1966,  1971),  on  the 
basis  of  similiar  penial,  pelvic  girdle,  and 
hind  limb  morphology  for  the  two  genera 
considered  them  congeneric,  and  Parsons 
(1960,  1968)  found  the  choanal  structures 
of  both  genera  to  be  so  variable  that  the 
evidence  did  not  particularly  support  or 
refute  the  congeneric  idea.  Several  other 
authors  (Ernst  and  Barbour,  1972; 
McKown,  1972;  Dundee,  1974;  Killebrew, 
1979;  Dobie  and  Jackson,  1979;  Pritch- 
ard,  1979;  Vogt,  1978,  1980)  have  not 
supported  the  synonymy  of  Graptemys 
with  Malaclemys;  they  evidently  must 
believe  that  sufficient  evidence  has  not 
been  presented  to  lump  the  two  genera 
together. 

The  purpose  of  this  study  is  to  clarify 
the  generic  status  of  Malaclemys  and 
Graptemys. 

MATERIALS  AND  METHODS 

Representatives  of  each  of  the  ten  ex- 
tant Graptemys  species  (Vogt,  1980)  and 
their  subspecies  and  individuals  of  several 
subspecies  of  the  monotypic  Malaclemys 
were  examined.  External  features,  includ- 


Editorial  Committee  for  this  Paper: 

Dr.   Eugene  S.    GAFFNEY,   Associate  Curator,   Department  of  Vertebrate 
Paleontology,  American  Museum  of  Natural  History,  New  York,  New  York  10024 

Dr.   John   J.    IVERSON,   Assistant   Professor  of  Biology,   Earlham  College, 
Richmond,  Indiana  47374 

85 


86 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


ng  scute  contracts,  plastral  patterns,  and 
striping  on  the  head  and  leg  were  analyzed 
in  juvenile  and  adult  turtles  of  both  sexes. 
Skull  and  shell  characters  were  analyzed 
on  large  sub-adult  and  adult  females. 
Skull  terminology  is  that  of  Gaffney  (1972 
a);  scute  and  bone  terminology  is  that 
used  by  Zangerl  (1969). 

The  method  used  to  elucidate  the  rela- 
tionship between  Malaclemys  and  Grapt- 
emys  and  to  other  North  American  emy- 
did  turtles  is  the  search  for  taxa  that  have 
shared  derived  characters.  This  method 
was  described  by  Hennig  (1966),  and  has 
been  used  by  others  (Gaffney,  1972  b, 
1975;  W.E.  Clark,  1978)  and  is  called 
phylogenetic  systematics  or  cladism. 

DIAGNOSTIC  CHARACTERISTICS 

The  diagnostic  characteristics  of  Grapt- 
emys,  Malaclemys  and  an  outgroup  com- 
parison of  those  genera  with  the  other 
North  American  emydid  genera  are  listed 
in  Table  1 .  Each  feature  is  also  designated 
as  either  ancestral  (primitive)  or  advanced 
(derived). 

SIGNIFICANCE  OF  DIAGNOSTIC 
CHARACTERISTICS 

The  number  (s)  in  a  bracket  refers  to 
the  number  of  the  diagnostic  features  in 
Table  1. 

SKULL  FEA  TURES 

(1)  Quadratojugal  -  maxilla  contact.  If 
the  absence  of  contact  between  these  two 
bones  represents  the  primitive  state,  then 
the  possession  of  the  derived  condition  in 
three  Graptemys  species  (in  one  pseudo- 
geographica  and  in  all  pulchra  and  bar- 
bourij,  in  M.  terrapin,  and  in  some 
Pseudemys  species  suggests  that  M. 
terrapin  could  have  been  derived  from 
one  of  these  Graptemys  or  Pseudemys 
species.  Graptemys  could  have  come  from 
any  group  lacking  contact  between  the 
two  bones. 

(2)  Spoon-shaped  symphysis  of  lower  jaw 


(Fig.  1).  The  flattened  spoon-shaped 
nature  of  the  symphyseal  part  of  the  lower 
jaw  apparently  is  a  derived  feature  in 
Graptemys.  The  absence  of  such  a  struc- 
ture in  Malaclemys  suggests  that  Grapt- 
emys was  not  ancestral  to  Malaclemys  and 
that  Malaclemys  may  have  arisen  from 
some  Pseudemys  species. 


Figure  1.  Shape  of  the  symphyseal  area  of  the 
lower  jaw  in  mature  females  of  (A)  Malaclemys  ter- 
rapin. (B)  Graptemys  pseudogeographica,  (C)  G. 
geographica,  (D)  G.  pulchra,  (E)  G.  barbouri,  (?)  G. 
caglei,  (G)  G.  versa,  (H)  G.  ouachitensis  sabinensis, 
(I)  G.  o.  ouachitensis,  and  (J)  G.  flavimaculata  (the 
shape  of  the  symphysis  is  the  same  for 
flavimaculata,  oculifera,  and  nigrinoda). 


No.  1 


Malaclemys-  Graptemys  Relationship 


87 


(3)  Bones  surrounding  the  foramen  pala- 
tinum  posterius  (Fig.  2).  The  bones  sur- 
rounding that  foramen  in  Terrapene  and 
in  the  species  of  the  Pseudemys  rubriven- 
tris  complex  are  the  same  as  Graptemys; 
the  other  species  of  Pseudemys  and  the 
other  N.A.  emydid  genera  are  like  Mala- 
clemys. Therefore,  Graptemys  and  Mala- 
clemys were  possibly  derived  from  differ- 
ent species  of  Pseudemys. 

(4)  The  absence  of  contact  between  the 
ophisthotic  and  pterygoid  due  to  the  in- 
volvement of  the  exoccipital.  If  the  condi- 
tion in  Malaclemys  and  Deirochelys  repre- 
sents a  derived  feature,  this  would  strong- 
ly suggest  that  Malaclemys  was  not  the 
ancestral  stock  from  which  Graptemys 
evolved.  It  could  also  indicate  that  a 
Graptemys,  Deirochelys,  or  any  other 
species  of  North  American  emydid  turtle 
could  have  been  ancestral  to  Malaclemys. 

(5)  The  lack  of  a  notch  in  the  premaxil- 
lary  bones.  The  lack  of  a  notch  in  those 
bones  in  Graptemys  and  the  presence  of  a 
notch  in  Malaclemys  and  the  rest  of  the 
N.A.  emydids,  precludes  determination 
of  the  possible  ancestor  for  Graptemys 
and  Malaclemys  based  on  this  feature. 

SHELL  FEA  TURES 

(6)  Flaring  of  carapace.  The  presence  of 
such  in  Graptemys  and  to  varying  degrees 
in  all  other  N.A.  emydids  except  Mala- 
clemys and  some  Terrapene,  may  indicate 
that  flaring  is  an  ancestral  feature.  If  so, 
the  upturning  of  the  carapace  in  the  last 
two  genera  would  be  a  derived  feature. 
This  implies  that  Graptemys  did  not  come 
from  a  Malaclemys  stock. 

(7)  Double  notching  of  some  peripherals. 
The  double  notching  of  some  of  the  per- 
ipherals is  found  only  in  Graptemys  and 
in  some  individuals  of  Pseudemys  scripta 
and  P.  concinna.  This  could  indicate  that 
Graptemys  was  not  ancestral  to  Mala- 
clemys and  that  a  Pseudemys  species  was 
ancestral  to  Graptemys. 

(8  and  9)  The  keel  and  its  associated 
bosses  (Fig.  3).  A  number  of  reports  have 
dealt  with  the  extent  and  development  of 
the  keel  in  Malaclemys.  The  last  vertebral 


scute  is  variable  with  respect  to  keel  devel- 
opment. Say  (1825)  reported  that  the  last 
vertebral  in  M.  terrapin  centrata  was  un- 
keeled;  Wied  (1865)  noted  that  all  of  the 
vertebrals  of  M.  t.  pileata  have  a  well 
developed  keel.  The  keel  in  Malaclemys  t. 
centrata  was  stated  by  W.P.  Hay  (1904) 
"to  be  rather  low  and  rounded,"  whereas 
it  was  "always  well  developed,"  in  M.  t. 
macrospilota.  A  keel  is  thus  not  always 
present  on  the  last  vertebral,  and  I  have 
not  observed  the  end  of  the  keel  (the  fifth 
boss  area)  to  extend  more  than  four-fifths 
the  length  of  the  last  vertebral  scute.  W.P. 
Hay's  (1904)  statement  about  the  keel  and 
bosses  of  M.  t.  littoralis  was:  "the  first 
vertebral  plate  is  raised  on  the  middle  line 
to  form  a  broad,  low  carina;  on  the 
second  plate  the  elevation  is  greater,  and 
stands  out  as  a  smooth  boss  .  .  .  ;  the 
elevation  on  the  third  plate  has  the  form 
of  a  hemispherical  button  with  a  well- 
marked  constriction  around  the  posterior 
half  of  the  base  .  .  .  ;  on  the  fourth  plate 
the  elevation  is  raised  into  a  knob-like 
protuberance  from  a  base  which  is  con- 
stricted all  around  .  .  .  ;  the  fifth 
vertebral  plate  is  flat  or  with  only  a  trace 
of  an  elevation."  Thus  Hay's  statement 
suggests  that  four  or  five  bosses  are 
present  on  the  keel  in  Malaclemys.  This  is 
not  always  the  case.  Auburn  University 
Museum  of  Paleontology  (AUMP)  speci- 
men 2179  has  only  three  bosses,  and  its 
shell  structures  are  normal. 

Concerning  the  total  number  of  bosses 
on  the  keel  in  Graptemys  pulchra,  Carr 
and  Coin  (1955)  said,  "the  dorsal  keel 
.  .  .  comprises  a  boss  on  each  of  the  first 
four  centrals,  .  .  .  weak  to  nearly  lacking 
on  the  first  and  completely  lacking  on  the 
fifth."  A  boss  on  the  fifth  central  (verte- 
bral) is  not  lacking  \n  pulchra.  Although  it 
is  not  prominent  in  G.  pulchra  or  in  any 
other  species  of  Graptemys,  a  terminal 
boss  can  be  detected  in  all  species.  Cagle 
(1954),  p.  182,  Fig.  11)  illustrated  a 
juvenile  G.  flavimaculata  that  had  five 
bosses  on  the  carapace.  I  have  never 
examined  any  specimen  of  Graptemys, 
including  G.  flavimaculata,  in  which  the 


88 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


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No.  1 


Malaclemys-  Graptemys  Relationship 


91 


Figure  2.  The  location  of  the  foramen  palatinum  posterius.  The  foramen  is  bounded  on  its  mediolateral 
and  outer  lateral  sides  by  the  palatine  in  Graptemys  (A)  pseudogeographica,  (B)  geographica,  (C)  pulchra, 
(D)  barbouri,  (E)  eaglet  and  versa,  (F)  o.  sabinensis,  (G)  o.  ouachitensis  and  (H)  nigrinoda,  oculifera,  and 
flavimaculala.  It  is  bounded  on  its  mediolateral  and  outer  sides  by  the  palatine  and  maxilla,  respectively,  in 
Malaclemys  terrapin  (I).  Palatine  (p).  Maxilla  (m).  Foramen  (0-  Vomer  (v).  Pterygoid  (pt). 


92 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


fifth  boss  was  located  in  the  position  il- 
lustrated by  Cagle;  the  fifth  boss  is  always 
at  the  posterior  end  of  the  last  vertebral 
scute.  The  similar  location  of  each  boss  in 
Graptemys  and  Malaclemys  indicates 
their  close  relationship. 
(10)  Amount  of  ventrolateral  extension  of 
the  nuchal  bone  and  the  costiform  process 
of  the  nuchal  bone.  Graptemys  normally 
lacks  a  costiform  process;  Malaclemys  has 
one.  Even  though  the  nuchal  of  Grapt- 
emys is  as  wide  as  the  same  bone  in  Mala- 
clemys, the  distance  the  nuchal  extends 
ventrolaterally  is  less  in  Graptemys  than 
in  Malaclemys.  Therefore,  the  degree  of 
such  extension  must  not  be  solely  a  func- 
tion of  the  width  of  the  nuchal  bone.  This 


Nu 


seems  to  be  the  case  since  the  distal  width 
of  the  first  peripheral  is  proportionately 
greater  in  Graptemys  than  in  Malaclemys. 
Therefore,  the  presence  of  a  narrower 
first  peripheral  and  a  costiform  process  in 
Malaclemys  results  in  a  greater  ventro- 
lateral extension  of  the  nuchal  in  that 
genus  than  in  Graptemys. 

The  other  North  American  emydids 
that  have  a  costiform  process  are  Pseud- 
emys,  Terrapene,  some  Clemmys  and 
Deirochelys,  and  the  latter  genus  is  the 
only  group  that  has  a  ventrolateral  exten- 
sion similar  to  that  of  Malaclemys.  I  think 
it  unlikely  that  Deirochelys  was  ancestral 
to  Malaclemys;  therefore,  perhaps  some 
Pseudemys  turtle  was  the  stock  from 
which  Malaclemys  arose.  The  ancestral 
stock  for  Graptemys  can  not  be  determi- 
ned with  respect  to  this  feature. 
(11  and  12)  The  notching  of  the  postero- 
lateral borders  of  the  nuchal  bone  and  the 
anterior  border  of  the  costal  bone  (Figs.  4 
and  5).  The  presence  of  such  notching  in 
Graptemys,  Terrapene  and  in  most 
Clemmys  (14  of  16),  Pseudemys  (29  of 
31),  and  Chrysemys  (15  of  20),  and  not  in 
Malaclemys  (except  in  one  specimen), 
Emydoidea,  and  most  Deirochelys  sug- 
gests that  Malaclemys  was  not  ancestral  to 


Figure  3.  The  location  of  the  bosses  in  Graptemys 
(A)  pulchra,  (B)  nigrinoda,  and  (C)  Malaclemys 
lerrapin  and  the  contact  of  the  eighth  costal  with  the 
seventh  neural  in  some  G.  pulchra  due  to  the  loss  of 
the  eighth  neural  bone.  The  normal  contact  is 
between  eighth  costal  and  eighth  neural  in  Grapt- 
emys and  eighth  costal  and  seventh  and  eight  neurals 
in  Malaclemys.  Nuchal  bone  (Nu).  Bosses  )B  1-5). 

Neural  bones  (N  1-8).  Suprapygal  bones  (S  1-2).  Py- 
gal  bone  (P).  Costal  bones  (C  1-8). 


Figure  4.  Dorsal  view  of  the  nuchal  bone  in 
Graptemys  (A)  pseudogeographica,  (B)  pulchra  and 
(C  and  D)  Malaclemys  terrapin.  Arrows  indicate 
notches.  The  position  of  the  anteromedial  edge  of 
the  first  pleural  scute  and  the  anterolateral  borders 
of  the  first  vertebral  scute  are  not  on  the  nuchal  bone 
in  some  Malaclemys  (D). 


No.  I 


Malaclemys-  Graptemys  Relationship 


93 


Graptemys  if  the  absence  of  notching  is  a 
derived  feature.  However,  Graptemys 
could  have  given  rise  to  Malaclemys,  as 
could  have  Clemmys,  Chrysemys,  Pseud- 
emys,  Terrapene,  Emydoidea,  and  Deiro- 
chelys.  Emydoidea  and  Deirochelys  pre- 
sumably would  be  the  best  candidates 
from  which  to  derive  Malaclemys  if  rela- 
tionships are  based  on  the  presence  of 
shared  derived  features.  In  spite  of  the 
presence  of  a  shared  derived  feature 
between  those  genera  and  Malaclemys,  I 
do  not  believe  that  either  one  is  a  good 
candidate  for  being  the  progenitor  of 
Malaclemys.  Therefore,  Graptemys, 
Pseudemys,  and  Chrysemys  are 
considered  to  be  more  likely  candidates. 
(13  and  14)  The  amount  of  pleural  scute 
overlap  on  the  nuchal  bone  and  first 
vertebral  scute  -  nuchal  bone  relation- 
ships. A  great  deal  of  pleural  scute  over- 
lap exists  in  Graptemys,  Pseudemys,  and 


Figure  5.  Dorsal  view  of  the  first  left  costal  bone  in 
Graptemys  (A)  pseudogeographica,  (B)  pulchra  and 
(C  and  D)  Malaclemys  terrapin.  That  part  of  the  an- 
terior borde:  of  the  costal  bone  that  would  adjoin 
the  nuchal  generally  is  straight  and  unnotched  in 
Malaclemys  as  in  (D).  Arrows  indicate  notches. 


in  some  Terrapene  and  the  pleural  scute 
always  contacts  the  margin  of  the  first 
vertebral  scute  on  the  nuchal  bone  in  the 
first  two  of  the  the  three  above  (Dobie 
and  Jackson,  1979).  Malaclemys  resem- 
bles rriost  Chrysemys  and  some  Terra- 
pene, Clemmys,  and  Deirochelys  in  that 
there  is  little  overlap  of  the  pleural  scute 
on  the  nuchal  and  the  pleural  scute  does 
not  always  contact  the  first  vertebral  scute 
on  the  nuchal  bone  (Dobie  and  Jackson, 
1979). 

Malaclemys  terrapin  could  have 
evolved  from  Chrystemys  in  which  the 
extent  of  pleural  scute  overlap  was  mini- 
mal and  the  margin  of  the  first  vertebral 
scute  did  not  always  meet  the  pleural  scute 
on  the  nuchal  bone.  If  M.  terrapin  evolved 
from  any  species  of  Graptemys  or  Pseud- 
emys that  had  a  large  amount  of  pleural 
scute  overlap  and  contact  between  the  two 
scutes  on  the  nuchal  bone,  then  presum- 
ably a  reduction  in  the  amount  of  pleural 
scute  overlap  must  have  occurred.  Grapt- 
emys could  have  arisen  from  a  Pseudemys 
stock. 

(15  and  16)  Amount  of  nuchal  scute  over- 
lap and  underlap  and  the  width-length 
relationships  of  the  underlap  part  of  the 
nuchal  scute  (Figs.  6  and  7).  The  amount 
of  nuchal  scute  overlap  is  small  in  Mala- 
clemys, in  some  Terrapene,  and  in  all 
extant  species  of  Graptemys,  except  G. 
geographica  (Dobie  and  Jackson,  1979). 
Both  Malaclemys  and  Graptemys  have 
smaller  amounts  of  nuchal  scute  underlap 
than  any  other  North  American  emydid 
turtle,  and  the  distal  width  of  the  under- 
lap part  of  the  nuchal  scute  is  broader 
than  its  length  in  both  of  those  genera  and 
in  some  Pseudemys  and  Terrapene  (Dobie 
and  Jackson,  1979).  Based  on  these  fea- 
tures, Malaclemys  would  seem  to  be  more 
closely  related  to  Graptemys  than  to  any 
other  extant  North  American  emydid 
genus. 

(17)  Contact  of  the  eighth  costal. bone 
with  the  seventh  and  eighth  neurals  (Fig. 
3).  The  presence  of  such  contacts  in  Mala- 
clemys and  the  contact  of  the  eighth  costal 
with  only  the  eighth  neural  in  Graptemys 


94 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


(except  for  a  single  population  of  G. 
pulchra)  and  in  all  other  North  American 
emydid  genera  except  Terrapene  (the 
eighth  neural  is  absent  in  some  Terra- 
pene }  indicates  that  contact  with  the 
seventh  neural  is  a  derived  character.  The 
stock  from  which  Malaclemys  was  derived 
presumably  could  have  been  any  genus  of 
North  American  emydid  turtles;  Grapt- 
emys  could  have  come  from  Pseudemys 
or  from  any  other  North  American 
emydid  genus  except  Malaclemys. 
(18)  Lateral  ridges  on  undersides  of  first 
and  fifth  costals  (Fig.  8).  The  lateral 
ridges  extending  toward  the  midline  of  the 
carapace  from  the  anterior  and  posterior 
ends  of  the  bridge  are  well  developed  in 
Graptemys  in  constrast  to  those  of  Mala- 
clemys and  the  rest  of  the  North  Ameri- 
can emydid  genera.  The  functional  sig- 


nificance of  those  ridges  is  not  known  but 
they  may  serve  as  supportive  units  for  the 
carapace.  Malaclemys  and  Graptemys 
presumably  could  have  been  derived  from 
any  one  of  those  genera. 

(19)  Distal  widths  of  the  three  widest 
costal  bones.  An  attempt  to  indicate  the 
degree  of  relationships  of  Malaclemys  to 
any  other  emydid  genus  on  the  basis  of 
this  character  would  be  impractical 
because  of  the  extremely  variable  nature 
of  the  widths  of  the  costal  bones.  The 
fairly  consistent  widths  in  the  species  of 
Graptemys  does  indicate  that  they  are 
closely  related. 

(20)  Sculpturing  on  the  carapace.  The 
sculpturing  on  the  carapacial  bones  in 
Graptemys  is  similar  to  that  of  some 
species  of  Pseudemys  (P.  floridana  and  P. 
concinna)  although  the  degree  of  sculp- 


26|- 

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10 


•  Srapttmyt   (38) 

o  Maiaeltmyt    (II) 

■  CHrytemyt    ( Psfudfmyt 

and    rreeH»myt)    (36) 

o  Chrystmyt  picta     (17) 

A  D»iroclt»ly$   (16) 

A  Emydotdta    (H) 

+  r»rrap»n»   (23) 

«  Clfmmys    (!•) 

»  Fossil  Malaeltmya    (i) 


Distal  Width  of  Nuchol  Scute  Overlap  (mm) 


Figure  6.     Length  of  nuchal  scute  overlaps  versus  distal  width  of  nuchal  scute  overlap  in  various  emydines 
including  Graptemys  (35)  and  Malaclemys  (11). 


No.  1 


Malaclemys-  Graptemys  Relationship 


95 


turing  in  Graptemys  is  generally  less  than 
in  any  species  of  Pseudemys  and  more 
than  that  of  Chrysemys.  The  type  of 
concentric  sculpturing  in  Malaclemys  is 
unique  and  represents  a  derived  feature 
(the  species  of  Terrapene,  some  Antillean 
Pseudemys,  and  Clemmys  insculpta  also 
have  concentric  sculpturing  (Zangerl, 
1969;  Dobie  and  Jackson,  1979)  but  the 
sculpturing  patterns  in  the  species  of 
Terrapene,  Antillean  Pseudemys,  and  in 
C.  inscultpa  are  not  the  same  as  that 
demonstrated  by  Malaclemys.  Graptemys 
may  have  arisen  from  Pseudemys; 
Malaclemys  from  any  one  of  these  genera 
including  Graptemys. 
(21)  Carapacial  pattern.  The  patterns  on 
the  carapace  of  the  various  Graptemys 
justify  the  name,  "map  turtle".  Those 
patterns,  although  more  similar  to  those 
patterns  found  in  other  North  American 
emydids,  except  Clemmys  guttata,  are 
distinctive  and  were  probably  modified 
from  a  less  elaborate  carapacial  pattern. 
The  lack  of  similarity  of  the  carapacial 
patterns  of  Graptemys  and  Malaclemys 
could  mean  that  the  patterns  of  both  were 
independently  derived  from  different  an- 


cestors or  that  they  came  from  the  same 
ancestor  that  had  a  less  elaborate  pattern. 
(22)  Bridge  width  (Fig.  9).  The  width  of 
the  bridge  in  Graptemys  resembles  that  of 
most  aquatic  emydids.  The  relatively 
narrow  bridge  in  M.  terrapin  is  distinc- 
tive, presumably  derived,  and  perhaps  is 
an  adaptation  for  increasing  the  animal's 
ability  for  bottom  walking  in  that  a 
narrow  bridge  could  allow  the  limbs  to  be 
advanced  to  a  greater  degree  anteriorly 
than  in  a  turtle  having  a  wide  bridge. 
Malaclemys  could  have  come  from  any 
one  of  several  different  genera  on  the 
basis  of  this  feature. 

(23  and  24)  The  separation  of  the  seventh 
marginal  scute  from  the  abdominal  scute 
by  the  inguinal  scute  and  the  sizes  of  the 
inguinal  and  axillary  scutes.  The  separa- 
tion of  the  two  scutes  by  the  inguinal  scute 
in  Graptemys  indicates  that  the  size  of  the 
inguinal  scute  is  about  the  same  size  as 
that  found  in  most  other  North  American 
emydids.  The  contact  between  the  abdom- 
inal and  seventh  marginal  scutes  in  Mala- 
clemys is  due  to  the  small  size  of  the 
inguinal  scute  or  the  absence  of  that  scute. 
The  condition  in  Malaclemys  is  probably 
derived. 


-     13 

E 

i   12 


O  " 


•  Graptemys  (55) 
o    Malaclemys  ( 1 1 1 

■   Chrysemys  (Pseudemys 

ond  Trachemys)  Ci^) 
a   Chrysemys  picta  ( 1 7 ) 
A  Oeirochelys  (16) 
A  Emydoidea  (il) 
•♦■   Terrapene  (24) 

*  Clemmys  (I I) 

»   Fossil  Malaclemys  (I) 


5  6  7  8  9  10  II  12 

DIstol  Width  of  Nuchol  Scute  Underlop  (mm) 


Figure  7.     Length  of  nuchal  scute  un(Jerlap  versus  distal  wi(Jth  of  nuchal  scute  uncjerlap  in  various  emydines 
including  Graptemys  (35)  and  Malaclemys  (11). 


96 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Figure  8.  Lateral  "extensions  of  ridges  on  the 
ventral  sides  of  the  first  and  fifth  costal  bones  in  (A) 
Graptemys  nigrinoda  and  (B)  Malaclemys  terrapin. 
The  arrows  indicate  the  ridges.  Nuchal  bone  (Nu). 
Costal  bone  (C  1).  Costal  bone  (C  5). 


The  size  of  the  axillary  scute  in  Grap- 
temys is  like  that  of  most  other  emydids. 
It  is  either  absent  or  very  small  in  Mala- 
clemys. The  reduction  in  the  size  or  loss  of 
both  the  axillary  and  inguinal  scutes  is 
perhaps  a  result  of  the  decrease  in  bridge 
width.  Based  on  these  features,  Grapt- 
emys and  Pseudemys  are  more  similar 
than  either  is  to  Malaclemys. 
(25  and  26)  Plastral  formulae  and  the 
lengjh  of  the  abdominal  plastral  scute. 
The  two  genera  are  more  similar  to  each 
other  in  these  two  features  than  either  is 
to  any  other  North  America  emydid 
genus;  they  would  thus  appear  to  be  close- 
ly related. 

(27)  Plastral  patterns.  The  ancestral  plas- 
tral pattern  of  Graptemys  was  probably 
ornate  because  to  varying  degrees  ornate 
plastral  patterns  appear  in  all  species  of 
Graptemys  except  G.  barbouri.  The 
plastral  patterns  in  Malaclemys,  although 
ornate,  do  not  resemble  the  pattern  of  any 
Graptemys  species   except   for   a  single 


specimen  of  G.  nigrinoda.  The  ornate 
plastral  patterns  of  both  were  probably 
derived  from  different  ancestral  stocks. 

HEAD,  NECK  AND  LIMB  STRIPING 

(28)  Head,  neck  and  limbs  striped.  The 
striping  of  such  units  is  a  typical  emydid 
condition  and  Graptemys  is  no  exception. 
According  to  Wood  (1977),  Malaclemys  is 
striped  although  I  and  evidently  Pritchard 
(1979)  have  never  seen  a  striped  individual 
and  Ernst  and  Barbour  (1972)  use  the 
absence  of  ^triping  in  Malaclemys  as  a 
feature  in  their  key  to  U.S.  turtles.  If 
striping  does  occur  in  Malaclemys,  it  must 
be  a  rare  condition.  The  absence  of  strip- 
ing in  Malaclemys  is  a  derived  feature. 
Malaclemys  could  have  been  derived  from 
Graptemys  or  from  any  other  North 
American  emydid  genus. 

DIPLOID  CHROMOSOME  NUMBER 

(29)  Chromosome  count.  Because  all 
emydines  presumably  have  50  chromo- 
somes (Killebrew,  1977),  Graptemys  and 
Malaclemys  could  have  been  derived  from 
each  other,  from  any  one  of  several  dif- 
ferent groups,  or  perhaps  from  a  bata- 
gurine  if  in  fact  the  50  chromosome 
number  of  emydines  is  a  derivation  of  the 
52  chromosome  number  of  the  batagur- 
ines. 

DISCUSSION  AND  CONCLUSION 

All  indications  are  that  Graptemys 
represents  a  distinct  group  of  closely  re- 
lated turtles.  Malaclemys  is  undoubtedly 
more  closely  related  to  Graptemys  than  it 
is  to  any  other  extant  genus,  as  would  be 
evidenced  by  (1)  the  pterygoid  forming  a 
suture  with  the  exoccipital  except  in  some 
species  of  Graptemys  (G.  nigrinoda  for 
example)  and  in  some  individuals  of  M. 
terrapin,  (2)  similarities  in  penial,  pelvic 
girdle  and  hind  limb  morphology,  (3)  sim- 
ilarity in  carapacial  seam  contacts 
(Tinkle,  1962),  (4)  similarity  in  the 
amount  of  nuchal  scute  underlap,  and  (5) 
similarity   in   the   width-length   relation- 


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97 


ships  of  the  underlap  of  the  nuchal  scute. 
In  addition,  the  plastral  scute  formulae 
are  the  same  for  the  two  genera  as  are, 
generally,  the  locations  of  the  bosses  on 
the  carapace. 

The  Oligocene  species  of  Graptemys, 
G.  inornata  (Loomis,  1904)  and  G.  cordi- 
fera  (J.  Clark,  1937)  do  not  have  shell 
characteristics  that  indicate  a  close  rela- 
tionship with  Malaclemys.  No  other 
remains  of  G.  inornata  and  G.  cordifera 
are  known.  No  fossils  intermediate  be- 
tween Graptemys  and  Malaclemys  are 
known,  and  only  recently  were  fossil  re- 
mains for  M.  terrapin  discovered  (Pleisto- 
cene age:  South  Carolina,  [Dobie  and 
Jackson,    1979]  ).    Examination    of    an 


Eocene  specimen  (South  Dakota  School 
of  Mines  and  Technology,  SDSM&T, 
59187)  identified  as  Graptemys  by  Bjork 
(1967),  reveals  that  it  is  not  Graptemys  or 
Malaclemys  because  it  lacks,  among  other 
things,  a  keel  and  bosses.  The  absence  of 
the  uniform  fine  granular  tubercles  on  the 
external  surface  of  the  carapace  of  the 
Eocene  fossil  prevents  its  inclusion  within 
Compsemys  (a  baenid  turtle,  Gaffney, 
1972b)  and  the  absence  of  a  keel  and 
rugosities  rules  out  its  inclusion  within 
any  genus  of  North  American  emydids 
except  Chrysemys  (some  Chrysemys  do 
have  a  slight  keel).  On  the  basis  of  the 
absence  of  the  latter  two  features  it  is  like 
Chrysemys  picta.  However,  it  cannot  be 


85 


80 


75 


70 


^65 

E 


60 


55 


50 


45 


40 


35 


30 


100 


•  Gfoptemys 
o  Moloclemys 


120 


140 


160  180 

PlOStron  Length  (miTi) 


200 


220 


Figure  9.     Relative  bridge  width  in  Graptemys  and  Malaclemys.  The  solid  line  depicts  the  separation  of  the 
two  genera. 


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Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


included  within  Chrysemys  picta  as  the 
length  of  the  sixth  neural  in  C  picta  is 
about  twice  as  long  as  that  of  the  fossil 
and  the  posterior  width  of  the  first 
suprapygal  of  the  fossil  is  about  twice  the 
width  of  the  same  bone  in  C.  picta.  The 
neural  bones  of  the  fossil  are  narrow  as 
compared  to  those  of  Deirochelys  carri, 
D.  reticularia,  Emydoidea  blandingi, 
Malaclemys  terrapin,  Clemmys  guttata, 
and  C.  insculpta  and  this  rules  out  the 
inclusion  of  the  fossil  into  any  of  those 
genera. 

The  features  possessed  by  the  Eocene 
fossil  do  not  fit  those  of  Graptemys, 
Malaclemys  or  any  extant  North  Ameri- 
ican  emydid  genus,  thus,  it  may  be  a  new 
taxon. 

Although  Graptemys  and  Malaclemys 
have  several  characteristics  in  common 
with  some  of  the  species  of  the  Eocene 
emydid  fossil  turtles  assigned  to  the  genus 
Echmatemys  (Table  2),  I  do  not  believe 
that  either  one  of  the  two  taxa  nor  any 
other  new  world  emydine  genus  came 
from  Echmatemys.  O.P.  Hay  (1908)  and 
Weaver  and  Rose  (1967)  proposed  that 
Chrysemys  came  from  Echmatemys  and 
Hay  (1908)  also  believed  that  Echma- 
temys was  the  ancestral  stock  for  most 
other  North  American  emydine  genera.  I 
reject  the  ancestral  status  of  Echmatemys 
because  to  me  many  if  not  most  of  the 
species  of  Echmatemys  appear  to  be 
members  of  Rhinoclemmys  (e.g., 
McDowell,  1964,  believed  that  E.  pusilla 
belonged  in  the  Neotropical  batagurine 
genus  Rhinoclemmys)  and  because  most 
of  the  characters  used  to  indicate  relation- 
ships between  Echmatemys  and  Chrys- 
emys (in  the  sense  of  Weaver  and  Rose, 
1967)  were  primitive  characters  and  such 
can  never  be  used  to  determine  relation- 
ships. The  Graptemys  line  may  have 
arisen  from  some  Eocene  pve-Pseudetnys 
of  Pseudemys  stock;  Malaclemys  may  be 
an  additional  derivation  of  a  Pseudemys 
stock  or  of  a  Graptemys  stock,  but  its 
origin  was  probably  somewhat  later  in  the 
Tertiary  (post-Miocene  or  later). 

Loveridge  and  Williams  (1957)  believed 


that  Graptemys  may  have  arisen  from  a 
Pseudemys  stock,  as  did  McDowell 
(1964),  Ernst  (1974),  and  Pritchard 
(1979),  and  that  the  ancestral  Malaclemys 
was  close  to  a  Graptemys  stock.  The 
former  is  in  disagreement  with  O.P. 
Hay's  (1908)  conclusion  that  Graptemys 
was  from  Malaclemys.  Wood  (1977)  also 
considered  Graptemys  a  Malaclemys 
derivative,  and  according  to  him,  "most 
or  all  of  these  species  evolved  indepen- 
dently and  perhaps  also  at  different  times 
during  the  latter  part  of  the  Pleistocene 
from  Malaclemys  rather  than  giving  rise 
to  one  another."  Assuming  that  each 
species  of  Graptemys  was  independently 
derived  from  M.  terrapin  as  Wood 
believes,  then  each  feature  common  to 
two  or  more  Graptemys  but  absent  in  M. 
terrapin  must  exemplify  convergence.  A 
total  of  24  features,  at  least  10  of  which 
appear  to  be  derived,  are  shared  by  all 
Graptemys,  only  six  of  these  feature,  at 
least  three  of  which  appear  to  be  derived, 
are  possessed  by  Malaclemys.  It  is  highly 
unlikely  that  the  remaining  18  features 
(seven  derived  and  11  ancestral)  would 
have  arisen  independently  in  all  Grapt- 
emys species. 

Because  of  the  number  of  features  held 
in  common  by  the  species  of  Graptemys 
and  because  it  is  obvious  to  me  and  to 
other  individuals  (Cagle,  1952,  1953a, 
1953b,  1954;  McKown,  1972;  Dundee, 
1974;  Killebrew,  1977;  Vogt,  1978,  1980) 
that  there  are  closely  related  complexes  of 
Graptemys  i\xri\Q^,  e.g.,  G.  nigrinoda,  G. 
flavimaculata,  and  G.  oculifera;  G. 
pulchra  and  G.  barbouri;  G.  pseudogeo- 
graphica,  G.  ouachitensis,  G.  versa,  and 
G.  caglei,  (G.  geographica  belongs  in  a 
group  by  itself),  I  conclude  that  the 
various  species  of  the  Graptemys  turtles 
were  derived  from  other  species  of  Grapt- 
emys. (The  species  of  Graptemys  are  thus 
more  closely  related  to  each  other  than 
any  one  species  is  to  M.  terrapin.) 

Wood  (1977)  apparently  was  unaware 
that  there  are  two  Oligocene  fossil  species 
of  Graptemys.  If  the  fossils  are  correctly 
assigned,  the  various  species  of  Grapt- 


No.  1 


Malaclemys-  Graptemys  Relationship 


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100 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


emys  obviously  could  not  have  been 
derived  independently  from  M.  terrapin 
during  the  Pleistocene. 

Adult  female  Malaclemys  terrapin  and 
adult  females  of  some  species  of  Grapt- 
emys  (pseudogeographica,  pulchra,  bar- 
bouri  and  geographica)  resemble  one 
another  closely  in  general  skull  shape.  The 
resemblance  of  M.  terrapin  to  those 
Graptemys  species  is  probably  not  due  to 
common  ancestry  but  rather  to  the  devel- 
opment by  each  species  of  similar  kinds  of 
anatomical  features  (e.g.,  broad  heads)  as 
adaptations  for  feeding  on  similar  kinds 
of  food  items  (mussels.)  Graptemys 
pulchra,  barbouri,  and  geographica  are 
also  farther  from  the  base  of  the  Grapt- 
emys phylogenetic  tree  than  is  G.  pseudo- 
geographica (a  species  which  is  presumed 
to  represent  more  nearly  the  ancestral-like 
stock)  and  both  G.  geographica  and  G. 
barbouri  appear  to  be  highly  specialized, 
derived  terminal  end  forms  with  respect  to 
skull  features.  None  of  those  species 
appears  to  be  closely  related  to  Malaclem- 
ys terrapin  even  though  all  have  broad 
heads. 

Mature  females  of  some  of  the  species 
of  Graptemys,  G.  nigrinoda,  G.  oculifera, 
G.  flavimaculata,  G.  versa,  G.  caglei,  G. 
ouachitensis  and  some  G.  pseudogeo- 
graphica, have  narrow  alveolar  surfaces. 
The  genus  Graptemys  cannot  be  differ- 
entiated, therefore,  from  Malaclemys  on 
the  basis  of  wide  alveolar  surfaces,  as 
O.P.  Hay  (1908)  contended. 

The  evidence  is  clearly  against  the 
lumping  of  Graptemys  and  Malaclemys. 
A  subsequent  paper  will  clarify  the  phyl- 
ogenetic relationships  of  the  Graptemys 
turtles. 

Acknowledgments 

1  am  grateful  to  Drs.  Robert  Mount  and 
George  Folkerts  for  their  advice  on 
various  aspects  of  this  study.  Several 
museums  and  one  individual  loaned  me 
specimens  and  Robert  Mount,  John 
Pritchett  and  Lacy  Hyche  reviewed  this 
manuscript.  Theresa  Rodriguez  and  Dr. 
Jeanne  Stuart  did  most  of  the  drawings. 


SPECIMENS  EXAMINED 

Chrvsemys  picla:  (74)  (AUM  426,  605,  749,  829, 
1170,  1553,  1915,  2062,  3827,  3872-73,  3875-76, 
3884-85,  3999,  5669,  5885,  7072,  9514,  9747,  10091, 
10126,  12587,  12589,  13616,  14133-34,  16231, 
17366-67,  17871-72,  18033-34,  18218,  18812-14, 
23478,  24109,  25088);  (AUMP  132,  1713-23,  1965, 
1967,  1983,  1985,  1990,  2117,  2171-76,  2318-20, 
2351-54,  2405). 

Clemmys  guttata:  (9)  (AUM  21554,  22433,  26741, 
three  classroom  specimens);  (AUMP  308,  2251); 
(UF/FSM  41018). 

C  insculpta:  (5)  (AUM  29257);  (AUMP  279); 
(UF/FSM  19016,  41525-26). 

C.  marmorata:  (9)  (AUMP  2260-62,  2264-66, 
2310-11);  (UF/FSM  41523). 

C.  muhlenbergi:  (1)  (UF/FSM  14116). 

Deirochelys  reticularia:  (44)  (AUM  1705,  1733, 
3378,  3898,  8747-48,  9320,  10090,  10109,  10152, 
11564,  12394,  13495,  15791,  18236,  18484,  18999, 
19729,  22706,  22998,  23001);  (AUMP  125-26,  897, 
935,  1924,  2315,  2910);  (UF/FSM  T736,  6530,  7744, 
14192,  14244-48,  30348,  34880,  35026,  38433,  40824, 
41524,  41533). 

Emydoidea  blandingi:  (17)  (AUMP  1724-26, 
1959,  1962,  1971,  2014-15,  2017,  2115,  2117,  2119, 
2252-54,  2417-18). 

Graptemys  barbouri:  (35)  (AUM  3380-81,  5956, 
6238,  6326-27,  6329,  6388,  6621,  8793,  8966, 
9470-71,  9500,  9548,  9659,  10101,  10104-05,  10276, 
11231,  12694-95,  13653-54,  14278,  21606,  22662); 
(AUMP  297,  325,  328-29,  931,  1733,  2357). 

G.  caglei:  (10)  (TNHC)  36066,  36071,  36084, 
36088,  36093,  36097,  36103,  36621,  36627-28). 

G.  flavimaculata:  (48)  (AUM  5941 ,  5968-74,  6147, 
6387,  8792,  8941-43,  8982-83,  9238-31,  9348, 
9492-95,  9538-40,  9542-46,  10150-51,  10294,  10296- 
98,  13660-61,  23664);  (AUMP  925,  940,  998-99, 
2129,  2247). 

G.  geographica:  (31)  (AUM  5976-77,  6622,  9319, 
9446-47,  10858,  11805,  11830,  12410-18,  12240-41, 
13002,  21613,  22910,  23111,  23242,  29574);  (AUMP 
300,  909,  1940,  2355);  (NLSC  622). 

G.  nigrinoda:  (33)  (AUM  5665,  5939,  5942,  5964, 
5983,  5989,  8948,  8968,  8970,  9233,  9235,  9237, 
9261-62,  9268,  10127,  10143-44,  10149,  10292, 
10301,  12562,  12575,  12630,  12635,  21553, 
22988-89);  (AUMP  927,  1730,  2255-56,  2419). 

G.  oculifera:  (23)  (AUM  5951-53,  5979,  9333, 
14289,  23665-69,  25136-39);  (AUMP  304,  2125-28, 
2215-16,  2248). 

G.  ouachitensis  ouachitensis:  (27)  (AUM  9136-38, 
25983-84,  25988,  26431-34,  26648);  (AUMP  278, 
309,  1738,  1997,  2131-32,  2136,  2200-04,  2273-75): 
(NLSC  9383). 

G.  ouachitensis  sabinensis:  (32)  (AUM  24019, 
24022-23,  24239-46,  24253-55,  25129-35);  (AUMP 
2121-24,  2244-46);  (NLSC  10137-39,  10142). 

G.  pseudogeographica  pseudogeographica:  (24) 
(AUM  25985,  27090,  27101,  27113),  (AUMP  2905, 
2902,   2277-84);   (SUSD    1520,   2855,   2860,   2862, 


No.  1 


Malaclemys-  Graptemys  Relationship 


101 


2880-83,  two  uncatalogued  specimens). 

G.  pseudogeographica  kohni:  (81)  (AUM  6843, 
20715,  23985,  23989,  23991-97,  24020,  24191, 
24224-25,  24247-52,  24259-60,  24263,  25989,  26385, 
26401-02,  26406,  26422-25,  27093-98);  (AUMP 
305-08,  326-27,  2118,  2133-35,  2161-66,  2185-88, 
2191-99,  2221,  2267-72,  2276,  2402,  2901);  (KU 
1183);  (NLSC2304,  5263). 

G.  pulchra:  (37)  (AUM  4997,  5000-01,  5004-06, 
5597,  5742,  5961,  6302,  6311,  9467-69,  9532,  9535, 
12556,  19898,  23482,  25140-44,  25977);  (AUMP  301, 
443,  926,  930,  936,  943-44,  989-91,  1000,  1960). 

G.  versa:  (14)  (AUM  16653,  22816,  23984,  24202, 
24222,  26030-34,  29302);  (AUMP  924,  2130  2137). 

Malaclemys  terrapin:  (23)  (AUM  8839,  14277,  a 
classroom  specimen);  (AUMP  706,  932,  954,  963, 
1732,  1734-37,  1956,  1980,  2157-58,  2179,  2403); 
(TU  15194,  .2,  15195.1);  (UF/FSM  22849a-49b). 

Pseudemys  alabamensis:  (41)  (AUM  4840,  9346, 
9957,  10072,  11598-99,  11601-02,  11608,  11813-14, 
12580,  12591,  16870-71,  17032-33,  19362,  26998, 
27003-05,  27007,  27009-10,  27018,  27020,  27023); 
(AUMP  277,  298,  938,  1706,  1710,  1906,  2285,  2356, 
2360-62);  (USA  1501-02). 

P.  concinna:  (142)  (AUM  4560,  5901,  5994,  7432, 
7567,  8918,  10140,  10147,  10396,  11294,  12650, 
13553,  13639,  13743,  16906,  17139,  18483,  18975, 
19140,  21802-05,  22825,  23248,  24201,  24208, 
24214-16,  24223,  24227-28,  24280-81,  25126-28, 
26413,  26416,  29298-01);  (AUMP  17,  284,  288,  290, 
311,  318-19,  693-94,  697,  881,  900-01,  911-12, 
917-19,  933-34,  950,  1707-09,  1904-05,  1941,  1976, 
1989,  1993,  2000,  2148,  2156,  2167-69,  2181-84, 
2189-90,  2221  2286-90,  2292-94,  2316  2410-12); 
(FMNH  55646,  55649-52);  (KU  33526);  (SFA  2769, 
2803,  2858,  2989,  3460);  (TCWC  13735,  13965-67, 
42345);  (TNHC  536-37);  (TU1637,  3605-06,  11940, 
13464,  14414,  14421-22,  .1-.3,  .9-. 10,  14441,  .2-. 3, 
.10,  14451,  .2-.3,  14506.1,  14541,  16030);  (UNM465, 
30345). 

P.  floridana:  (53)  (AUM  1670,  7672,  8976,  9505, 
9563,  10102,  10290-91,  10725-29,  11596,  12428, 
12430,  12602,  13834,  17133-34,  19000,  19927-29, 
21609,  21831,  22658,  23201,  23490,  23703,  27706, 
27945);  (AUMP  289,  440-42,  447-48,  700,  1703, 
1712,  1727-29,  1902,  1948,  1963,  1981,  1998,  2249, 
2291  2309,  2404). 

P.' nelsoni:  (19)  (AMNH  80234);  (AUM  27948); 
(AUMP  299,  446,  449,  913,  1702,  1946,  1964,  1982, 
1992,  1994,  2200,  2413-16);  (USNM  101393, 
101398). 

P.  rubrivenlris:  (25)  (AMNH  69909-12,  77114, 
77587,  77613,  99145);  (AUMP  445,  2116,  2120); 
(CM  14022-29);  (UF/FSM  1821  -  six  specimens). 

P.  scripla:  (84)  (AUM  3828,  6993-97,  7574-76, 
7578-80,  11557-58,  11560,  13319,  21540,  24203, 
24258,  24261-62,  24264-68,  25125,  27016);  (AUMP 
11.0-11.21,  12-15,  16.1-.5,  285-87,  317,  692,  1720, 
1969-70,  1972-73,  1984,  1988,  1999,  2001.  2149, 
2155,  2173,  2214,  2222-24,  2406-09). 


LITERATURE  CITED 

BAUR,  G.  1890.  Two  new  species  of  tortoises  from 
the  South.  Science  16:  262-263. 

BJORK,  P.R.  1967.  Late  Eocene  vertebrates 
from  northwestern  South  Dakota.  J.  Paleontol- 
ogy, 41:  227-436. 

BOULENGER,  G.A.  1889.  Catalogue  of  the  chel- 
onians,  rhynchocephalians,  and  crocodiles  in  the 
British  Museum  (Natural  History).  Taylor  and 
Frances,. London.  541  pp. 

CAGLE,  F.R.  1952.  The  status  of  the  turtles 
Graptemys  pulchra  Baur  and  Graptemys  barbouri 
Carr  and  Marchand  with  notes  on  their  natural 
history.  Copeia  1952:  223-234. 

.  1953a.  Two  new  subspecies  of  Grapt- 
emys pseudogeographica.  Occ.  Pap.  Mus.  Zool. 
Univ.  Mich.,  546:  1-17. 

.  1953b.  The  status  of  the  turtle  Grapt- 


emys pseudogeographica.  Occ.  Pap.  Mus.  Zool. 
.   1954.  Two  new  species  of  the  genus 


Graptemys.  Tulane  Stud.  Zool.,  1:  167-186. 
CARR,  A.F.    1949.  The  identity  of  Malacoclem- 

mys  kohnii  Baur.  Herpetologica,  5:  9-10. 
.  1952.  Handbook  of  turtles.  Comstock 

Publ.  Assoc,  Ithaca,  New  York,  542  pp. 
and  C.J.  COIN.  1955.  Guide  to  the 


reptiles,   amphibians,   and   freshwater  fishes  of 
Florida.  Univ.  Florida  Press,  Gainesville.  341  pp. 

CLARK,  J.  1937.  The  stratigraphy  and  paleontol- 
ogy of  the  Chadron  Formation  in  the  Big  Bad- 
lands of  South  Dakota.  Ann.  Carnegie  Mus.,  25: 
261-350. 

CLARK,  W.E.  1978.  The  weevil  genus  Sibinia 
Germar:  natural  history,  taxonomy,  phylogeny, 
and  zoogeography,  with  revision  of  the  New 
World  species  (Coleoptera;  Curculionidae). 
Quaest.  Ent.,  14:  91-387. 

DOBIE,  J.L.  and  D.R.  JACKSON.  1979.  First 
fossil  record  for  the  diamondback  terrapin,  Mala- 
clemmys  terrapin  (Emydidae),  and  comments  on 
the  fossil  record  of  Chrysemys  nelsoni  (Emydi- 
dae). Herpetologica,  35:  139-145. 

DUNDEE,  H.A.  1974.  Evidence  for  specific 
status  of  Graptemys  kohni  and  Graptemys  pseud- 
ogeographica. Copeia  1974:  540-542. 

ERNST,  C.H.  1974.  Observations  on  the  court- 
ship of  male  Graptemys  pseudogeographica. 
Herpetol.  8:  377-378. 

and  R.W.  BARBOUR.  1972.  Turtles 

of  the  United  States.  The  Univ.  Press  of  Ky., 
Lexington,  Ky.  347  pp. 

GAFFNEY,  E.S.  1972a.  An  illustrated  glossary 
of  turtle  skull  nomenclature.  Am.  Mus.  Novit., 
2486:  1-33. 

.  1972b.  The  systematics  of  the  North 

American  family  Baenidae  (Reptilia,  Crytodira). 
Bull.  Amer.  Mus.  Nat.  Hist.,  147:  241-319. 

1975.  A  phylogeny  and  clasification  of 


the  higher  categories  of  turtles.  Bull.  Amer.  Mus. 
Nat.  Hist.,  155:  387-436. 


102 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Hay  O.P.  1908.  The  fossil  turtles  of  North 
America.  Publ.  Carnegie  Inst.  Washington,  75: 
IV  +  568  pp. 

Hay,  W.P.  1904.  A  revision  of  Malaclemmys,  a 
genus  of  turtles.  Bull.  U.S.  Bur.  Fisheries,  24: 
1-20. 

HENNIG,  W.  1966.  Phylogenetic  systematics. 
Univ.  IL.  Press,  Urbana,  IL.  263  pp. 

KILLEBREW,  F.C.  1977.  Miotic  chromosomes  of 
turtles.  IV.  The  Emydidae.  Texas  J.  Sci.,  29: 
245-253. 

.   1979.  Osteological  variation  between 

Graptemys  flavimaculata  and  Graplemys  nigri- 
noda  (Testudines:  Emydidae).  Herpetologica,  35: 
146-153. 

LOOMIS,  F.B.  1904.  Two  new  river  reptiles  from 
the  Titanothere  Beds.  Am.  J.  Science,  18:  427- 
432. 

LOVERIDGE,  A.  and  E.E.  WILLIAMS.  1957. 
Revision  of  the  African  tortoises  and  turtles  of 
the  suborder  Cryptodira.  Bull.  Mus.  Comp. 
Zool.,  115:  163-557. 

McDowell,  S.B.  1964.  Partition  of  the  genus 
Clernmys  and  related  problems  in  the  taxonomy 
of  the  aquatic  Testudinidae.  Proc.  Zool.  Soc. 
London,  143:  239-279. 

McKOWN,  R.R.  1972.  Phylogenetic  relation- 
ships within  the  turtle  genera  Graptemys  and 
Malaclemys.  Unpublished  Ph.D.  Dissertation, 
University  of  Texas  at  Austin.  Ill  pp. 

PARSONS,  T.S.  1960.  The  structure  of  the  choan- 
ae  of  the  Emydinae  (Testudines,  Testudinidae). 
Bull.  Mus.  Comp.  Zool,  123:  111-127. 

.  1968.  Variation  in  the  choanal  struc- 
ture of  recent  turtles.  Canadian  J.  Zool.,  46:1235- 
1263. 

PRITCHARD,  P.C.H.  1979.  Encyclopedia  of 
turtles.  T.F.H.  Publ.  Inc.  of  Neptune,  N.J.  895 
pp. 

SAY,  T.  1825.  On  the  fresh  water  and  land 
tortoises  of  the  United  States.  J.  Acad.  Nat.  Sci- 
ences, Philadelphia,  1:  203-219. 


TINKLE,  D.W.  1962.  Variation  in  shell  morphol- 
ogy of  North  American  turtles  I.  The  carapacial 
seam    arrangement.    Tulane    Stud.     Zool..    9: 

331-     349. 

VOGT,  R.C.  1978.  Systematics  and  ecology  of 
the  false  map  turtle  complex  Graptemys  pseudo- 
geographica.  Ph.D.  dissertation.  Univ.  of  Wis- 
consin -  Madison.  375  pp. 

.  1980.  Natural  history  of  the  map  tur- 
tles Graptemys  pseudogeographica  and  G.  ouach- 
itensis  in  Wisconsin.  Tulane  Stud.  Zool.  and  Bot., 
22:  17-48. 

and  C.J.  McCOY.  1980.  Status  of  the 


emydine  turtle  genera  Chrysemys  and  Pseudemys. 
Ann.  Carnegie  Mus.,  49:  93-102. 

WEAVER,  W.G.  and  F.L.  ROSE.  1967.  Syste- 
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Chrysems.  Tulane  Stud.  Zool.,  14:  63-73. 

WIED,  PRINZ  MAXIMILIAN  Z.  1865.  Ver- 
zeichniss  der  Reptilien  welche  auf  einer  Reise  im 
ndrdlichen  American  beobachtet  wiirden.  Nova. 
Act.  Acad.  Leopold  Carol.  Nat.  Curios,  32:  viii  + 
143. 

WOOD,  R.C.  1977.  Evolution  of  the  emydine 
turtles  Graptemys  and  Malaclemys  (Reptilia, 
Testudines).  J.  Herpetol.,  11:  415-421. 

ZANGERL,  R.  1969.  The  turtle  shell.  In:  Biology 
of  the  reptilia,  I.  Gans,  C,  and  T.S.  Parsons 
(eds.):  Academic  Press,  pp.  311-339. 

ZUG,  G.R.  1966.  The  penial  morphology  and  the 
relationships  of  cryptodiran  turtles.  Occ.  Pap. 
Mus.  Zool.  Univ.  Mich.,  647:  1-24. 

.  1971.  Buoyance,  locomotion,  morph- 
ology of  the  pelvic  girdle  and  hind  limb,  and 
systematics  of  cryptodiran  turtles.  Misc.  Publ. 
Mus.  Zool.  Univ.  Mich.,  142:  1-98. 


December  30,  1981 


ADDENDUM 

Tulane  Studies  in  Zoology  and  Botany  Volume  23,  number  I 

The  addendum  below  is  a  continuation  of  the  left  hand  paragraph  of  page  101  in  the  article 
by  Dobie.  It  ends  where  the  right  hand  paragraph  begins  LITERATURE  CITED. 

I'viU  -OOL 

LIBRARY 

OEC  201982 

HARVAKD 
UNIVIRRSITY 


p.  stejnegeri:  (5)  (AUMP  2363,  four  uncatalogued 
specimens). 

Rhinodemmys  areolata:  (1)  (AUMP  2111). 

R.  pukherrima:  (2)  (AUMP  910,  uncatalogued 
specimen). 

R.  unidentified  species:  (1)  (AUMP  2299). 

Terrapene  ornata:  (AUM  10732);  (AUMP  122-23, 
962,  1939). 

T.  Carolina:  (52)  (AUM  551,  1394.  1899,  3909-11, 
4998,  5925,  8866,  9414,  11611,  14295,  17634,  20942. 
23851,  25096-97);  (AUMP  116-20,  124,  128,  130-31, 
136-42,  702,  712,  914-16,  2250,  2257,  2312,  2317): 
(UF/FSM  7570,  14204,  35023,  38341,  40388, 
41508-09,  41518,  41521-22). 

Unidentified  genus  and  species:  (1)  (SDSM  &  T 
59187). 

Specimens  came  from  the  following  collections: 
Amerian  Museum  of  Natural  History  (AMNH); 
Auburn  University  Museum  (AUM);  Auburn  Uni- 
versity Museum  of  Paleontology  (AUMP);  Carnegie 
Museum  (CM);  Field  Mueum  of  Natural  History 
(FMNH);  University  of  Kansas  Museum  of  Natural 
History  (KU);  The  Vertebrate  Museum,  Northeast 
Louisiana  State  College  (NLSC);  South  Dakota 
School  of  Mines  and  Technology  (SDSM  &  T); 
Stephen  F.  Austin  State  University  Vertebrate 
Collection  (SFA);  State  University  of  South  Dakota 
(SUSD);  Texas  Cooperative  Wildlife  Collection, 
Texas  A&M  University  (TCWC);  Texas  Natural 
History  Collection,  Austin  (TNHC);  Tulane  Uni- 
versity Museum  (TU);  University  of  Florida,  Florida 
State  Museum  (UF/FS);  Museum  of  Southwestern 
Biology,  The  University  of  New  Mexico  (UNM); 
University  of  South  Alabama  (USA);  United  States 
Museum  of  Natural  History,  Smithsonian  Institu- 
tion (USNM). 


iaai>  UU0Z.-O/0Z. 


T&- 


Volume23  .  Number  2  $3.50  ^  ,,    .D^eember  15,  1982 


OtC2  0ii^82 

MARvArxD 
I  lMI\/irR~lTY 

CHANGES  IN  MELANIN  MIGRATION  INDUCED  BY  NORADRENERGIC 
AND  HISTAMINERGIC  AGENTS  IN  THE  FIDDLER  CRAB,  UCA  PUGILA  TOR 

MUKUND  M.  HANUMANTE  AND  MILTON  FINGERMAN  p.  103 

ADDITIONAL  TREMATODES  OF  MAMMALS  IN  LOUISIANA 

WITH  A  COMPILATION  OF  ALL  TREMATODES  REPORTED  FROM 

WILD  AND  DOMESTIC  MAMMALS  IN  THE  STATE 

WESLEY  L.  SHOOP  AND  KENNETH  C.  CORKUM  p.  109 

COMPARATIVE  VISCERAL  TOPOGRAPHY  OF  THE 

NEW  WORLD  SNAKE  TRIBE 

THAMNOPHIINI  (COLUBRIDAE,  NATRICINAE) 

NITA  J.  ROSSMAN  ,  DOUGLAS  A.  ROSSMAN 
and 
NANCY  K.  KEITH  P-  123 


TULANE  UNIVERSITY 
NEW  ORLEANS 


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TULANE  STUDIES  IN  ZOOLOGY  AND  BOTANY 


Volume  23,  Number  2 


December  15,  1982 


CHANGES  IN  MELANIN  MIGRATION  INDUCED  BY  NORADRENERGIC 
AND  HISTAMINERGIC  AGENTS  IN  THE  FIDDLER  CRAB,  UCA  PUGILA  TOR* 


MUKUND  M.  HANUMANTE  AND  MILTON  FINGERMAN 

Department  of  Biology,  Tulane  University 

New  Orleans,  Louisiana  70118  U.S.A. 


Abstract 

The  effects  of  the  H,  receptor  blocker  SA-97,  the 
Hi  receptor  blocker  cimetidine,  the  tyrosine  hydroxy- 
lase inhibitor  a -methyl-para-tyrosine  and  the  H, 
receptor  and  norepinephrine  uptakci  blocker  diphen- 
hydramine on  histamine-  or  4-methyl  histamine-in- 
duced  inhibition  of  melanin  dispersion  in  the  fiddler 
crab,  Uca  pugilator  undergoing  a  background  trans- 
fer from  white  to  black  were  determined.  Only  cimeti- 
dine significantly  antagonized  the  4-methyl  histamine- 
evoked  decrease  in  melanin  dispersion.  a-Methyl- 
para-tyroslne  by  itself  significantly  diminished  where- 
as diphenhydramine  by  itself  significantly  potentiated 
the  amount  of  this  centrifugal  melanin  migration  in 
the  fiddler  crabs.  None  of  these  drugs  affected 
melanin  migration  in  vitro.  The  results  are  consistent 
with  the  hypotheses  that  norepinephrine  triggers  re- 
lease of  a  melanin-dispersing  hormone  and  that  H,  re- 
ceptor activation  decreases  impulse-mediated  nore- 
pinephrine release  in  this  crab. 

INTRODUCTION 

Translocation  of  the  melanin  in  the 
melanophores  of  the  fiddler  crab,  Uca 
pugilator,  is  regulated  by  antagonistic 
neurohormones,  a  melanin-dispersing 
hormone  (MDH)  and  a  melanin-concen- 
trating hormone  (Carlson,  1935;  Sandeen, 
1950;  Fingerman,  1956).  Norepinephrine 
(NE)  triggers  release  of  MDH  in  this  crab 
(Fingerman  et  al.,  1981;  Hanumante  and 
Fingerman,  1981a,b;  1982a,b,c;  Hanu- 
mante et  al.,  1981).  Recently  histamine 
(HA)  has  been  shown  to  inhibit  melanin 

♦Supported  by  Grant  PCM-8 1-08864  from  the 
National  Science  Foundation. 


dispersion  in  a  dose-dependent  manner 
(Hanumante  and  Fingerman,  1981b).  Use 
of  a  variety  of  histaminergic  agonists  and 
antagonists  led  to  the  hypothesis  that  two 
types  of  HA  receptors,  called  H,  and  H2, 
are  present  on  NE  neurons  that  trigger 
MDH  release  and  that  HA  exerts  its  inhibi- 
tory action  by  stimulating  the  H2  recep- 
tors. The  present  investigation  was  devised 
to  obtain  further  support  for  this 
hypothesis.  This  objective  was  carried  out 
by  observing  the  effects  of  specific  mam- 
malian histaminergic  and  noradrenergic 
agents  not  used  previously  on  the  inhibi- 
tory action  of  HA  and  4-methyl  histamine 
(4-MeHA;  a  selective  H2  receptor  agonist, 
Owen  et  al.,  1979;  Douglas,  1980;  Polanin 
et  al.,  1981)  on  melanin  dispersion  in  Uca 
pugilator  transferred  from  a  white  to  a 
black  background. 

Materials  and  Methods 
Adult  male  fiddler  crabs,  Uca  pugilator, 
from  the  vicinity  of  Panacea,  Florida, 
(Gulf  Specimen  Company)  were  used. 
Their  melanophores  were  staged  according 
to  the  system  of  Hogben  and  Slome  (1931) 
whereby  stage  1 .0  represents  maximal  pig- 
ment concentration,  stage  5.0  maximal 
pigment  dispersion  and  stages  2.0,  3.0,  and 
4.0  the  intermediate  conditions.  When 
intact  crabs  were  used,  the  melanophores 
seen  through  the  cuticle  on  the  anteroven- 
tral  surface  of  the  second  walking  leg  on 
the  right  side  were  staged  at  the  time  a  sub- 


EDITORIAL  COMMITTEE  FOR  THIS  PAPER: 

DR.  RAY  W.  FULLER,  Research  Advisor,  Eli  Lilly  and  Company,  Indianapolis, 
Indiana  46206 

DR.  WILLIAM  S.  HERMAN,  Professor  and  Head,  Department  of  Genetics  and 
Cell  Biology,  University  of  Minnesota,  MinneapoHs,  Minnesota  55108 


103 


104 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


stance  was  injected  and  15,  30,  60,  90,  and 
120  minutes  thereafter.  To  facilitate  com- 
parison of  the  responses  of  the  experimen- 
tal and  control  crabs,  mean  differences 
between  the  15  through  120  minute 
melanophore  stages  for  the  control  and  ex- 
perimental groups  were  calculated  for  use 
in  Table  1 .  The  depicted  data  are  based  on 
the  mean  melanophore  stages  of  20  intact 
crabs  (10  experimental  and  10  control)  or 
20  isolated  legs  (10  experimental  and  10 
control).  When  assays  were  performed  on 
isolated  legs,  the  melanophores  were 
staged  only  at  the  time  the  legs  were 
removed  from  the  crab  (at  which  time  the 
legs  were  perfused  with  the  test  or  control 
solution)  and  15,  30,  45,  and  60  minutes 
thereafter.  The  second  and  third  walking 
legs  from  both  sides  of  the  crab  were 
removed;  the  legs  from  the  right  served  as 
experimentals  and  the  legs  from  the  left 
side  received  control  solution;  the  melano- 
phores on  the  anteroventral  surface  of 
these  isolated  legs  were  observed  for 
staging.  The  assays  were  performed  using 
isolated  legs  having  initially  either  maxi- 
mally concentrated  melanin  (stage  1.0)  or 
maximally  dispersed  melanin  (stage  5.0). 
Melanophores  in  isolated  legs  of  this  crab 
remain  responsive  for  at  least  120  minutes 
(Herman  and  Dallmann,  1975).  The  statis- 
tical significance  of  the  data  was  deter- 
mined using  Standard  Errors  of  the  Means 
(SEM)  the  Student's  t  test  with  significance 
set  at  the  95%  confidence  interval.  None 
of  the  data  for  isolated  legs  were  statis- 
tically significant. 

The  volume  of  the  solution  injected  into 
each  crab  or  isolated  leg  was  always  0.05 
ml.  The  experiments  with  intact  crabs  and 
isolated  legs  were  performed  at  24  °C  under 
an  illumination  of  1190  Ix.  4-MeHA  dihy- 
drochloride  (Smith,  Khne  and  French), 
cimetidine  (N"-Cyano-N-methyl-N'-{2- 
(5-methylimidazol-4-yl)  methylthioethyl} 
guanidine)  (Smith,  Kline  and  French)  and 
SA-97  (homochlorcyclizine)  (Eisai)  were 
generous  gifts.  In  addition,  HA,  amethyl- 
para-tyrosine  (a-MPT)  and  diphenhydra- 
mine HCl  (all  from  Sigma)  were  used.  The 
concentration  used  for  each  drug,  whether 


injected  alone  or  in  combination,  was  20 
ug/dose  of  the  free  compound.  All  drugs 
except  cimetidine  were  dissolved  in 
Pantin's  physiological  saline  (Pantin, 
1934).  Cimetidine  was  dissolved  in  acidi- 
fied (a  drop  of  1,2  M  HCl)  saline.  Conse- 
quently, a  drop  of  HCl  (1.2  M)  was  added 
to  control  saline  for  the  cimetidine  exper- 
iments. The  rest  of  the  controls  received 
pure  saline. 

Results  and  Discussion 

4-MeHA,  an  H2  receptor  agonist, 
slowed  the  rate  of  melanin  dispersion,  as 
observed  earlier  by  Hanumante  and  Fin- 
german  (1981b),  in  intact  crabs  transferred 
from  a  white  to  a  black  background  (Table 
1).  Cimetidine,  which  selectively  blocks 
mammalian  H2  receptors  (Douglas,  1980; 
Polanin  and  McNeill,  1981)  significantly 
antagonized  the  4-MeHA.  On  the  other 
hand,  the  H,  receptor  blocker  SA-97  not 
only  did  not  antagonize  the  4-MeHA  but 
the  combination  of  4-MeHA  plus  SA-97 
resulted  in  significantly  further  inhibition. 
None  of  these  drugs  affect  melanin  migra- 
tion in  vitro  nor  do  SA-97  and  cimetidine 
by  themselves  have  an  effect  on  the  rate  of 
melanin  dispersion  in  crabs  undergoing  a 
background  change  from  white  to  black 
(Hanumante  and  Fingerman,  1981b),  a 
black  background  fostering  melanin  dis- 
persion (Brown  and  Hines,  1952)  which 
will  be  effected  by  MDH. 

a-MPT  selectively  inhibits  tyrosine 
hydroxylase.  This  enzyme  catalyzes  the 
synthesis  of  dihydroxyphenylalanine  from 
tyrosine.  At  least  in  mammals  this  is  the 
rate-limiting  step  in  the  biosynthesis  of 
NE  (Terrasawa  et  al.,  1975;  Lofstrom  and 
Backstrom,  1978).  a  MPT  by  itself  signi- 
ficantly decreased  melanin  dispersion. 
HA  by  itself,  as  reported  earlier 
(Hanumante  and  Fingerman,  1981b),  sig- 
nificantly reduced  centrifugal  melanin 
migration  in  intact  crabs  transferred  from 
a  white  to  a  black  background.  However, 
in  the  crabs  that  were  co-administered 
either  4-MeHA  and  a-MPT  or  HA  and 
a-MPT  (Table  1),  4-MeHA  and  HA  were 
not  able  to  produce  further,  significant 


No.  2 


Melanin  Migration  in  Crabs 


105 


reduction  of  the  melanin  dispersion. 
Diphenhydramine,  a  blocker  of  H,  recep- 
tors and  NE  uptake,  in  mammals  (Isaac 
and  Goth,  1965;  Fantozzi  et  al.,  1975; 
Marco  et  al.,  1980),  by  itself  significantly 
enhanced  melanin  dispersion.  However, 
when  HA  was  co-administered  with 
diphenhydramine,  the  HA-induced  inhi- 
bition in  melanin  dispersion  was  still 
evident  (Fig.  1). 

The  present  data,  in  light  of  our  earlier 
report  (Hanumante  and  Fingerman, 
1981b)  and  the  pharmacological  actions 
of  noradrenergic  and  histaminergic  agents 
in  mammals,  further  strengthen  the  hypo- 
thesis that  (a)  NE  serves  as  a  neurotrans- 
mitter triggering  release  of  MDH  and  that 
(b)  activation  of  H2  receptors  located  on 
NE  neurons  which  control  MDH  release 
results  in  a  decrement  of  melanin  disper- 
sion in  Uca  pugilator  transferred  from  a 
white  to  a  black  background.  The 
observations  that  cimetidine,  a  selective 
H2  receptor  blocker,  antagonized  the 
4-MeHA-induced  inhibition  in  melanin 
dispersion,  whereas  the  Hi  blocker  SA-97 
did  not,  reveal  that  this  effect  is  mediated 
specifically  by  activation  of  HA  H2  recep- 
tors. The  marked  increase  in  inhibitory 
effect  of  4-MeHA  when  co-administered 
with  the  Hi  antagonist  SA-97  was 
probably  due  to  the  fact  that  excitation  of 
H,  receptors  evokes  enhanced  melanin 
dispersion  (Hanumante  and  Fingerman, 
1981b),  blocking  them  would  prevent  any 
endogenous  Hi  stimulation  of  the  crabs. 
This  would  enable  4-MeHA,  an  agonist  of 
H2  receptors,  to  produce  an  even  greater 
inhibition  of  the  melanin  dispersion.  On 
the  contrary,  in  the  crabs  whose  H2  recep- 
tors were  blocked  by  cimetidine,  4-MeHA 
was  unable  to  significantly  decrease  the 
action  potential-mediated  release  of  NE, 
which  in  turn  resulted  in  a  near  normal 
quantity  of  MDH  being  released  into  the 
hemolymph  of  these  crabs  transferred  to 
the  black  background.  The  fact  that 
metiamide,  another  H2  receptor  blocker, 
significantly  antagonized  the  4-MeHA- 
stimulated  decrease  in  centrifugal  melanin 
migration  (Hanumante  and  Fingerman, 


1981b)  in   vivo  further  strengthens  this 
conclusion. 

NE  has  been  found  (0.51  pg/g)  in  the 
supraesophageal  ganglia  of  male  fiddler 
crabs  (Hanumante  and  Fingerman, 
1982b).  Also,  we  have  provided  evidence 
that  Hi  and  H2  receptors  occur  on  NE 
neurons  because  in  fiddler  crabs  pretreated 
with  6-hydroxydopamine  (which 
presumably  destroys  NE  neuroterminals  in 
Uca  as  it  does  in  vertebrates)  (Hanumante 
and  Fingerman,  1982b,c)  HA  is  unable  to 
significantly  reduce  further  the  melanin 
dispersion  (Hanumante  and  Fingerman 
1981b).  We  have  not  determined  (i)  the 
levels  of  NE  in  a-MPT  injected  crabs  or  (ii) 
the  exact  mechanism  of  action  of  a-MPT 
in  Uca  puilator.  However,  data  that  we 
obtained  using  noradrenergic  and  histadre- 
nergic  agents  (Hanuamante  and  Finger- 
man, 1981b)  reveal  that  20  MPT  clearly 
interferes  with  NE  neurotransmission. 
This  probably  was  either  by  way  of  its  well- 
established  (at  least  in  mammals)  pharma- 
cological NE  synthesis-inhibiting  effect 
(Terraswawa  et  al.,  1975;  Lofstrom  and 


HOURS 

Figure  1.  Relationships  between  melanophore  stage 
and  time.  Circles  with  bottom-half  darkened,  crabs 
that  received  diphenhydramine;  circles  with  top-half 
darkened,  crabs  that  received  histamine;  solid  circles, 
crabs  that  received  histamine  plus  diphenhydramine; 
open  circles,  salme-injected  controls.  Vertical  bars 
indicate  SEM. 


106 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Table  i .  The  means  ( ±  SEM)  of  the  differences  between  the  melanophore  stages  de- 
termined at  15,  30,  60,  90,  and  120  minutes  of  the  intact  crabs  that  received  a 
drug  versus  the  saline-injected  controls.  The  minus  sign  indicates  decreased 
melanin  dispersion  relative  to  the  controls.  *Statistically  significant  p^  .05 
relative  to  respective  controls. 


4-Methyl  histamine  (4-MeHA) 

Cimetidine 

4-MeHA  plus  cimetidine 

4-MeHA  plus  SA-97 

a  Methyl-p-Tyrosine  (a-MPT) 

4-MeHA  plus  a-MPT 

Histamine  (HA) 

HA  plus  a-MPT 


-0.67*(±  0.08) 
-0.17  (±  0.01) 
-0.39  (±  0.07) 
-1.43*(±  0.12) 
-1.15*(±  0.15) 
-1.44*(±  0.21) 
-1.18*(±  0.18) 
-1.01*(±  0.12) 


Backstrom,  1978;  Douglas,  1980)  or  by 
stimulating  H2  receptors,  thereby  leading 
to  the  observed  decrement  in  MDH  release 
(Table  1).  Hence,  the  melanin  of  these 
a -MPT-treated  crabs  did  not  disperse  to 
the  extent  it  did  in  the  control  animals. 

As  stated  above,  in  the  crabs  co-injected 
with  4-MeHA  and  a-MPT  or  HA  and 
a-MPT,  neither  4-MeHA  nor  HA  signifi- 
cantly affected  the  melanin  dispersion 
compared  with  that  which  occurred  in  re- 
sponse to  a-MPT  alone  (Table  1).  This  pre- 
sumably was  due  to  the  interference  with 
NE  neurons  by  a-MPT  in  such  a  way  that 
the  impulse-mediated  decrement  in  NE 
secretion  by  the  H2  stimulators  4-MeHA 
and  HA  was  not  large  enough  to  affect 
significantly  the  NE-mediated  MDH 
release. 

The  diphenhydramine-evoked  increment 
in  melanin  dispersion  (Fig.  1)  was  pre- 
sumably due  to  its  blocking  action  on  NE 
uptake,  (Marco  et  al.,  1980).  NE  uptake, 
inhibitors  like  nisoxetine  (Koe,  1976)  have 
already  been  shown  to  potentiate  MDH 
release  (Hanumante  and  Fingerman, 
1981a).  Diphenhydramine  antagonizes  H, 
receptors  (Isaac  and  Goth,  1965;  Fantozzi 
et  al.,  1975;  Marco  et  al.,  1980)  also.  How- 
ever, because  H,  receptor  blockers  do  not 
significantly  abolish  HA-  or  4-MeHA-  (an 
H:  receptor  agonist)  mediated  inhibition  of 
melanin  dispersion,  we  suggest  that  the  NE 
uptake,  blocking  action  of  diphenhydra- 


mme  is  responsible  for  the  potentiation  of 
melanin  dispersion.  The  observation  that 
even  when  HA  is  co-administered  with  di- 
phenhydramine there  is  still  a  decrease  in 
melanin  dispersion  (Fig.  1)  indicates  that 
HA  does  not  evoke  its  effect  by  stimulating 
NE  uptake,;  uptake,  being  the  major 
mechanism  of  inactivating  the  postsyn- 
aptic actions  of  monoamines  including  NE 
(Fuller  and  Wong,  1977).  That  none  of 
these  drugs  affect  significantly  melanin 
migration  in  isolated  legs  (Hanumante  and 
Fingerman,  1981b)  is  consistent  with  the 
hypothesis  that  these  drugs  elicit  changes 
in  melanin  dispersion  indirectly  by  inter- 
acting with  the  neuroendocrine  system  of 
Uca  pugilator. 

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108 


Tulane  Studies  in  Zoology  and  Botany  Vol .  23 


ADDITIONAL  TREMATODES  OF  MAMMALS  IN  LOUISIANA 

WITH  A  COMPILATION  OF  ALL  TREMATODES  REPORTED  FROM 

WILD  AND  DOMESTIC  MAMMALS  IN  THE  STATE 

WESLEY  L.  SHOOP  AND  KENNETH  C.  CORKUM 

Department  of  Zoology  and  Physiology,  Louisiana  State  University 

Baton  Rouge,  Louisiana  70803 


Abstract 

The  following  trematodes  were  collected  from 
hunter-trapped  mammals  in  the  Atchafalaya  basin  of 
Louisiana  during  the  winters  of  1981  and  1982:  Alaria 
alarioides  (Dubois,  1937)  Dubois,  1970  from  mink, 
Mustela  vison  Schreber,  and  river  otter,  Lutra  cana- 
densis (Schreber);  Alaria  marcianae  (La  Rue,  1917) 
Walton,  1949  from  raccoon,  Procyon  lotor  (Linn.) 
and  bobcat,  Lynx  rufus  (Schreber);  Alaria  mustelae 
Bosma,  1931  from  raccoon  and  mink;  Amphimerus 
speciosus  (Stiles  and  Hassal,  1896)  Barker,  1911  from 
raccoon  and  the  domestic  cat.  Fells  domesticus  Linn.; 
Baschklrovitrema  incrassatum  (Dies.,  1850)  Skrjabin, 
1944  from  mink  and  river  otter;  Brachylaima  virgin- 
iana  Dickerson,  1930  from  opossum,  Dldelphis  vir- 
giniana  Kerr;  Carneophallus  basodactylophallus 
Bridgman,  1969  from  raccoon;  Cryptocotyle  concava 
(Creplin,  1825)  Lube,  1899  from  mink;  Fibricola 
cratera  (Barker  and  Noll,  1915)  Dubois,  1932  from 
mink,  opossum,  and  raccoon;  F.  lucida  (La  Rue  and 
Bosma,  1927)  Dubois  and  Rausch,  1950  from  mink 
and  opossum;  Gyrosoma  stngulare  Byrd,  Bogitsh, 
and  Maples,  1%1  from  raccoon  and  mink;  Hasstllesia 
texensis  Chandler,  1929  from  muskrat.  Ondatra  zibe- 
thica  (Linn.);  Heterobllharzia  americana  Price,  1929 
from  mink,  raccoon,  and  bobcat;  Isthmiophora  mells 
(Schrank,  1788)  Luhe,  1909  from  raccoon  and  mink; 
Linstowiella  szldati  (Anderson,  1944)  Anderson  and 
Cable,  1950  from  opossum  and  raccoon;  Marltremtn- 
oides  nettae  (Gower,  1938)  Rankin,  1939  from  rac- 
coon and  mink;  Microphallus  opacus  (Ward,  1894) 
Ward,  1901  from  raccoon  and  mink;  Paragonlmus 
kellkottl  Ward,  1908  from  opossum;  Pharyngosto- 
moldes  procyonis  Harkema,  1942  from  raccoon; 
Quinqueserialis  qulnqueserialis  (Barker  and  Laughlin, 
1911)  Harwood,  1939  from  muskrat;  Rhopalias  ma- 
cracanthus  Chandler,  1932  from  opossum;  and  Sella- 
cotyle  vitellosa  Sogandares-Bernal,  1961  from  mink. 


Alaria  alarioides,  A.  marcianae,  Amphimerus  spe- 
ciosus, Cryptocotyle  concava,  Isthmiophora  mells. 
Microphallus  opacus,  Paragonlmus  kellkottl,  and 
Qulnqueserialis  qulnqueserialis  have  not  been  pre- 
viously reported  from  Louisiana  mammals.  Diag- 
noses are  presented  for  the  species  representing  state 
records  along  with  pertinent  notes  on  the  biology  of 
each.  New  host  records  include  Heterobllharzia 
americana,  Cryptocotyle  concava,  and  Maritremi- 
noides  nettae  from  mink;  Alaria  marcianae,  Amphi- 
merus speciosus,  and  Linstowiella  szldati  from  rac- 
coon; and  Hasstllesia  texensis  from  muskrat.  A  com- 
pilation of  trematodes  previously  reported  from  Loui- 
siana mammals  is  presented. 


INTRODUCTION 

Recently,  we  reported  some  trematodes 
collected  from  mammals  in  south  Louisi- 
ana (Shoop  and  Corkum,  1981a).  Since 
that  time  we  have  continued  our  examina- 
tion of  hunter-trapped  mammals  from  the 
Atchafalaya  basin  of  Louisiana  during  the 
winters  of  1981  and  1982.  The  following 
mammals  were  examined  for  trematodes: 
42  minks,  Mustela  vison  Schreber;  37  rac- 
coons, Procyon  lotor  (Linn.);  seven  river 
otters,  Lutra  canadensis  (Schreber);  five 
muskrats,  Ondatra  zibethica  (Linn.);  three 
bobcats,  Lynx  rufus  (Schreber);  four 
domestic  cats,  Felis  domesticus  Linn.;  two 
opossums,  Dldelphis  virginiana  Kerr;  and 
three  red  foxes,  Vulpes  fulva  (Desmarest). 
The  red  foxes  were  found  uninfected  with 
trematodes. 

Trematodes  were  fixed  in  steaming  10% 


EDITORIAL  COMMITTEE  FOR  THIS  PAPER: 

DR.  BERT  B.  BABERO,  Professor  of  Biological  Sciences,  University  of  Nevada, 
Las  Vegas,  Las  Vegas,  Nevada  89154 

DR.  WALTER  E.  WILHELM,  Associate  Professor  of  Biology,  Memphis  State 
University,  Memphis,  Tennessee  38152 


109 


no 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


formalin  and  stained  in  Semichon's  aceto- 
carmine.  All  measurements  are  in  micro- 
meters unless  otherwise  stated;  means  are 
followed  by  the  ranges  in  parentheses.  Line 
drawings  were  prepared  with  the  aid  of  a 
microprojector.  Representative  specimens 
of  the  species  for  which  diagnoses  are 
given  were  deposited  in  the  Manter  Lab- 
oratory, University  of  Nebraska  State 
Museum,  Lincoln,  Nebraska. 

Table  I  lists  the  trematodes  recovered 
from  the  eight  species  of  mammals. 
Lumsden  and  Zischke  (1961)  reported  and 
diagnosed  Fibricola  cratera,  F.  lucida, 
Hasstilesia  texensis,  Brachylaima  virgin- 
iana,  and  Rhopalias  macracanthus  from 
Louisiana  mammals.  Our  specimens  agree 
in  all  respects  with  Lumsden  and  Zischke's 
(1961)  diagnoses.  Our  specimens  of  Hassti- 
lesia texensis  from  the  muskrat  represent  a 
new  host  record.  Shoop  and  Corkum 
(1981a)  reported  and  diagnosed  Alaria 
mustelae,  Baschkirovitrema  incrassatum, 
Gyrosoma  singulare,  Maritreminoides 
nettae,  and  Pharyngostomoides  procyonis 
from  Louisiana  mammals.  In  that  report 
we  noted  M.  nettae  in  raccoons;  it  is  herein 
reported  from  the  mink  as  well  (new  host 
record).  In  a  more  recent  note,  we  (Shoop 
and  Corkum,  1982)  commented  further  on 
the  status  of  G.  singulare  in  this  state.  He- 
terobilharzia  americana  has  been  reported 
from  Louisiana  mammals  by  Malek  et  al. 
(1961)  and  Kaplan  (1964).  Our  collections 
of  H.  americana  from  mink  represent  a 
new  host  record.  Carneophallus  basodac- 
tylophallus  was  originally  described  by 
Bridgman  (1969)  from  raccoon  in  Louisi- 
ana as  was  Sellacotyle  vitellosa  from  mink 
by  Sogandares-Bernal  (1961).  Lumsden 
and  Winkler  (1962)  reported  Linstowiella 
szidati  from  opossum.  We  have  found  it  in 
opossum  as  well  as  in  raccoon.  In  addition 
to  these  trematodes,  we  identified  eight 
other  species  that  have  not  been  previously 
reported  from  Louisiana  mammals  and 
that  are  of  importance  from  epidemiolo- 
gical or  zoogeographical  standpoints. 
Table  II  compiles  all  trematodes  reported 
heretofore  from  mammals  in  the  state  of 
Louisiana. 


Family  DIPLOSTOMIDAE  Poirier,  1886 

Alaria  alarioides  (Dubois,  1937) 

Dubois,  1970 

(Figure  1) 

Synonyms:    Diplostomum    alarioides 
Dubois,  1937;  Enhydrodiplostomum  alar- 
ioides (Dubois,  1937)  Dubois,  1944. 
Hosts:  Mustela  vison  Schreber  and  Lutra 
canadensis  (Schreber). 
Location:  Small  intestine. 
Locality:  Belle  River,  Assumption  Parish, 
Louisiana. 

Deposition:  Univ.  Nebraska  State  Mus., 
Manter  Lab.  Coll.  No.  21367. 
Diagnosis  (based  on  ten  mature 
specimens):  Body  elongate,  distinctly 
bisegmented,  1650  (1400-1800)  long  by  540 
(450-650)  at  the  widest  point.  Forebody 
spathulate,  777  (640-940)  long  by  540 
(450-650)  wide;  pseudosuckers  present  as 
depressions  on  either  side  of  the  oral 
sucker,  never  observed  evaginated.  Hind- 
body  claviform,  907  (760-1050)  long  by 
430  (400-480)  wide,  containing  reproduc- 
tive organs.  Forebody  tegument  covered 
with  small  spines;  hindbody  smooth.  Oral 
sucker  terminal,  92  (80-100)  long  by  106 
(90-120)  wide;  acetabulum  weak,  spher- 
ical, 75  (60-80)  long  by  76  (60-90)  wide, 
often  covered  by  the  tribocytic  organ; 
tribocytic  organ  broadly  elliptical  when 
evaginated,  348  (240-400)  long  by  280 
(240-330)  wide,  with  a  longitudinal  cleft. 

Prepharynx  and  esophagus  extremely  short 
or  absent;  pharynx  usually  in  contact  with 
oral  sucker,  77  (70-90)  long  by  65  (50-80) 
wide;  paired  ceca  extend  to  the  posterior 
end  of  body.  Testes  tandem,  not  equal; 
anterior  testis  asymmetrical,  laterally  dis- 
posed on  either  side  of  midline,  215 
(200-250)  long  by  317  (290-350)  wide;  pos- 
terior testis  symmetrical,  dumbbell- 
shaped,  much  wider  than  anterior  testis, 
218  (190-250)  long  by  394  (350-410)  wide, 
with  a  ventro-median  groove  to  allow  pas- 
sage of  ceca,  uterus,  and  vitellaria;  ejacula- 
tory  duct  opens  into  the  genital  atrium; 
genital  atrium  opens  posterior,  subterm- 
inally  on  the  dorsal  surface.  Ovary  spher- 
ical, located  in  hindbody  just  in  front  of 


No.  2 


Trematodes  of  Mammals 


111 


Table  I.  Trematodes  recovered  from  hunter-trapped  mammals  in  Louisiana  during  the  winters  of  1981 
and  1982. 


Trematode 


Hosts 


No. 
Examined 


No. 
Infected 


% 


Location 


Alaria  alarioides  (Dubois,  1937) 

Dubois,  1970 
A.  marcianae  (La  Rue,  1917) 

Walton,  1949 
A.  mustelae  Bosma,  1931 

Amphimerus  speciosus  (Stiles  and 

Hassal,  1896)  Barker,  1911 
Baschkirovitrema  incrassatum 

(Dies.,  1850)  Skrjabin,  1944 
Brachylaima  virginiana 

Dickerson,  1930 
Carneophallus  basodactylophallus 

Bridgman,  1969 
Cryptocotyle  concava  (Creplin,  1825) 

Luhe,  1899 


otter 

7 

mink 

42 

raccoon 

37 

bobcat 

3 

raccoon 

37 

mink 

42 

raccoon 

37 

domestic  cat 

4 

otter 

7 

mink 

42 

opossum 

2 

raccoon 

37 

mink 


Fibricola  cratera  (Barker  and  Noll, 
1915)  Dubois,  1932 

F.  lucida  (La  Rue,  and  Bosma,  1927) 
Dubois  and  Rausch,  1950 

mink 

raccoon 

opossum 

mink 

opossum 

Gyrosoma  singulare  Byrd,  Bogitsh, 
and  Maples,  1961 

raccoon 
mink 

Hasstilesia  texensis  Chandler, 

1929 
Heterobilharzia  americana 

Price,  1929 

muskrat 

raccoon 
mink 

bobcat 

Isthmiophora  melis  (Schrank,  1788) 
Luhe,  1909 

raccoon 
mink 

Linstowiella  szidati  (Anderson,  1944) 
Anderson  and  Cable,  1950 

raccoon 
opossum 

Maritreminoides  nettae  (Gower,  1938) 
Rankin,  1939 

mink 
raccoon 

Microphallus  opacus  (Ward,  1894) 
Ward,  1901 

raccoon 
mink 

Paragonimus  kellicotti  Ward, 
1908 

opossum 

Pharyngostomoides  procyonis 
Harkema,  1942 

raccoon 

Quinqueserialis  quinqueserialis 
(Barker  and  Laughlin,  1911) 
Harwood,  1939 

muskrat 

Rhopalias  macracanthus 
Chandler,  1932 

opossum 

Sellacotyle  vitellosa  Sogandares- 
Bernal,  1961 

mink 

42 

42 

37 
2 

42 
2 

37 

42 
5 

37 

42 

3 

37 
42 
37 

2 
42 
37 
37 
42 

2 

37 


2 
42 


2 

24 
2 
2 
1 
1 
1 
1 
2 

21 
1 


22 

4 

12 
2 

26 

2 
7 
2 
1 

20 
2 
1 
6 

2 
1 
1 
3 
6 
5 
4 
1 

31 


29 

57 

5 

67 

3 

2 

3 

25 

29 

50 

50 


52 


Sm.  Int. 


Liver 
Sm 


Int. 


10 

32 

100 

62 

100 

19 

5 

20 

Cecum 

54 

Mes.  Ven 

5 

33 

16 

Sm.  Int. 

5 

3 

50 

7 

16 

14 

10 

50 

Lungs 

84 

Sm.  Int. 

40 

Cecum 

50 

Sm.  Int. 

5 

» 

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Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


the  anterior  testis,  103  (90-120)  long  by  1 14 
(110-120)  wide;  uterus  courses  anteriad 
into  the  forebody  and  turns  immediately 
posteriad  where  it  opens  in  the  genital 
atrium;  vitellaria  penetrate  the  forebody 
and  extend  in  two  bands  through  the  ven- 
tro-medial  grooves  of  the  testes  to  the  level 
of  the  genital  atrium;  vitelline  reservoir 
median,  intertesticular.  Eggs  large, 
operculate,  101  (90-1 10)  long  by  55  (50-60) 
wide.  Excretory  system  not  observed. 
Discussion:  Dubois  (1937)  originally  des- 
cribed Diplostomum  alarioides  from  a 
Brazilian  otter.  He  (Dubois,  1944)  subse- 
quently purged  the  genus  Diplostomum  of 
all  mammalian  parasites,  retaining  it  for 
avian  parasites,  and  erected  the  new  genus 
Enhydrodiplostomum  for  D.  alarioides 
and  a  second  otter  parasite,  D.  fosteri. 
Chandler  and  Rausch  (1946)  assigned  two 
additional  species,  Alaria  clathrata  and  A. 
pseudoclathrata,  both  also  parasites  of  the 
otter,  to  the  genus  Enhydrodiplostomum. 
In  a  later  revision,  Dubois  (1970)  agreed 
that  these  four  species  are  closely  related, 
but  reassigned  them  to  the  genus  Alaria 
where  additional  mustelid  parasites  are 
found. 

Sawyer's  (1961)  collection  of  A.  alari- 
oides from  river  otter  in  Georgia  was  the 
first  report  from  North  America.  Since 
then.  Miller  and  Harkema  (1964,  1968) 
reported  y4.  alarioides  from  both  mink  and 
river  otter  in  North  Carolina,  and  Fleming 
et  al.  (1977)  reported  it  from  river  otter  in 
Alabama.  A.  alarioides  is  also  a  common 
parasite  of  mink  and  river  otter  in  Louisi- 
ana. Measurements  oi  A.  alarioides  from 
the  two  hosts  compare  favorably  with  the 
descriptions  of  Dubois  (1937,  1970). 

Alaria  marcianae  (La  Rue,  1917) 

Walton,  1949 

(Figure  2) 

Synonyms:  Cercaria  marcianae  La  Rue, 
1917;  Agamodistomum  marcianae  (La 
Rue,  1917)  Cort,  1918;  Alaria  americana 
Hall  and  Wigdor,  1918;  Alaria  canis  La 
Rue  and  Fallis,  1934;  Alaria  minnesotae 
Chandler,  1954. 


Hosts:  Lynx  rufus  (Schreber)  and  Procyon 
lotor  (Linn.). 
Location:  Small  intestine. 
Locality:  Pierre  Part,  Assumption  Parish, 
Louisiana. 

Deposition:  Univ.  Nebraska  State  Mus., 
Manter  Lab.  CoU.  No.  21368. 
Diagnosis  (based  on  ten  mature 
specimens):  Body  elongate,  distinctly  bi- 
segmented,  1375  (1000-1600)  long  by  478 
(350-600)  at  the  widest  point.  Forebody 
spathulate  with  lateral  margins  folded  ven- 
trally  where  they  meet  at  the  midline,  the 
entire  forebody  serving  as  an  organ  of 
attachment,  883  (650-1050)  long  by  478 
(350-600)  wide;  ear-like  appendages  pre- 
sent on  either  side  of  the  oral  sucker,  rarely 
observed  invaginated  to  form  pseudo- 
suckers.  Hindbody  conical,  535  (400-650) 
long  by  363  (280-500)  wide,  containing  re- 
productive organs.  Forebody  tegument 
covered  with  small  spines,  hindbody  tegu- 
ment smooth.  Oral  sucker  terminal  90 
(60-105)  long  by  73  (60-81)  wide;  aceta- 
bulum weak,  spherical,  74  (60-95)  long  by 
75  (60-95)  wide,  rarely  covered  by  the 
tribocytic  organ;  tribocytic  organ  elongate 
when  evaginated,  453  (310-550)  long  by 
200  (155-225)  wide,  with  a  longitudinal 
cleft.  Prepharynx  present,  5  (4-6)  long; 
pharynx  pyriform,  102  (75-215)  long  by  64 
(55-85)  wide;  esophagus  6  (4-10)  long; 
paired  ceca  extend  to  the  posterior  end  of 
the  body.  Testes  tandem,  not  equal;  ante- 
rior testis  asymmetrical,  typically  wedge- 
shaped,  laterally  disposed  on  either  side  of 
the  midUne,  160  (128-215)  long  by  225 
(175-300)  wide;  posterior  testis  symmetri- 
cal, dumbbell-shaped  much  wider  than 
anterior  testis,  210  (165-276)  long  by  340 
(275-425)  wide,  with  a  ventro-medial 
groove  to  allow  passage  of  ceca  and  uterus; 
muscular  ejaculatory  pouch  lies  posterior 
to  the  testes  and  empties  into  the  genital 
atrium;  genital  atrium  located  in  the  poste- 
rior end  of  the  body,  opening  on  the 
dorso-subterminal  side.  Ovary  reniform, 
located  in  front  of  the  anterior  testis  on 
either  side  of  midline,  72  (60-99)  long  by 
167  (100-180)  wide;  Mehlis'  gland  opposite 


No.  2 


Trematodes  of  Mammals 


113 


the  ovary;  uterus  courses  briefly  into  the 
forebody  and  turns  immediately  posteriad 
where  it  empties  into  the  genital  atrium; 
vitellaria  located  only  in  the  forebody, 
from  just  in  front  of  the  acetabulum  to  the 
forebody-hindbody  juncture;  vitelline  re- 
servoir prominent,  located  in  the  hindbody 
at  the  level  of  the  anterior  testis.  Eggs  few, 
large,  operculate,  122  (110-128)  long  by  65 
(60-75)  wide.  Excretory  pore  terminal, 
remainder  of  excretory  system  not 
observed. 

Discussion:  Apparently,  adult  Alaria 
marcianae  have  not  previously  been 
reported  from  Louisiana.  A  single  speci- 
men of  A.  americana  (=  A.  marcianae) 
from  a  dog  from  Baton  Rouge  was  de- 
posited by  G.  Dikmans  (USNM  Helm. 
Coll.  No,  25159).  We  examined  that  speci- 
men and  identify  it  as  /I .  marcianae,  being 
similar  to  our  material  from  the  bobcat. 

In  a  previous  report,  the  epidemiology 
of  A.  marcianae  mesocercariae  was  studied 
in  Louisiana  and  evidence  was  presented 
that  this  species  was  responsible  for  an 
authochtonous  human  infection  (Shoop 
and  Corkum,  1981b).  In  experimental  in- 
fections only  juvenile  raccoons  served  as 
definitive  hosts  for  A.  marcianae.  Adult 
raccoons  proved  to  be  refractory  to  the 
development  of  the  mesocercarial  stage, 
which  remained  undifferentiated  in  the 
subcutaneous  fat.  These  findings  were  cor- 
roborated in  the  present  study  because  no 
adult  raccoons  were  found  infected.  Two 
yearlings,  however,  harbored  several  adult 
A.  marcianae  in  their  duodena.  This  is  the 
first  report  of  raccoon  naturally  infected 
with  this  species.  Though  these  worms 
from  the  yearlings  exhibited  no  morpholo- 
gical anomalies,  they  were  smaller  than 
specimens  from  the  bobcat. 

The  known  definitive  hosts  for  A .  mar- 
cianae in  Louisiana  now  include  the 
domestic  dog,  bobcat,  and  juvenile  rac- 
coons. In  experimental  laboratory  infec- 
tions we  have  found  that  the  domestic  cat 
is  a  suitable  definitive  host  and  that  it,  as 
well  as  feral  cats,  may  play  a  significant 
role  in  the  maintenance  of  A.  marcianae  in 
Louisiana. 


Family  OPISTHORCHIIDAE 

Braun,  1901 

Amphimerus  speciosus 

(Stiles  and  Hassal,  1896)  Barker,  1911 

(Figure  3) 

Synonyms:  Amphimerus  caudalitestis 
Caballero,  Grocott,  and  Zerecero,  1953; 
A.  guayaquilensis {Rodriguez,  Gomez,  and 
Montalvan,  1948)  Caballero,  Grocott,  and 
Zerecero.  1953;  A.  interruptus  (Braun, 
1901)  Barker,  1911;  A.  minimus  Thatcher, 
1970;  A.  neotropicalis  Caballero,  Mon- 
tero-Gei,  and  Caballero,  1963;  A.  parcio- 
vatus  Franco,  1967;  A.  pricei  (Foster, 
1939)  Yamaguti,  1958;  A.  pseudofelmeus 
(Ward,  1901)  Barker,  1911. 
Hosts:  Felis  domes ticus  Linn,  and  Procyon 
lotor  (Linn.). 

Location:  Liver  and  bile  ducts. 
Locality:  Ramah,  Iberville  Parish,  Louisi- 
ana. 

Deposition:  Univ.  Nebraska  State  Mus., 
Manter  Lab.  Coll.  No.  21369. 
Diagnosis  (based  on  ten  mature 
specimens):  Body  elongate,  sharply  ta- 
pered anterior  to  the  acetabulum,  10.25 
(8.0-12.25)  mm  long  by  2010  (1150-2400) 
at  the  widest  point.  Tegument  beset  with 
small,  stout  spines.  Oral  sucker  268 
(240-300)  long  by  313  (270-340)  wide; 
acetabulum  200  (150-240)  long  by  218 
(170-250)  wide.  Prepharynx  absent; 
pharynx  183  (160-200)  long  by  173 
(150-190)  wide;  esophagus  170  (120-200) 
long;  paired  ceca  extend  to  the  posterior 
end  of  body.  Testes  tandem,  in  posterior 
Vi  of  body,  transversely  elongate,  sHghtly 
lobed;  anterior  testis  498  (410-600)  long  by 
925  (550-1150)  wide;  posterior  testis  573 
(450-720)  long  by  925  (550-1150)  wide; 
seminal  vesicle  elongate,  coiled,  opens  into 
the  genital  atrium  which  is  immediately 
preacetabular.  Ovary  oval  to  reniform, 
may  be  slightly  lobed,  325  (240-450)  long 
by  470  (370-610)  wide;  seminal  receptacle 
large,  lying  immediately  postovarian,  525 
(200-700)  long  by  473  (320-600)  wide; 
Laurer's  canal  present,  opening  on  dorsal 
surface;  Mehlis'  gland  preovarian,  sinistral 
to  midline;  uterus  forming  transverse, 
intercecal  coils  between  the  ovary  and  ace- 


114 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


tabulum;  vitellaria  lateral,  extracecal,  con- 
sisting of  two  pairs  of  disjunct  bundles  on 
each  side,  each  pair  separate  at  level  of  the 
ovary;  four  vitelline  ducts  fuse  mesially  at 
the  level  of  the  ovary  to  form  a  vitelline 
reservoir.  Eggs  small,  28  (25-32)  long  by  12 
(11-14)  wide.  Excretory  pore  terminal  or 
slightly  subterminal;  excretory  vesicle 
sigmoid,  coursing  anteriorly  between  the 
testes  and  bifurcating  immediately  poste- 
rior to  the  seminal  receptacle. 
Discussion:  Reports  of  species  of  Amphi- 
merus  from  North  American  mammals 
have  almost  exclusively  been  A.  pseudofe- 
lineus  and  this  name  has  become  well  en- 
trenched in  veterinary  literature.  However, 
Nasir  and  Diaz  (1972)  synonymized  the 
following  species  with  A.  speciosus:  A. 
caudalitestis;  A.  guayaquilensis;  A.  inter- 
ruptus;  A.  minimus;  A.  neotropicalis;  A. 
parciovatus;  A.  pricei;  and  A.  pseudofe- 
lineus. 

Lumsden  and  Zischke  (1963)  reported 
A:  <;;himerus  interruptus  from  a  yellow- 
crowned  night  heron,  Nyctanassa  violacea. 
Their  measurements  fall  within  the  ranges 
we  recorded  and  the  specimen  figured  is 
remarkably  similar  to  ours,  indicating  that 
they  are  the  same  species.  Lumsden  and 
Zischke  also  noted  similarities  between 
their  specimens  and  the  description  of  A. 
speciosus.  These  observations  corroborate, 
in  part,  Nasir  and  Diaz's  (1972)  synony- 
mies and  further  indicate  the  ability  of 
these  organisms  to  live  in  both  avian  and 
mammaUan  hosts. 

A.  speciosus  has  been  reported  in  cats 
and  dogs  from  several  states  in  the  United 
States  (Rothenbacher  and  Lindquist, 
1963).  Chronic  morbidity  associated  with 
infection  includes  liver  and  biliary  cirrhosis 
and  pancreatitis.  Also,  Thatcher  (1970) 
commented  on  the  unassessed  possibility 
of  human  infection  with  this  species.  A. 
speciosus  was  collected  from  the  liver  and 
bile  ducts  of  one  of  four  domestic  cats  and 
two  of  37  raccoons  in  Louisiana.  The  rac- 
coon apparently  is  a  new  host  record  for 
this  species. 

Family  HETEROPHYIDAE 
(Leiper,  1909)  Odhner,  1914 


Cryptocotyle  concava  (Creplin,  1825) 

Luhe,  1899 

(Figure  4) 

Synonyms:    Distoma    concava    Creplin, 
1825;    Tocotrema  concava  Looss,    1899; 
Cryptocotyle  echinata  Linstow,  1878. 
Hosts:  Mustela  vison  Schreber. 
Location:  Small  intestine. 
Locality:  Belle  River,  Assumption  Parish, 
Louisiana. 

Deposition:  Univ.  Nebraska  State  Mus., 
Manter  Lab.  Coll.  No.  21370. 
Diagnosis  (based  on  ten  mature 
specimens):  Body  foliate,  904  (780-1050) 
long  by  612  (560-680)  wide.  Tegument  be- 
set with  small  spines.  Oral  sucker  terminal, 
47  (35-55)  long  by  54  (40-65)  wide;  aceta- 
bulum 41  (35-50)  in  diameter,  found  within 
the  genital  atrium  and  comprising  a  part  of 
the  acetabulogenital  apparatus;  acetabulo- 
genital  apparatus  67  (60-75)  long  by  91 
(70-125)  wide,  located  medially  and  equa- 
torially.  Prepharynx  10  (5-15)  long; 
pharynx  49  (40-55)  long  by  48  (45-60)  wide; 
esophagus  76  (65-100)  long;  paired  ceca 
extend  to  the  posterior  end  of  body  where 
they  turn  medially  just  posterior  to  the 
testes.  Testes  opposite,  distinctly  lobate, 
152  (125-175)  long  by  233  (210-250)  wide, 
located  in  posterior  end  of  body;  seminal 
vesicle  courses  from  testes  to  the  acetabu- 
logenital apparatus;  cirrus  pouch  absent. 
Ovary  wedge-shaped,  lobate,  93  (70-115) 
long  by  138  (100-175)  wide,  located  dextral 
to  the  midline,  between  the  ovary  and  right 
testis;  uterus  makes  3-4  intercecal  loops 
before  opening  into  the  acetabulogenital 
complex;  vitellaria  mostly  lateral,  com- 
mence behind  the  level  of  the  cecal  bifurca- 
tion and  extend  to  the  posterior  end  of 
body  where  they  meet  at  the  midline;  vitel- 
line reservoir  is  located  medially,  at  the 
level  of  the  seminal  vesicle.  Eggs  small, 
operculate,  36  (33-40)  long  by  15  (13-20) 
wide. 

Discussion:  Wootton  (1957)  first  reported 
Cryptocotyle  concava  from  North  Amer- 
ica and  elucidated  the  Ufe  cycle.  It  included 
an  operculate  snail,  Amnicola  longiqua,  in 
which  rediae  gave  rise  to  pleurolophocer- 
cous    cercariae;     these    penetrated    and 


No.  2 


Trematodes  of  Mammals 


115 


Figures  1-7.  1.  Alarm  alarioides  from  mink  and  river  otter.  2.  Alaria  marcianae  from  bobcat  and  raccoon  3 
Amphimerus speciosus  from  raccoon  and  the  domestic  cat.  4.  Cryptocotyle  concava  from  mink  5  Isthmio- 
phora  melis  from  raccoon  and  mink.  6.  Microphallus  opacus  from  raccoon  and  mink.  7.  Quinquesenalis 
quinqueserialis  from  muskrat.  Scales  in  micrometers. 


116 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


encysted  in  three-spined  sticklebacks,  Cas- 
ter osteus  aculeatus.  When  infected  fish 
were  fed  to  both  chicks  and  ducklings 
adult  worms  were  recovered.  Hoffman 
(1957)  found  metacercariae  of  C.  concava 
in  suckers,  Catostomus  commersoni,  and 
also  obtained  adults  from  experimentally 
infected  chicks. 

The  only  other  report  of  C.  concava 
from  North  America  was  that  of  Burrows 
and  Lillis  (1965)  who  collected  specimens 
from  a  dog  in  New  Jersey.  We  compared 
our  specimens  with  theirs  (USNM  Helm. 
Coll.  No.  60902)  and  find  no  differences 
between  them. 

Our  report  is  the  first  record  of  C 
concava  from  mink.  Its  occurrence  in  them 
is  not  surprising  due  to  the  prevalence  of 
fish  in  their  diet  and  the  lack  of  definitive 
host  specificity  common  in  heterophyids. 
Quite  possibly,  Louisiana  veterinarians 
may  encounter  eggs  of  this  trematode  in 
routine  stool  examination  of  pets.  In  addi- 
tion, the  possibility  of  human  infection  can 
not  be  overlooked  because  Cryptocotyle 
eggs  have  already  been  reported  from 
humans  elsewhere  (Babbot  et  al.,  1961). 

Family  ECHINOSTOMATIDAE 

(Looss,  1902)  Poche,  1926 

Isthmiophora  melis  (Schrank,  1788) 

Luhe,  1909 

(Figure  5) 

Synonyms:  Fasciola  putori  Gmelin,  1790; 

Fasciola    trigonocephala    Rud.,     1802; 

Euparyphium     melis    (Schrank,     1788) 

Railliet,     1919;     Echinocirrus    melis 

(Schrank,  1788)  Mendheim,  1943. 

Mendheim,  1943. 

Hosts:  Procyon  lotor  (Linn.)  and  Mustela 

vison  Schreber. 

Location:  Small  intestine. 

Locality:  Belle  River,  Assumption  Parish, 

Louisiana. 

Deposition:  Univ.  Nebraska  State  Mus., 

Manter  Lab.  Coll.  No.  21371. 

Diagnosis     (based     on     ten     mature 

specimens):     Body    lanceolate,     2450 

(2000-3500)  long  by  650  (520-700)  wide. 

Anterior  tegument  densely  covered  with 

spines  until  the  posterior  level  of  the  aceta- 


bulum, where  they  diminish  in  number 
towards  the  posterior  end  of  the  body. 
Head  collar  reniform,  bearing  27  spines; 
each  side  with  4  corner  spines,  59  (57-61) 
long  by  13  (12-14)  wide;  six  marginals  on 
each  side,  46  (43-48)  long  by  11  (9-13) 
wide;  and  a  double,  uninterrupted  row  of 
dorsal  spines  composed  of  four  oral  and 
three  aboral  spines,  40  (36-44)  long  by  1 1 
(8-12)  wide.  Acetabulum  large  relative  to 
the  oral  sucker,  380  (350-410)  long  by  385 
(350-430)  wide.  Prepharynx  not  discern- 
ible; pharynx  130  (110-160)  long  by  115 
(110-140)  wide;  esophagus  173  (110-210) 
long;  ceca  bifurcate  immediately  anterior 
to  the  cirrus  sac  and  extend  to  the  posterior 
end  of  the  body.  Testes  tandem,  irregular 
in  shape,  from  strongly  indented  to  com- 
pletely lobed,  posterior  testis  always  more 
indented  or  lobate  than  the  anterior  testis, 
both  testes  wider  than  long;  anterior  testis 
242  (200-310)  long  by  348  (310-370)  wide; 
posterior  testis  285  (220-410)  long  by  341 
(320-360)  wide;  cirrus  sac  ovate,  extending 
from  middle  of  the  acetabulum  to  just  pos- 
terior to  the  cecal  bifurcation,  265 
(220-300)  long  by  168  (130-200)  wide; 
seminal  vesicle  distinct;  cirrus  long,  coiled 
when  withdrawn,  beset  with  minute  spines. 
Ovary  spherical,  dextral  to  midline.  111 
(90-130)  long  by  1 14  (90-130)  wide,  located 
between  the  acetabulum  and  anterior  tes- 
tis; MehHs'  gland  broadly  oval  to 
reniform,  lying  immediately  in  front  of  the 
anterior  testis;  seminal  receptacle  absent; 
uterus  short,  with  3-5  intercecal  coils;  vitel- 
laria  extend  from  the  level  of  the  ovary  to 
the  posterior  end  of  body;  vitelline  reser- 
voir well  developed,  at  the  anterior  half  of 
the  anterior  testis.  Eggs  large,  operculate, 
97  (95-100)  long  by  53  (50-60)  wide.  Excre- 
tory pore  dorsal  and  subterminal. 
Discussion:  Dawes  (1946)  and  Skrjabin 
and  Bashkirova  (1956)  transferred  all  the 
species  of  Isthmiophora  to  the  genus 
Euparyphium,  however,  Yamaguti  (1971) 
retained  the  former  based  on:  (1)  body 
shape  (lanceolate  in  Isthmiophora  whereas 
Euparyphium  is  subcylindrical);  and  (2) 
shape  of  testes  (irregular  with  lateral  in- 
dentations  in   Isthmiophora   whereas   in 


No.  2 


Trematodes  of  Mammals 


117 


Euparyphium  they  are  longitudinally 
elongated).  Based  upon  a  comparative 
study  of  several  hundred  specimens  from 
Louisiana  mink  and  raccoons,  our  speci- 
mens agree  with  the  generic  diagnosis  of 
Isthmiophora  as  presented  by  Yamaguti, 

This  is  the  first  report  of  Isthmiophora 
melis  from  the  raccoon  and,  to  our  know- 
ledge, the  only  report  of  this  species  from 
North  America.  We  have  found  this  spe- 
cies in  the  small  intestine  of  six  of  37  rac- 
coons and  two  of  42  minks.  The  only  other 
echinostomes  found  in  raccoon  are 
Euparyphium  beaveri  reported  by 
Harkema  and  Miller  (1964)  and  Bufundo 
et  al.  (1980)  and  Echinostoma  revotutum 
which  was  regarded  as  an  aberrant  condi- 
tion (Larson  and  Scharf,  1975).  Because 
Euparyphium  beaveri  is  also  found  in 
minks  we  compared  the  type  material  de- 
posited by  Beaver  (1941)  to  our  specimens. 
We  find  they  are  very  similar  in  head  collar 
spination  and  body  anatomy,  but  that  they 
differ  strikingly  in  two  respects:  (1)  the 
range  in  size  of  our  specimens  (2000-3500) 
is  not  concordant  with  the  ranges  provided 
by  Beaver  (3860  -10500)  and  the  averages 
are  markedly  dissimilar  (2450  for  our 
material  to  6100  for  that  of  Beaver's);  and 
(2)  the  testes  in  our  specimens  are  broader 
than  long  with  either  deep  marginal  inden- 
tations or  completely  lobate,  whereas  that 
oi  Euparyphium  is  longitudinally  oval  with 
only  slight  evidence  of  indentations  in  the 
larger  specimens.  We  conclude  that  our 
material  is  distinct  from  Euparyphium 
beaveri. 

Lumsden  and  Zischke  (1961)  rediag- 
nosed  Euparyphium  beaveri  from  Louisi- 
ana minks.  A  close  inspection  of  their 
diagnosis  indicates  they  probably  were  not 
dealing  with  E.  beaveri  but  with  the  closely 
related  Baschkirovitrema  incrassatum.  At 
the  time  of  their  diagnosis  B.  incrassatum 
had  not  been  reported  from  North  Amer- 
ica. It  is  now  known  to  be  a  common  inha- 
bitant of  mustelids  from  the  Gulf  and 
Atlantic  coasts  (Sawyer,  1961;  Miller  and 
Harkema,  1964;  Fleming  et  al.,  1977; 
Shoop  and  Corkum,  1981a).  At  the  time 
we  diagnosed  B.  incrassatum  from  a  river 


otter  in  Louisiana  we  had  only  specimens 
from  a  single  otter.  We  now,  however, 
have  a  large  series  of  B.  incrassatum  from 
both  river  otter  and  mink  and  they  include 
the  ranges  of  both  our  previous  material 
and  that  given  by  Lumsden  and  Zischke 
(1961).  We,  therefore,  regard  Eupary- 
phium beaveri  of  Lumsden  and  Zischke, 
1961  conspecific  with  Baschkirovitrema 
incrassatum. 

Family  MICROPHALLIDAE 

Travassos,  1920 

Microphallus  opacus  (Ward,  1894) 

Ward,  1901 

(Figure  6) 

Synonyms:  Microphallus  ovatus  Osborn, 
1919. 

Hosts:  Procyon  lotor  (Linn.)  and  Mustela 
vison  Schreber. 
Location:  Small  intestine. 
Locality:  Belle  River,  Assumption  Parish, 
Louisiana. 

Deposition:  Univ.  Nebraska  State  Mus., 
Manter  Lab.  CoU.  No.  21372. 
Diagnosis  (based  on  ten  mature 
specimens):  Body  oval  to  pyriform,  1233 
(1160-1300)  long  by  664  (620-700)  wide. 
Tegument  spined  throughout.  Oral  sucker 
subterminal,  67  (60-70)  long  by  80  (75-90) 
wide;  acetabulum  86  (80-90)  long  by  91 
(90-100)  wide.  Prepharynx  60  (35-85)  long; 
pharynx  weak,  36  (35-40)  long  by  29 
(25-30)  wide;  esophagus  340  (275-400) 
long;  ceca  short,  rarely  extending  beyond 
the  seminal  vesicle,  occasionally  with  a 
single  sac.  Testes  two,  opposite,  190 
(150-230)  long  by  135  (75-190)  wide,  very 
often  the  testes  are  not  discernible  in 
gravid  specimens;  seminal  vesicle  saccular, 
preacetabular,  opening  into  the  genital 
atrium;  genital  atrium  lies  sinistral  to  the 
acetabulum,  62  (55-75)  long  by  74  (65-80) 
wide.  Ovary  spherical  to  oval  in  shape, 
dextral  to  midline,  150  (110-175)  long  by 
160  (130-205)  wide;  oviduct  sinistral  to 
ovary,  courses  posteriad  to  the  Mehlis' 
gland;  Mehlis'  gland  prominent,  on  the 
midline  of  the  body  between  the  two 
bundles  of  vitellaria;  uterus  makes  several 
loops  in  posterior  half  of  body  and  opens 


118 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


into  the  genital  atrium;  vitellaria  in  two 
symmetrical  clusters  of  spherical  follicles, 
located  in  the  posterior  Vi  of  body;  vitel- 
line ducts  fuse  in  the  middle  of  the  body  at 
the  level  of  the  Mehlis'  gland  to  form  a 
viteUine  reservoir.  Eggs  small,  numerous, 
25  (25-26)  long  by  13  (12-14)  wide. 
Excretory  vesicle  V-shaped,  extending  to 
the  anterior  level  of  the  vitellaria;  a  single 
collecting  duct  arises  from  each  side  of  the 
vesicle  and  courses  anteriad  to  the  level  of 
the  pharynx. 

Discussion:  Though  Microphallus  opacus 
is  generally  regarded  as  a  fish  parasite 
(Yamaguti,  1971)  it  has  been  experimental- 
ly established  in  various  reptilian  species  as 
well  as  opossum  and  raccoon  by  Rausch 
(1947)  and  in  white  mice  by  Sogandares- 
Bernal  (1965a).  Rausch  (1946)  also 
reported  it  from  a  naturally  infected  rac- 
coon from  Ohio  and  provided  a  brief  diag- 
nosis. Our  material  from  raccoon  and 
mink  agrees  well  with  that  description. 

Sogandares-Bernal  (1965a)  surveyed  the 
crayfish  parasites  in  Louisiana  and  found 
Cambarellus  puer  and  Procambarus  clarkii 
naturally  infected  with  the  metacercariae 
of  Microphallus  opacus.  He  noted  that 
snails  of  the  genus  Amnicola,  "probably 
Integra",  released  several  different  types  of 
microphallid  cercariae  at  his  study  site 
(Rosedale,  Louisiana),  one  of  which  he  be- 
lieved to  be  M.  opacus.  The  definitive  host 
at  that  time  was  unknown.  The  life-cycle 
of  M.  opacus  in  Louisiana  can  be  postu- 
lated using  Sogandares-Bernal's  report  and 
that  of  the  present  work  to  include  the  fol- 
lowing; an  amnicolid  snail  as  first  interme- 
diate host;  several  crayfish  species  as 
second  intermediate  hosts;  and  the  raccoon 
and  mink  as  definitive  hosts.  At  present, 
the  extent  to  which  M.  opacus  uses  fishes 
as  definitive  hosts  in  Louisiana  is  unas- 
sessed  as  it  has  yet  to  be  reported  from 
fishes  in  this  state. 

Family  PARAGONIMIDAE 

DoUfus,  1939 

Paragonimus  kellicotti  Ward,  1908 

Hosts:  Didelphis  virginiana  Kerr. 

Location:  Lungs. 

Locality:  Baton  Rouge,  East  Baton  Rouge 


Parish,  Louisiana. 

Deposition:  Univ.  Nebraska  State  Mus., 
Manter  Lab.  Coll.  No.  21394. 
Discussion:  We  have  recovered  three  ma- 
ture Paragonimus  kellicotti  from  the  lungs 
of  a  single  opossum.  We  have  not  figured 
or  diagnosed  P.  kellicotti  owing  to  the 
paucity  of  specimens  in  our  possession  and 
to  the  fact  that  our  specimens  are  similar  to 
those  described  by  Byrd  et  al.  (1942)  which 
came  from  the  lungs  of  a  Tennessee  opos- 
sum. 

Paragonimus  kellicotti  metacercariae 
were  reported  from  crayfish  in  Louisiana 
by  Ameel  (1934)  and  La  Rue  and  Ameel 
(1937).  Sogandares-Bernal  (1965b)  re- 
ported natural  infections  of  the  snail, 
Pomatiopsis  lapidaria,  with  Paragonimus 
kellicotti.  Since  those  accounts,  P.  kelli- 
cotti is  commonly  acknowledged  to  be  pre- 
sent in  Louisiana  although  neither  the 
adult  nor  the  definitive  host  have  been 
reported  from  this  state. 

That  the  infected  opossum  was  trapped 
in  residential  Baton  Rouge  is  epidemiolog- 
ically  significant.  The  location  was  an 
upper  middle  class  neighborhood  which 
borders  on  the  flood  plain  of  the  Mississip- 
pi River.  The  area  of  the  flood  plain  in 
heavily  treed,  with  numerous  bayous,  and 
low  lying  grounds  which  are  nearly  always 
water  laden.  This  scenario  is  a  classical 
nidus  capable  of  maintaining  all  of  the 
hosts  essential  to  the  life-cycle  of  P.  kelli- 
cotti and  has  the  potential  of  including 
man  into  the  life-cycle  owing  to  his  close 
proximity  and  crustacean  cuisine. 

Family  NOTOCOTYLIDAE 

Luhe,  1909 
Quinqueserialis  quinqueserialis 
(Barker  and  Laughhn,  1911) 
Harwood,  1939 
(Figure  7) 
Synonyms:    Notocotylus    quinqueserialis 
Barker  and  LaughUn,  1911;  Quinqueser- 
ialis   hassali    (Mcintosh    and    Mcintosh, 
1934)  Harwood,  1939;  Notocotylus  urban- 
ensis  of  Harrah,  1922. 
Hosts:  Ondatra  zibethica  (Linn.). 
Location:  Cecum. 
Locality:  Belle  River,  Assumption  Parish, 


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No.  2 


Trematodes  of  Mammals 


119 


Louisiana. 

Deposition:  Univ.  Nebraska  State  Mus., 
Manter  Lab.  Coll.  No.  21373. 
Diagnosis     (based     on     ten     mature 
specimens):  Body  elongate,  oval,  slightly 
attenuated    anteriorly,    3850    (3420-4150) 
long  by   1050  (960-1300)  at  the  greatest 
width.  Tegument  aspinous.  Ventral  sur- 
face with  five  longitudinal  rows  of  spher- 
ical glands.  Oral  sucker  subterminal,  335 
(320-350)  in  diameter;  acetabulum  absent. 
Pharynx  absent;  esophagus  short,  paired 
ceca  extend   to   posterior   end   of  body. 
Testes  opposite,  highly  branched,  in  pos- 
terior end  of  body,  513  (405-610)  long  by 
305     (260-390)    wide;     external    seminal 
vesicle  tubular,  coursing  anteriad  to  the 
base  of  the  cirrus  sac;  cirrus  sac  elongate, 
claviform,  1277  (1050-1500)  long  by  145 
(125-170)  at  the  greatest  width;  cirrus  often 
extruded  and  much  coiled,  densely  beset 
with   spines;   genital   pore   median,   near 
intestinal  bifurcation.  Ovary  deeply  lobed, 
intertesticular,  334  (300-390)  long  by  210 
(150-250)  wide;  Mehlis'  gland  immediately 
anterior   to   ovary;   uterus  comprised   of 
transverse  loops  which  may  extend  beyond 
the  ceca;  metraterm  distinct,  775  (700-900) 
long;  vitellaria  pretesticular,  in  two,  extra- 
cecal  bands.  Eggs  oval,  17  (16-18)  long  by 
8    (7-9)    wide,    without    polar    filaments. 
Excretory  system  not  observed. 
Discussion:  Penn  (1942)  examined  1,780 
muskrats  from  coastal  Louisiana  and  re- 
covered    the     trematodes     Nudacotyle 
novicia,    Echinochasmus  schwartzi,    and 
Paramonostomum  pseudalveatum.    Byrd 
and  Reiber  (1942)  examined  three  musk- 
rats    from    the    New    Orleans    area    and 
reported  E.  schwartzi  and  Phagicola  nana 
( =  P.  angrense).  Because  of  their  declining 
numbers,  we  were  unable  to  obtain  a  large 
series  of  muskrats  from  trappers,  but  we 
were  successful  in  obtaining  five  carcasses. 
Two  of  the  muskrat  harbored  hundreds  of 
Quinquesehalis    quinqueserialis    in    their 
ceca.  Although  this  species  is  considered  a 
ubiquitous  parasite  of  muskrats  in  North 
America,  this  is  the  first  report  of  it  from 
Louisiana.  Our  measurements  agree  well 
with  those  provided  by  Kinsella  (1971)  in 


his  study  of  intraspecific  variation  of  Q. 
quinserialis.  The  life-cycle  has  been  eluci- 
dated by  Herber  (1942)  and  includes  the 
freshwater  snail,  Gyraulis  parvus,  from 
which  monostome  cercariae  are  released 
and  encyst  on  vegetation.  The  muskrat 
becomes  infected  while  grazing  on  vegeta- 
tion containing  the  cysts. 

Incidentally,  one  muskrat  was  infected 
with  thousands  of  Hasstilesia  texensis  in 
the  cecum  (new  host  record).  We  have 
found  H.  texensis  in  all  of  the  swamp  rab- 
bits, Sylvilagus  aquaticus,  that  we  have 
examined  in  Louisiana.  As  all  of  the  speci- 
mens from  the  muskrat  were  gravid  and 
showed  neither  stunting  nor  any  anoma- 
lies, we  presume  that  the  muskrat  may 
serve  occasionally  as  a  normal,  definitive 
host  for  this  species. 

SUMMARY 

The  following  trematodes  were  collected 
from  hunter-trapped  mammals  in  the 
Atchafalya  basin  of  Louisiana  during  the 
winters  of  1981  and  1982:  Alaria  alarioides 
(Dubois,  1937)  Dubois,  1970;  Alaria  mar- 
cianae  (La  Rue,  1917)  Walton,  1949; 
Alaria  mustelae  Bosma,  1931;  Amphi- 
merus  speciosus  (Stiles  and  Hassal,  1896) 
Barker,    1911;    Baschkirovitrema    incras- 

satum  (Dies.  1850)  Skrjabin,  1944;  Brachy- 
laima  virginiana  Dickerson,  1930; 
Carneophallus  basodactylophallus  Bridg- 
man,  1969;  Cryptocotyle  concava 
(Creplin,  1825)  Luhe,  1899;  Fibricola  cra- 
tera  (Barker  and  Noll,  1915)  Dubois,  1932; 
Fibricola  lucida  (La  Rue  and  Bosma,  1927) 
Dubois  and  Rausch,  1950;  Gyrosoma  sin- 
gulare  Byrd,  Bogitsh,  and  Maples,  1961; 
Hasstilesia  texensis  Chandler,  1929; 
Heterobilharzia  americana  Price,  1929; 
Isthmiophora  metis  (Schrank,  1788)  Luhe, 
1909;  Linstowiella  szidati  (Anderson, 
1944)  Anderson  and  Cable,  1950;  Mari- 
treminoides  nettae  (Gower,  1938)  Rankin, 
1939;  Microphallus  opacus  (Ward,  1894) 
Ward,  1901;  Paragonimus  kellicotti  Ward, 
1908;  Pharyngostomoides  procyonis 
Harkema,  1942;  Quinqueserialis  quinque- 
serialis (Barker  and  Laughlin,  1911)  Har- 
wood,     1939;     Phopalias    macracanthus 


120 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


(Chandler,  1932;  and  Sellacotyle  vitellosa 
Sogandares-Bernal,  1961. 

Adult  trematodes  reported  from  Louisi- 
ana for  the  first  time  are:  Alaria  alarioides, 
A.  marcianae,  Amphimerus  speciosus, 
Cryptocotyle  concava,  Isthmiophora 
melis,  Mircophallus  opacus,  Paragonimus 
kellicotti,  and  Quinqueserialis  quin- 
queserialis. 

New  host  records  include  Heterobilhar- 
zia  americana,  Cryptocotyle  concava,  and 
Maritreminoides  nettae  from  mink;  Alaria 
marcianae,  Amphimerus  speciosus,  and 
Linstowiella  szidati  from  raccoon;  and 
Hasstilesia  texensis  from  muskrat. 

Natural  infections  of  adult  Alaria  mar- 
cianae were  found  only  in  juvenile  rac- 
coons. This  substantiates  previous  experi- 
mental work  which  demonstrated  that 
adult  raccoon  are  unsuitable  for  the 
maturation  of  this  trematode.  The  larvae, 
however,  are  able  to  employ  the  adult  rac- 
coon as  a  paratenic  host  where  they  remain 
undifferentiated  in  the  subcutaneous  fat. 

Amphimerus  speciosus  is  a  well  known 
pathogen  of  dogs  and  cats  in  North 
America,  being  herein  reported  from  a 
domestic  cat  and  a  raccoon.  Synonymiza- 
tion  of  the  better  known  A .  pseudofelineus 
with  A.  speciosus  is  corroborated  by  our 
observations. 

Whether  Isthmiophora  is  distinct  from 
Euparyphium  has  been  debated  by  several 
authors.  We  place  our  specimens  in  the 
genus  Isthmiophora  on  the  basis  of  body 
shape  and  testicular  morphology.  We  com- 
pared our  specimens  to  those  of  Eupary- 
phium beaveri  and  conclude  they  are  dis- 
tinct. This  is  the  first  report  of  /.  melis 
from  a  raccoon  and,  to  our  knowledge,  the 
only  report  of  this  species  from  North 
America.  We  consider  Euparyphium  bea- 
veri of  Lumsden  and  Zischke,  1961  to  be  a 
synonym  of  Baschkirovitrema  incras- 
satum. 

Microphallus  opacus  is  a  common  para- 
site in  the  mink  and  raccoon  in  Louisiana. 
Sogandares-Bernal  (1965a)  stated  that  the 
aquatic  snail,  Amnicola,  probably  served 
as  first  intermediate  host  and  that  several 
species  of  crayfish  served  as  second  inter- 


mediate hosts.  Therefore,  a  hypothetical 
life-cycle  of  M.  opacus  from  Louisiana  can 
be  proposed:  the  first  intermediate  host  is 
the  aquatic  snail,  Amnicola;  several  cray- 
fishes serve  as  second  intermediate;  and 
the  raccoon  and  mink  are  definitive  hosts. 
Fishes  have  yet  to  be  reported  with  M. 
opacus  from  Louisiana. 

The  first  and  second  intermediate  hosts, 
as  well  as  the  larval  stages,  of  Paragonimus 
kellicotti  have  been  previously  reported 
from  Louisiana.  However,  this  is  the  first 
report  from  this  state  of  the  adult  fluke  in 
a  naturally  infected  definitive  host,  the 
opossum.  The  locality  of  the  infection  is 
noteworthy  in  that  it  was  found  in  an 
upper  middle  class  Baton  Rouge  residential 
area. 

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December  15,  1982 


COMPARATIVE  VISCERAL  TOPOGRAPHY  OF  THE 

NEW  WORLD  SNAKE  TRIBE 

THAMNOPHIINI  (COLUBRIDAE,  NATRICINAE) 

NITA  J.  ROSSMAN  and  DOUGLAS  A.  ROSSMAN 

Museum  of  Zoology,  Louisiana  State  University 
Baton  Rouge,  Louisiana  70893 

NANCY  K.  KEITH 

Dept.  of  Experimental  Statistics,  Louisiana  State  University 
Baton  Rouge,  Louisiana  70893 


Abstract 

The  positions  and  lengths  of  a  variety  of  visceral 
organs  in  631  preserved  adult  thamnophiine  snakes 
were  determined  in  terms  of  ventral  scute  number  and 
converted  into  a  per  cent  of  total  ventral  number;  a 
mean  was  calculated  for  each  taxon  to  allow  compar- 
ison with  other  taxa.  Dice-Leraas  diagrams  were  then 
constructed  for  the  following  organ  positions  and 
lengths:  posterior  end  of  heart,  anterior  and  posterior 
ends  of  Uver,  posterior  end  of  pancreas,  anterior  and 
posterior  ends  of  right  and  left  kidney,  Uver  length, 
right  and  left  kidney  lengths,  heart-liver  interspace, 
and  kidney  overlap.  Sexual  dimorphism  is  apparent  in 
many  of  the  characters  examined.  Apparently  corre- 
lated with  their  need  for  space  to  accommodate 
developing  young,  females  tend  to  have  their  anterior 
and  midbody  organs  placed  more  anteriorly  and  their 
kidneys  more  posteriorly  than  those  in  males. 

Stepwise  discriminant  analysis  was  performed  on 
the  following  four  variables  in  male  thamnophiine 
snakes:  posterior  end  of  heart,  anterior  end  of  right 
kidney,  posterior  end  of  left  kidney,  and  kidney 
overlap.  The  294  specimens  represented  11  groups  — 
7  genera  plus  Ruthven's  four  species  groups  of 
Thamnophis.  Two  of  four  linear  discriminant 
functions  were  retained  as  they  explain  83.2 l<^o  of  the 
relative  variation.  Function  1  is  generally  an  anterior 
end  of  right  kidney  dimension,  and  function  2  is  a 
kidney  overlap  and  posterior  end  of  heart  dimension. 
More  than  66%  of  the  specimens  were  correctly  classi- 
fied by  use  of  the  model.  All  groups  except  Clonophis 
could  be  classified  with  greater  success  than  the  21% 
prior  probability  obtained  by  placing  them  all  in  the 


Elegans  group  of  Thamnophis,  the  numerically 
largest  sample.  The  discriminant  analysis  was  able  to 
distinguish  among  the  seven  genera  (as  well  as  among 
Ruthven's  four  species  groups  of  Thamnophis)  at  the 
0.05  level  except  that  Clonophis  and  Tropidoclonion 
could  not  be  distinguished  from  each  other. 

Although  visceral  topographic  data  alone  do  not 
clearly  delimit  thamnophiine  genera  nor  establish 
inter-  or  intrageneric  relationships,  some  trends  are 
apparent  that  serve  to  support  taxonomic  conclusions 
based  on  other  kinds  of  characters.  Clonophis  and 
Regina  can  be  distinguished  from  Nerodia,  in  which 
genus  they  were  formerly  included.  Thamnophis  (less 
proximus  and  sauritus)  can  also  be  distinguished  from 
Nerodia  (less  erythrogaster  and  valida).  The  Sauritus 
group  of  Thamnophis  differs  markedly  from  the 
other  three  species  groups  established  by  Ruthven  in 
most  visceral  topographic  features.  The  ribbon  snakes 
(Sauritus  group)  frequently  tend  to  have  a  posterior 
displacement  of  organs,  a  condition  often  occurring 
also  in  the  short,  semifossorial  genera  (Clonophis, 
Seminatrix,  Storeria,  Tropidoclonion,  Virginia).  One 
unique  feature  shared  by  all  of  the  semifossorial 
genera  is  the  possession  of  a  relatively  long  liver. 

Introduction 

The  technique  of  determining  snake  vis- 
ceral topography  using  ventral  scutes  as  re- 
ference points  has  received  little  attention 
since  its  introduction  by  Thompson 
seventy  years  ago.  Although  a  moderate 
amount  amount  of  descriptive  anatomical 
work  has  appeared  in  print,  very  little  has 


EDITORIAL  COMMITTEE  FOR  THIS  PAPER: 

DR.  SAMUEL  B.  McDOWELL,  Professor  of  Zoology,  Rutgers  University, 
Newark,  New  Jersey  07102 

DR.  JAMES  S.  ROGERS,  Associate  Professor  of  Biology,  University  of  New 
Orleans,  New  Orleans,  Louisiana  70122 

DR.  ROBERT  A.  THOMAS,  Director,  Louisiana  Nature  Center,  New  Orleans, 
Louisiana  70127 

123 


124 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


been  done  of  a  comparative  nature  that 
might  be  of  taxonomic  value,  and  none 
using  discriminant  analysis.  The  present 
study  was  undertaken  to  investigate  the 
possible  taxonomic  significance  of  visceral 
topography  in  the  tribe  Thamnophiini  of 
the  colubrid  subfamily  Natricinae. 

Beddard  (1908,  1909)  characterized  the 
position  of  visceral  organs  in  three  genera 
of  boid  snakes  in  terms  of  the  distance 
from  the  snout  to  the  organ.  He  also  mea- 
sured organ  length  and  the  distance  be- 
tween organs.  Beddard  was  convinced  that 
the  position  of  viscera  within  the  body  of 
snakes  generally  had  systematic  impor- 
tance. Subsequent  authors  who  also  used 
distance  measurements  were  Atwood 
(1916,  1918),  Bergman  (1941  et  seq.),  and 
Brongersma  (1951,  1957  a  &  b).  Bergman 
expressed  the  organ  positions  and  lengths 
as  a  per  cent  of  snout-vent  length,  and 
both  he  and  Brongersma  also  presented 
their  data  diagrammatically. 

Thompson  (1913a  &  b,  1914)  was  the 
first  to  relate  the  position  of  the  various 
visceral  organs  to  the  ventral  scutes  in  an 
attempt  to  provide  a  simple,  yet  objective, 
technique  for  stating  the  location  of  the 
organs.  The  position  of  an  organ  was 
expressed  as  a  percentage  of  the  total  num- 
ber of  ventrals  in  order  to  compensate  for 
individual,  sexual,  and  geographic  varia- 
tion in  ventral  number.  This  technique  has 
been  utilized  subsequently  only  by  Thorpe 
(1975),  Underwood  (1976),  and  Rasmussen 
(1979).  Thorpe  determined  the  midpoint  of 
an  organ  rather  than  the  anterior  and 
posterior  ends,  so  his  data  are  not 
comparable  to  ours  or  to  those  of  other 
authors.  Inasmuch  as  one  has  to  ascertain 
the  anterior  and  posterior  ends  in  order  to 
determine  the  midpoint,  the  latter  would 
appear  to  be  an  unnecessary  complication 
and  if  used  alone  it  also  results  in  a  loss  of 
information. 

Garrigues  (1962),  Bogert  (1968),  Collins 
and  Carpenter  (1970),  and  Frenkel  and 
Kochva  (1970)  also  gave  organ  positions 
and  lengths  in  terms  of  ventral  number, 
but  they  did  not  express  their  data  as  a  per 
cent  of  total  ventrals.  Also,  by  lumping  his 


samples  for  each  species,  Garrigues  failed 
to  take  sexual  dimorphism  into  account. 

Valle  (1944-45),  Bragdon  (1953),  and 
Camazine  et  al.  (1981)  used  ventral 
number  to  pinpoint  the  location  of  various 
posterior  organs  so  that  surgical  proce- 
dures could  be  carried  out  using  the 
smallest  incisions  possible.  In  each  case, 
the  investigator  counted  ventral  scutes 
from  the  vent  forward. 

Materials  and  Methods 

We  examined  63 1  preserved  adult  speci- 
mens, representing  8  thamnophiine  genera 
(only  Adelophis  was  omitted  because  of  its 
rarity)  and  35  species  (4  being  represented 
by  two  subspecies  or  populations).  Large 
subadults  were  used  only  if  their  data  fitted 
into  the  range  of  variation  for  the  taxon 
under  consideration.  Juveniles  were 
rejected  because  their  values  tend  to  lie 
outside  the  normal  range  of  variation  in 
adults  (see  Bergman,  1958a,  1961b). 

Only  nongravid  females  or  those  with 
undeveloped  eggs  were  used  because  of  the 
distortion  caused  by  developing  embryos 
(also  noted  by  Bergman,  1961a;  CoUins 
and  Carpenter,  1970;  Thorpe,  1975).  Be- 
cause females  tend  to  have  their  anterior 
organs  situated  more  anteriorly  and  their 
kidneys  more  posteriorly  than  those  of 
males,  each  sex  was  considered  separately 
(see  the  Sexual  Dimorphism  section  for 
further  discussion). 

Using  the  Dowling  method  for  counting 
ventral  scutes,  we  inserted  insect  pins  in  the 
20th  scute  and  in  every  15th  scute  there- 
after. Several  midventral  slits  were  made  to 
expose  the  organs  being  studied.  The  ven- 
tral scute  numbers  at  the  anterior  and 
posterior  ends  of  each  organ  were  re- 
corded; to  faciUtate  inter-  and  intraspecific 
comparisons,  a  percentage  was  calculated 
by  dividing  the  scute  number  by  the  total 
number  of  ventrals.  The  following  organs 
were  considered  where  possible:  heart, 
liver,  gall  bladder,  pancreas,  right  and  left 
kidneys.  Lungs,  thyroid,  spleen,  and 
adrenals  were  not  considered  because  they 
were  difficult  to  locate  in  many  specimens. 
Testes  and  ovaries  were  not  considered 


No.  2 


Visceral  Topography  of  Snakes 


125 


because  of  the  varying  size  depending  on 
whether  the  specimens  were  in  a  breeding 
or  non-breeding  state  (see  Matthews  and 
Marshall,  1956;  Manna  and  Sircar,  1978). 
Organ  lengths,  expressed  as  the  total 
number  of  ventral  scales  covered,  were 
also  recorded  and  treated  as  a  percentage 
of  total  number  of  ventrals.  The  following 
distances  were  measured  and  expressed  in 
the  same  manner:  posterior  end  of  heart  to 
anterior  end  of  liver,  posterior  end  of  liver 
to  anterior  end  of  gall  bladder,  distance 
between  or  overlap  of  the  right  and  left 
kidneys.  On  museum  material  other  than 
that  in  the  Louisiana  State  University 
Museum  of  Zoology  (LSUMZ),  only  the 
heart,  anterior  end  of  liver,  and  kidneys 
were  examined  in  order  to  minimize  the 
number  of  incisions.  Preliminary  data  on 
LSUMZ  specimens  had  indicated  that 
these  organs  were  the  most  relevant  to  the 
study. 

The  statistics  used  in  the  Inter-  and 
Intrageneric  Comparisons  section  con- 
sisted of  calculating  the  mean,  standard 
deviation,  and  standard  error  of  the  mean 
for  each  sex  of  each  taxon,  then  construct- 
ing graphs  by  the  Dice-Leraas  method  as 
discussed  in  Simpson  et  al.  (1960).  This 
method  presents  a  graphic  representation 
of  differences  between  populations,  and 
the  results  appear  in  Figs.  1-19.  The  95% 
confidence  interval  of  the  mean  was  deter- 
mined by  dividing  the  standard  deviation 
by  the  square  root  of  the  sample  size  and 
multiplying  this  figure  by  a  value  from  the 
Student's  t-test  table  using  n-1  degrees  of 
freedom  (Runyon  and  Harber,  1968).  Be- 
cause of  the  very  large  confidence  interval 
generated  by  a  sample  of  two  specimens, 
we  constructed  a  Dice-Leraas  diagram  only 
in  those  cases  where  we  had  a  minimum 
sample  of  three  specimens  of  the  same  sex. 
The  confidence  interval  results  in  a  plus  or 
minus  figure  relative  to  the  mean.  Where  a 
determination  of  the  statistical  significance 
of  the  differences  between  means  could  not 
be  obtained  from  this  graphic  representa- 
tion (using  the  three  general  rules  on  p.  353 
in  Simpson  et  al.,  1960),  then  a  Student's 
t-test  was  used.  When  data  are  stated  as 


being  significantly  different  in  this  paper, 
it  refers  to  the  fact  that  the  differences  are 
significant  at  the  p<  .05  level. 

To  minimize  the  possible  effects  of  geo- 
graphic variation,  we  attempted  to  sample 
populations  from  as  restricted  an  area  as 
possible.  In  four  instances  (Thamnophis 
couchii,  T.  elegans,  T.  sirtalis,  Tropido- 
clonion  lineatum)  we  treated  different  sub- 
species or  geographically  distant  popula- 
tions as  separate  taxon  samples.  Because 
enough  male  and  female  Thamnophis 
eques  could  not  be  obtained  from  one  geo- 
graphic area,  we  used  females  of  T.  e. 
megalops  and  males  of  T.  e.  virgatenuis. 

Due  to  the  existence  of  sexual  di- 
morphism, data  for  males  and  females 
could  not  be  combined  for  discriminant 
analysis.  We  chose  to  restrict  the  discri- 
minant analysis  to  the  data  for  males;  only 
a  relatively  few  confidence  intervals  could 
be  shown  for  females  on  the  Dice-Leraas 
diagrams  because  many  of  the  confidence 
intervals  exceeded  the  ranges  of  variation. 
Only  those  specimens  that  had  data  avail- 
able for  all  characters  were  used.  Six 
variables  (posterior  end  of  heart,  anterior 
and  posterior  ends  of  right  kidney,  ante- 
rior and  posterior  ends  of  left  kidney,  and 
kidney  overlap)  were  first  run  after  the 
values  were  standardized  at  the  mean  to 
allow  for  comparisons.  Because  the  poste- 
rior end  of  the  right  kidney  and  the  ante- 
rior end  of  the  left  kidney  were  signifi- 
cantly correlated,  those  characters  were 
eliminated  to  obtain  a  four-variable 
explanatory  and  predictive  model.  The 
posterior  end  of  the  right  kidney  and 
anterior  end  of  the  left  kidney  values  are 
reflected  in  the  kidney  overlap  figures. 

Because  of  the  relatively  small  number 
of  specimens  in  each  sample,  the  294  speci- 
mens were  placed  in  the  following  eleven 
groups  to  achieve  greater  statistical  signi- 
ficance of  the  discriminant  values: 

1 .  Clonophis  kirtlandii  —  6  specimens 

2.  Nerodia  (cyclopion,  erythrogaster, 
fasciata,  rhombifera,  sipedon,  valida) 
—  51 

3.  Regina  {alleni,  grahamii,  rigida,  sep- 
temvittata)  —  24 


126 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


4.  Seminatrix  pygaea  —  8 

5.  Storeria    {dekayi,     occipitomaculatd) 

—  15 

Thamnophis  (groups  from  Ruthven, 
1908) 

6.  Sauritus  group   (proximus,   sauritus) 

—  14 

7.  Radix  group  [brachystoma,^  butleri, 
eques  {  =  megalops  in  Ruthven),  mar- 
cianus,  radix]  —  44 

8.  Elegans  group  [couchii  couchii,^  c. 
hydrophilus,'  elegans  terrestris,^  e. 
vagrans,^  melanogaster,  nigronucha- 
lis,^  ordinoides,  rufipunctatus  {  =  an- 
gustirostris  in  Ruthven),  scalaris]  — 
63 

9.  Sirtalis  group  [chrysocephalus,^  cyr- 
topsis  ('eques  in  Ruthven),  godmani,^ 
sirtalis  fitchi, '  s.  sirtalis]  —  47 

10.  Tropidoclonion  lineatum  (Nebraska, 
New  Mexico,  Texas)  —  5 

11.  Virginia  {yaleriae,  striatula)  —  17 
Prior  probabilities  of  group  membership 

were  calculated  by  dividing  the  number  in 
any  group  by  the  total  number  in  the 
study.  These  prior  probabilities  are  used  in 
classifying  the  specimens  with  the  discri- 
minant model. 

Sexual  Dimorphism 

Details  on  sexual  dimorphism  in  this 
study  appear  in  Tables  I  and  II  and  in  Figs. 
1-19.  A  comparison  of  sexual  dimorphism 
data  from  this  study  with  other  studies 
appear   in  Table  III. 

Anterior  organ  positions 

The  posterior  end  of  the  heart  and  the 
anterior  end  of  the  liver  in  males  are 
located  posteriorly  to  those  positions  in  fe- 
males in  11^0  and  81  %  of  the  taxa,  respec- 
tively. Male  Clonophis,  Seminatrix,  Stor- 
eria, Tropidoclonion  (for  heart  only),  and 
Virginia  have  the  posterior  end  of  the  heart 
and  the  anterior  end  of  the  liver  located 
posteriorly  to  those  positions  in  females  in 
all  species.  In  Nerodia,  Regina,  and 
Thamnophis  there  is  interspecific  variabi- 
Uty  in  both  features.  Male  Thamnophis 

'taxon  described  since  Ruthven  (1908) 
Haxon  not  recognized  by  Ruthven  (1908) 


have  the  posterior  end  of  the  heart  situated 
posteriorly  to  that  of  females  in  74%  of  the 
taxa;  male  Nerodia  in  67<^o;  male  Regina  in 
67% .  The  anterior  end  of  the  liver  in  males 
lies  posteriorly  to  that  of  females  in  8O070 
of  the  species  of  Nerodia,  73%  of  the  taxa 
of  Thamnophis,  and  in  the  only  species  of 
Regina  for  which  data  are  available. 

Midbody  organ  positions 

Sexual  dimorphism  of  the  midbody 
organ  positions  is  not  pronounced.  Males 
have  the  posterior  end  of  the  liver  located 
posteriorly  to  that  of  females  in  50%  of  the 
taxa,  the  posterior  end  of  the  gall  bladder 
posteriorly  to  that  of  females  in  69%.  The 
posterior  end  of  the  liver  is  more  poste- 
riorly placed  in  males  in  60%  of  the  species 
of  Nerodia,  both  species  of  Storeria,  and 
in  the  one  species  of  Virginia  examined. 
Males  have  the  posterior  end  of  the  gall 
bladder  located  more  posteriorly  than  do 
females  in  60%  of  the  species  of  Nerodia, 
70%  of  the  taxa  of  Thamnophis,  and  in 
the  one  species  of  Storeria  examined. 

Posterior  organ  positions 

In  contrast  to  most  of  the  preceding 
characters,  the  kidneys  exhibit  marked 
sexual  dimorphism  in  many  of  their  fea- 
tures. The  anterior  ends  of  the  right  and 
left  kidneys  in  males  are  anterior  to  those 
of  females  in  all  taxa,  as  are  the  posterior 
ends  of  the  right  and  left  kidneys  in  86% 
and  73%  of  the  taxa,  respectively.  The 
posterior  end  of  the  right  kidney  in  males  is 
situated  anteriorly  to  that  of  females  in  all 
species  of  Regina,  Seminatrix,  Tropido- 
clonion, and  Virginia,  and  95%  of  the  taxa 
of  Thamnophis.  In  Nerodia  the  posterior 
end  of  the  right  kidney  of  males  is  situated 
posteriorly  to  that  of  females  in  67%  of  the 
species.  In  both  species  of  Storeria  and  in 
half  the  species  of  Nerodia,  the  posterior 
end  of  the  left  kidney  of  males  is  situated 
posteriorly  to  that  of  females.  The  poste- 
rior end  of  the  left  kidney  of  males  is 
anterior  to  that  of  females  in  95%  of  the 
taxa  of  Thamnophis,  67%  of  the  species  of 
Regina,  one  population  of  Tropidoclo- 
nion, and  in  both  species  of  Virginia. 


No.  2 


Visceral  Topography  of  Snakes 


127 


Table  1.  Sexual  dimorphism  in  certain  thamnophiine  snakes. 


Character 


post 
ant . 
post 
post 
ant . 
post 
ant . 
post 


,  heart^ 

liver 
.  liver 
i    gall  bladder 

r.  kidney 
•  r.  kidney 

1 .  kidney 
1.  kidney 


35 
27 
18 
16 
37 
37 
37 
37 


Position  in  eft? Position  in  99 

posterior  to  posterior  to 

that  in  99  (or  that  in  cfcC  (or 

o"  organ  longer')  9  organ  longer") 

7  77.  17% 

81%  15% 

50%  44% 

69%  31% 

0%  100% 

14%  86% 

0%  100% 

24%  73% 


liver  length  17 

gall  bladder  length  16 

r.  kidney  length  3  7 

1.  kidney  length  37 
post,  heart-ant. 

liver  interspace  26 

kidney  overlap  3  7 


76% 

69% 

5% 

0% 

2  7% 
13% 


-a'  significantly 
cfd=99       different  (pS05) 
from  9? 


6% 
4% 
6% 
0% 

0% 
0% 

0% 
3% 


6% 
0% 
3% 
0% 

0% 
3% 


:>7% 

41% 
22% 
25% 
81% 
49% 
86% 
32% 


18% 

6% 

46% 

73% 

12% 
22% 


means  of  the  taxa  were  used  in  computing  the  figures  in  this  table 


Organ  lengths  and  interspaces 

The  liver  and  gall  bladder  of  females  are 
longer  than  those  of  males  in  76%  and 
69%  of  the  taxa,  respectively.  However, 
the  right  and  left  kidneys  of  males  are 
longer  than  those  of  females  in  92%  and 
100%  of  the  taxa,  respectively,  probably 
due  to  the  presence  of  a  hypertrophied 
sexual  segment  in  males  (Matthews  and 
Marshall,  1956;  Prasad  and  Reddy,  1972). 
The  male  heart-liver  interspace  is  longer 
than  that  of  females  in  73%  of  the  taxa,  as 
is  the  male  kidney  overlap  in  84%  of  the 
taxa.  In  80%  of  the  taxa  of  both  Nerodia 
and  Thamnophis,  males  have  a  shorter 
liver  than  do  females;  the  hver  is  also 
shorter  in  male  Storeria  dekayi  (in  S.  occi- 
pitomaculata  the  Hver  shows  no  sexual 
dimorphism).  Data  were  available  for  both 
sexes  in  only  one  species  of  Regina  and  one 
of  Virginia.  Males  have  a  shorter  gall  blad- 
der than  do  females  in  80%  of  the  species 
of  Nerodia,  in  70%  of  the  taxa  of  Tham- 
nophis, and  in  Storeria  dekayi.  Males  of 
Clonophis,  Nerodia,  Regina,  Seminatrix, 
Storeria,  Tropidoclonion,  and  Virginia 
have  longer  right  and  left  kidneys  than  do 
females.  In  all  taxa  of  Thamnophis,  males 
have  a  longer  left  kidney  than  do  females; 
in  86%  of  those  taxa,  males  also  have  a 
longer    right    kidney.    In    all    species    of 


Clonophis,  Regina,  Seminatrix,  Storeria, 
and  Virginia,  males  have  a  longer  heart- 
Hver  interspace  than  do  females,  as  is  the 
case  for  60%  of  the  species  of  Nerodia  and 
64%  of  the  taxa  of  Thamnophis.  In  all 
species  of  Clonophis,  Nerodia,  Regina, 
Seminatrix,  Tropidoclonion,  and  Virginia, 
males  have  a  more  extensive  kidney 
overlap  than  do  females,  as  is  the  case  for 
76%  of  the  taxa  of  Thamnophis. 
Asymmetry  of  kidney  lengths 

In  76%  of  the  taxa,  females  have  the 
right  kidney  longer  than  the  left  (24% 
differ  significantly).  On  the  other  hand, 
males  have  the  left  kidney  longer  than  the 
right  in  55%  of  the  taxa  (5%  differ  signi- 
ficantly). In  all  species  of  Nerodia,  the 
right  kidney  is  the  longer  one  in  both  sexes. 
Summary  and  conclusions 

In  general,  the  anterior  and  midbody 
organs  are  placed  more  posteriorly  in 
males  than  in  females,  whereas  the  kidneys 
of  males  are  positioned  more  anteriorly 
than  those  of  females.  This  more  anterior 
positioning  of  the  anterior  organs  and 
more  posterior  positioning  of  the  posterior 
organs  in  females  would  allow  greater 
space  for  the  developing  young. 

Inter-  and  Intrageneric  Comparisons 
To  facilitate  comparisons,  each  set  of 


128 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


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oooooooooooooooooo  oo^ 

o 
o 

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tN   O 

o  o 
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1 
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1    1    1 

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llll                   cNinl                11^          III           1 
1     1     1     itntocoooitotoi     lotoi     1     igl 

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1     1     1     Itotototooitotoi     lOOl     1      1^' 
1      1      1      IZZZZ      -IZZl      l.-^-l      1       IZl 

o*.boo^       •bo            bo                 II 

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to  o  to  to  O    1 
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CO  -^  o  -rf  -pi  CO  CO  CO      -vi  c  tji  3  -tJ  g       s;  js  Y  ;5^  -^ 
^TOco^^oeetoa-rico^vH  tx-f*  «  5  5 

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No.  2 


Visceral  Topography  of  Snakes 


129 


Table  III.   Data  on  sexual  dimorphism  reported  In  the  literature.   X  indicates  that  the  organ  is  longer  or 
located  more  caudally  in  sex  indicated;   ND  that  there  is  no  appreciable  dimorphism. 


^ 

^ 

^"■^"--^^^                   Character 

^ 

u 

^ 

c 

c 

Xi 

^   > 

>. 

r. 

•o  £ 

TJ    X 

4J 

■M     U 

Taxon                               ^^■^'^^ 

M    OO 

J^     00 

Ul     > 

tfl    T3 

*j  -a 

Ul    T3 

CU    C 

>   c 

c 

O    tu 

C    -H 

c  ■-< 

0  ^ 

C    -H 

O'-H 

<u  fu 

D.  j: 

«  M 

D.^ 

O.  00 

n  M 

O.M 

(D  M 

C1.M 

^    i-H 

u  ^ 

^    -H 

<S    9 

a    9 

d    9 

d     d 

d     9 

d    9 

d    9 

d    9 

d     9 

d    9 

rf    9 

d    9 

Colubrldae,    Natrlcinae                        ^ 

Natrix   (=Phabdophis)   ahrysarga 

X 

X 

N.    (=Rhabdcphis)   subminiata^ 

X 

X 

X 

X 

X 

X 

X 

X 

X 

It.    ( =Sinonatrix )    trianguligera 
N.    (=Xenochi'ophis)   vittata 

X 

X 

X 

X 

X 

Colubrldae                                                         ^ 

Ablabee    (=aongiilosoma)   baliodeira 

X 

X 

X 

X 

X 

X 

X 

X 

ND 

X 

X 

Calamapia  multipunotata^ 

X 

X 

X 

Coluber  melanums    (=Elapke 

flavolineata)^ 

X 

X 

X 

X 

X 

X 

X 

Dendrophis    (=Dendrelaphis)  pictus 

ND 

ND 

X 

X 

Colubrldae,    Homalopslnae 

Enhydris  plwnbea° 

X 

X 

ND 

ND 

X 

X 

X 

X 

X 

X 

Fordonia   leucobatia 

X 

X 

X 

Homalopsis  bucaata^ 

ND 

ND 

TO 

Hypsirhina   (=Enhydris)  altemans 

X 

X 

X 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

Acrochordldae 

Acroahordus  gi-anulatus 
A.   Javaniaus'-^ 

X 

X 

X 

X 

X 

X 

X 

ND 

ND 

ND 

X 

X 

X 

X 

X 

X 

X 

ND 

Anllildae 

Cylindrophis  rufus 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

Boldae 

Xenopeltis  icniaolor 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

ND 

Elapldae 

Hydrophis  fasaiatus 

X 

Vlperidae 

AgkistTodon  pisaivorus 
Tpimeresurus  gramineus 

X 

ND 

ND 

ND 

ND 

X 

X 

X 

X 

X 

X 

X 

X 

X 

Bergman  (1959a);   Bergman  (1956b);   Bergman  (1950);   Bergman  (1963);   Bergman  (1965);   Bergman  (1961a);   Bergman  (1955b); 
^Bergman  (1960);  ^Bergman  (1951);  ^°Bergman  (1958a);  ^^Bergman  (1953);  ^^Bergman  (1955a);  ^^Bergman  (1962a); 
l^ColUns  and  Carpenter  (1970);  15Bergman  (1961b). 


Dice-Leraas  diagrams  (Figs.  1-19)  was 
divided  into  three  equal  triads,  using  the 
highest  and  lowest  individual  values  as 
outer  parameters.  In  these  figures  each  ver- 
tical bar  represents  the  mean,  each  hori- 
zontal line  the  range  expressed  as  a  per 
cent,  each  black  rectangle  the  95%  confi- 
dence interval,  each  number  the  quantity 
of  specimens  examined  in  that  taxon,  each 
horizontal  dotted  line  the  separation 
between  genera,  and  each  vertical  dashed 
Une  the  boundary  between  two  triads.  See 
Table  IV  for  a  comparison  of  taxa  assigned 
to  the  lowest,  middle,  and  highest  triads. 
Table  V  shows  the  degree  to  which  each 
taxon  differs  significantly  from  other  taxa 
in  this  study.  The  results  of  this  study  are 
compared  with  those  of  other  workers  in 
-Appendix  A. 

Posterior  End  of  Heart.  —  Assuming, 
strictly  for  the  sake  of  comparison,  that 
the  middle  triad  represents  the  normative 
condition,  there  is  a  clear  tendency  for  the 
heart  to  be  situated  more  posteriorly  than 


the  norm  in  both  sexes  of  Nerodia  rhom- 
bifera,  Regina  alleni,  Seminatrix  pygaea, 
and  Thamnophis  sauritus  (T.  melano- 
gaster,  T.  proximus,  and  T.  rufipunctatus 
exhibit  similar  tendencies,  but  to  a  lesser 
degree).  On  the  other  hand,  Tropidoclo- 
nian  lineatum  and  almost  half  the  taxa  of 
Thamnophis  (including  some  representa- 
tives from  three  of  Ruthven's  species 
groups)  tend  to  have  the  heart  displaced 
anteriorly  relative  to  the  norm. 

Anterior  End  of  Liver.  —  The  anterior 
end  of  the  liver  lies  markedly  farther  poste- 
riorly than  the  norm  in  both  sexes  of 
Regina  alleni  and  Seminatrix  pygaea.  It 
appears  to  extend  slightly  more  anteriorly 
than  the  norm  in  about  half  the  taxa  of 
Thamnophis  (the  same  ones  having  an 
anteriorly  displaced  heart)  and  in  female 
Clonophis  kirtlandii,  Nerodia  erythrogas- 
ter,  Storeria,  Tropidoclonion,  and 
Virginia.  The  most  posterior  placement  in 
Nerodia  is  again  found  in  TV.  rhombifera, 
and  in    Thamnophis  again   found  in    T. 


130 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Table  IV.   A  comparison  of  various  taxa  in  terms  of  the  triads  into  which  they 
fall.   L  represents  the  lowest  triad,  M  the  middle  triad,  and  H  the  highest  triad. 


^^■~--,..^^       Character 
Taxon       ^~~~^-,^^^ 

•   u 
0  OJ 

u 

•     (U 

4-'   > 

C  -H 
CO  M 

•  u 

■U  QJ 

m  > 

O  -H 

OJ 

•  >-i 

•U   U 

tfl  c 
o  « 

c 

T3 

■H 

•  J^ 

U 

C   • 

c 

•  -H 

o   • 
a.  u 

> 

C 

■a 

■H 

c  • 

<u 
c 
■v 

o   • 

CX.-H 

x: 

QJ  00 

>  c 

•H   Q> 

OJ 

c 

•H  U 

c 

•  QJ 

U     rH 

QJ 

c 
T3  x: 

•H  U 
J^     00 

c 

•  OJ 
1— 1  tH 

a, 
>.   n) 

QJ  r-\ 

C  u 

t3  QJ 
■H  > 
^  O 

QJ   OJ 

>  ^ 

H  O. 

1  tn 

■u  U 
U     QJ 
03  AJ 
QJ  C 

a  -H 

o"  9 

cf  9 

d  9 

(^  9 

a-  9 

d   9 

a-  9 

d  9 

Cf  9 

o"  9 

Cf   9 

Cf  9 

d  9 

Clonophis  kirtlandii 

L  M 

M  L 

-  - 

-  - 

H  H 

H  H 

H  H 

M  M 



M  M 

M  M 

M  M 

M  L 

Nerodia  cyolopion 

M  M 

M  M 

M  M 

M  M 

M  M 

H  M 

M  M 

M  M 

M  L 

M  M 

M  M 

H  H 

M  M 

erythrogaster 

L  M 

M  L 

M  M 

M  M 

M  L 

M  M 

M  L 

M  M 

M  L 

M  H 

H  H 

H  M 

M  M 

fasoiata 

M  M 

M  M 

M  M 

M  M 

M  M 

H  H 

M  M 

M  M 

M  L 

H  H 

H  M 

H  H 

M  M 

rhombifera 

H  H 

H  M 

M  M 

M  M 

M  M 

H  M 

M  M 

M  M 

L  L 

M  H 

M  M 

H  H 

M  M 

sipedon 

M  M 

M  M 

M  M 

M  M 

M  M 

H  M 

M  M 

M  M 

M  L 

M  H 

M  H 

H  M 

M  M 

valida 

M  M 

-  - 

-  M 

-  L 

M  M 

M  M 

M  M 

M  M 

-  - 

M  M 

M  M 

H  M 



Regina  alleni 

H  H 

H  H 

_  _ 

_  _ 

M  M 

M  M 

M  M 

M  M 

_  _ 

M  M 

M  M 

M  M 

H  H 

grahamii 

M  M 

-  M 

-  M 

-  M 

M  M 

H  H 

M  H 

M  H 

-  L 

M  M 

M  M 

M  M 

-  M 

rigida 

M  M 

-  M 

-  M 

-  H 

M  M 

M  M 

M  M 

M  M 

-  L 

M  M 

M  M 

M  M 

-  M 

septemvittata 

M  - 

M  - 

M  - 

-  - 

M  - 

M  - 

M  - 

M  - 

M  - 

M  - 

M  - 

M  - 

M  - 

Seminatvix  pygaea 

H  H 

H  H 

H  H 

H  H 

H  H 

M  H 

-  - 

L  M 

L  L 

M  M 

H  hI 

Storeria  dekayi 

M  M 

M  L 

H  M 

H  M 

M  M 

M  M 

H  H 

H  H 

H  M 

M  M 

M  M 

M  M 

M  M 

OGcipitomaoulata 

M  M 

M  L 

M  M 

H  - 

M  H 

M  M 

H  H 

H  M 

H  M 

M  L 

M  L 

L  L 

M  M 

Thamnophis  proximus 

M  M 

M  M 

M  M 

H  H 

H  H 

H  H 

H  H 

H  H 

M  L 

M  M 

M  M 

M  M 

M  M 

Sauritus       sauritus 

H  H 

-  M 

-  M 

-  M 

H  H 

H  H 

H  H 

H  H 

-  L 

M  M 

L  L 

M  M 

-  M 

group  1 

Thamnophis  brachystoma 

L  M 

_  _ 

_  _ 

_  _ 

M  M 

M  M 

M  M 

M  M 

_  _ 

M  M 

M  M 

M  M 

_  _ 

butleri 

M  M 

-  - 

-  - 

-  - 

M  M 

M  M 

M  M 

M  M 

-  - 

M  M 

M  M 

M  M 

-  - 

Radix              eques 

L  L 

L  L 

L  L 

L  L 

L  M 

L  M 

L  M 

L  M 

L  L 

M  H 

M  H 

M  M 

M  - 

group      marotanus 

L  L 

L  L 

L  L 

M  M 

M  M 

M  M 

M  M 

M  M 

M  L 

M  H 

M  M 

M  M 

M  M 

radix 

L  L 

L  L 

L  L 

M  M 

M  M 

M  H 

M  M 

M  H 

M  M 

M  H 

M  H 

M  H 

M  L 

Thamnophis  aouchii  A 

M  M 

L  - 

L  - 

M  - 

L  M 

L  M 

L  L 

L  M 

L  - 

M  M 

M  M 

M  M 

M  L 

couahii  B 

M  M 

M  L 

M  M 

M  M 

L  M 

L  M 

M  M 

M  M 

M  L 

M  M 

M  H 

M  M 

M  M 

elegans  A 

L  L 

L  L 

L  L 

M  - 

L  L 

L  M 

L  L 

L  M 

L  L 

M  H 

H  M 

M  M 

M  M 

elegans  B 

L  L 

L  L 

L  L 

M  M 

L  L 

L  M 

L  L 

L  M 

M  L 

M  M 

H  H 

M  M 

M  L 

Elegans         melanogaster 

M  H 

M  M 

M  M 

M  L 

M  M 

M  M 

M  M 

M  M 

M  L 

M  H 

M  H 

M  M 

M  M 

group      nigronuchalis 

M  M 

-  L 

-  M 

-  L 

L  L 

L  L 

L  L 

L  L 

-  L 

M  M 

M  M 

M  M 

-  M 

ordinoides 

L  M 

-  - 

-  - 

-  - 

M  M 

M  M 

M  M 

M  M 

-  - 

M  M 

M  M 

M  M 

-  - 

rufipunctatus 

M  M 

M  M 

-  - 

-  - 

L  M 

L  M 

L  L 

L  L 

-  - 

M  M 

M  M 

M  M 

M  M 

soalaris 

-  L 

M  L 

-  - 



M  M 

M  M 

M  M 

M  M 

-  - 

M  H 

M  H 

H  H 

-  L 

Thamnophis  chrysoaephalus 

M  M 

M  M 

-  _ 

_  _ 

M  M 

M  M 

M  M 

M  M 

_  _ 

M  M 

M  M 

M  M 

M  M 

ayrtopsis 

L  L 

L  L 

L  L 

M  L 

L  L 

M  M 

L  L 

M  M 

M  L 

M  H 

M  H 

H  H 

L  M 

Sirtalis       godmani 

L  M 

M  L 





L  M 

L  M 

L  M 

M  M 

-  - 

M  M 

H  M 

M  M 

M  M 

group      sirtalis  A 

L  L 

L  L 

L  L 

M  M 

M  M 

M  M 

M  M 

M  M 

M  L 

M  H 

M  M 

M  M 

M  M' 

sirtalis  B 

L  L 

L  L 

-  M 

-  M 

M  M 

M  M 

M  M 

M  M 

-  L 

M  H 

M  M 

M  M 

M  M 

Tropidoalonion  lineatum  A 

L  L 

-  - 

-  - 

-  - 

M  H 

M  M 

H  H 

M  M 

_  _ 

L  L 

L  L 

M  M 

_  _ 

_ linp.ntum   B 

-  L 

-  L 

-  M 

-  H 

-  H 

-  M 

-  H 

-  M 

-  M 

-  T, 

-  T, 

-  T, 

-  M 

Vvrg%nia  striatula 

M  M 

M  L 

-  M 

-  M 

H  H 

M  H 

H  H 

M  M 

-  M 

L  L 

L  L 

L  L 

M  L 

Valerias 

M  M  M  L 

H  H 

-  M 

H  H 

M  M 

H  H 

M  M 

H  H 

L  L 

L  L 

M  M 

M  L 

Ruthven's  species  groups 


No.  2 


Visceral  Topography  of  Snakes 


131 


"'  Males 

14      16      18     20     22      24     26 


C.      kirtlandii 

N.      eye lop  ion 

erythrogaster 

fasciata 
rhombifera 
sipedon 
valida 

R.  alleni 
grahamii 

rigida 
septemvittata 

Se,  pygaea 

St.  dekayi 

occipitomaculata 

Th.  brachystoma 
butleri 

chrysocephalus 

couchii  A 

couchii  B 

cyrtopsis 

elegans  A 

elegans  B 

eques 

godmani 

marcianus 

melanogaster 

nigronuchal is 

ordinoides 

proximus 

radix 

rufipunctatus 

sauritus 

scalaris 

sirtalis  A 

sirtalis  B 

Tr.  1  ineatum  A 
lineatum  B 

V.  striatula 
valeriae 


I 
I     - 


^• 


r 


-Hi' 


f^ 


-1-5 


Figure  1.     Location  of  the  posterior  end  of  the  heart  in  thamnophiine  snakes  (expressed  as  a  %  of  total 
ventrals).  Construction  of  this  and  subsequent  graphs  is  explained  on  pp.  127-129 


132 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


aoBdsja^ui: 


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No.  2 


Visceral  Topography  of  Snakes 


133 


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134 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


melanogaster  (except  females),   T.  proxi- 
mus,  T.  rufipunctatus,  and  T.  sauritus. 

Posterior  End  of  Liver.  —  The  poste- 
rior end  of  the  liver  extends  markedly 
farther  posteriorly  than  the  norm  in  both 
sexes  of  Virginia  valeriae.  In  general, 
Thamnophis  other  than  T.  proximus  and 
T.  sauritus  (and  female  T.  melanogaster) 
tend  to  have  the  posterior  end  of  the  liver 
lying  farther  anteriorly  than  in  any  other 
thamnophiines  save  Nerodia  erythrogaster 
and  N.  valida. 

Posterior  End  of  Pancreas.  —  In  males 
the  posterior  end  of  the  pancreas  extends 
farthest  posteriorly  in  Thamnophis  prox- 
imus and  the  two  species  of  Storeria;  in 
females  it  extends  farthest  posteriorly  in 
Regina  rigida,  Thamnophis  proximus,  T. 
sauritus,  Tropidoclonion  lineatum,  and 
the  two  species  of  Virginia.  About  half  the 
taxa  of  Thamnophis  tend  to  have  the  pan- 
creas located  more  anteriorly  than  in  any 
of  the  other  thamnophiines  except  female 
Nerodia  valida;  this  condition  is  most 
prounced  in  male  T.  eques.  Unfortunately, 
the  absence  of  data  for  one  of  the  sexes  in 
12  of  the  taxa  greatly  reduces  the  value  of 
the  pancreas  comparisons. 

Anterior  End  of  Right  Kidney.  —  In  all 
Thamnophis  except  T.  proximus  and  T. 
sauritus,  the  right  kidney  in  males  lies 
anterior  to  the  position  of  that  organ  in  all 
other  thamnophiines  except  Nerodia 
erythrogaster.  There  is  a  similar  tendency 
in  females,  but  it  is  neither  as  marked  nor 
as  consistent.  On  the  other  hand,  there  is 
marked  posterior  displacement  from  the 
norm  in  both  sexes  of  Seminatrix  pygaea, 
Thamnophis  proximus,  T  sauritus,  and 
Virginia  striatula,  and  a  similar  but  slightly 
less  pronounced  tendency  in  both  sexes  of 
Clonophis  kirtlandii  and  V.  valeriae  and  in 
females  of  Storeria  occipitomaculata  and 
Tropidoclonion  lineatum. 

Posterior  End  of  Right  Kidney.  —  The 
pattern  of  variation  here  is  generally  simi- 
lar to  that  described  in  the  preceding 
account.  The  most  notable  difference, 
however,  is  that  only  Thamnophis  proxi- 
mus, T.  sauritus,  and  female  Seminatrix 
pygaea  show  a  pronounced  extension  pos- 


teriorly. A  similar  but  less  pronounced 
trend  appears  in  males  of  Clonophis  kirt- 
landii, all  species  of  Nerodia  (except  N. 
erythrogaster  and  N.  valida),  Regina 
grahamii,  and  Seminatrix  pygaea. 

Anterior  End  of  Left  Kidney.  —  This 
position  lies  posterior  to  the  norm  in  both 
sexes  of  Clonophis  kirtlandii,  Seminatrix 
pygaea,  Tropidoclonion  lineatum,  both 
species  of  Storeria,  Thamnophis  proximus, 
T.  sauritus,  and  both  species  of  Virginia, 
and  in  females  of  Regina  grahamii.  Nero- 
dia erythrogaster  and  about  half  the  taxa 
of  Thamnophis  show  a  slight  tendency 
toward  anterior  displacement  from  the 
norm  (in  most  cases  this  tendency  is  better 
developed  in  males). 

Posterior  End  of  Left  Kidney.  —  The 
end  of  the  left  kidney  extends  more  poste- 
riorly than  the  norm  in  both  sexes  of 
Storeria  dekayi,  Thamnophis  proximus, 
and  T.  sauritus,  and  to  a  lesser  degree  in 
males  of  Storeria  occipitomaculata  and  fe- 
males of  Regina  grahamii,  Seminatrix 
pygaea,  and  Thamnophis  radix.  Males  of 
about  half  the  taxa  of  Thamnophis  show  a 
tendency  toward  anterior  displacement 
from  the  norm,  as  do  females  of  T.  nigro- 
nuchalis. 

Liver  Length.  —  The  liver  is  relatively 
long  in  the  genera  Storeria,  Tropidoclo- 
nion, and  Virginia.  Unfortunately  we  have 
no  data  for  males  of  the  latter  two  genera 
or  for  female  Seminatrix.  Male  Seminatrix 
have  an  even  longer  liver  than  is  found  in 
the  other  three  genera.  Two  male  and  two 
female  Clonophis,  although  not  shown  on 
the  Dice-Lerras  diagram  because  of  the 
small  sample  size,  also  have  a  relatively 
long  liver  (mean  values  of  26.3  and  26.0, 
respectively). 

Right  Kidney  Length.  —  The  right  kid- 
ney is  relatively  short  in  both  sexes  of  Tro- 
pidoclonion lineatum  and  in  both  species 
of  Virginia,  and  in  females  of  Storeria  oc- 
cipitomaculata and  males  of  Seminatrix 
pygaea.  In  males  there  is  a  tendency 
toward  a  greater  length  than  the  norm  in 
the  species  of  Nerodia  and  about  half  the 
taxa  of  Thamnophis;  the  same  tendency  is 
present  in  females  but  it  is  developed  to  a 


No.  2 


Visceral  Topography  of  Snakes 


135 


lesser  degree.  Notably,  Clonophis  and 
Regina  separate  completely  from  Nerodia 
on  this  character. 

Left  Kidney  Length.  —  The  left  kidney 
is  relatively  short  in  both  sexes  of  Semina- 
trix  pygaea,  Thamnophis  sauritus,  Tropi- 
doclonion  lineatum,  and  the  two  species  of 
Virginia,  and  in  females  of  Clonophis  kirt- 


landii,  Storeria  occipitomaculata,  and 
Thamnophis  proximus.  The  tendencies 
seen  with  regard  to  left  and  right  kidney 
lengths  are  generally  similar,  but  the  dis- 
tinction between  Nerodia  and  Clonophis- 
Regina  is  less  clearly  defined  in  the  left 
kidney  length  of  males. 


18   20  22  24   26 

28  30   32 

34  36  38 

c. 

kirtlandi  i 

cyclopion 

erythrogaster 

fasciata 

rhombifera 

sipedon 

al leni 

grahamii 

rigida 

septemvittata 

pygaea 

dekayi 
occipitomaculata 

chrysocephalus 

couchii  A 

couchii  B 

cyrtopsis 

elegans  A 

elegans  B 

eques 

godmani 

marcianus 

melanogaster 

nigronuchalis 

proximus 

radix 

rufipunctatus 

sauritus 

scalaris 

sirtalis  A 

sirtalis  B 

lineatum  B 

striatula 
valeriae 

K- 

\-  5    { 

N. 

1 

1 
1 

—  9    1 

9 
-6 

R 

1 

1 
1 
1 
1 

1 

i^-l 

1    c 

!  ^ 

Se. 

r 

i 

-^  7 

St. 

Th. 

— ■■■-  7 

1  +5 

■  6    1 

5      1 
-  3 

—  9 

1 

1  —. 

iiHm^ 

-his 

1  -^ 
1  ^ 

1±:.:i.. 

1^ —  ^1 

I 

Tr. 

1 
1 

1 
■ .1 

V. 

•  ^J  ^    ' 

~1  ""^ 

^^M ' 

Figure  2.     Location  of  the  anterior  end  of  the  liver  in  male  thamnophiine  snakes  (expressed  as  a  %  of  total 
ventrals). 


136 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Heart-Liver  Interspace.  —  Both  sexes 
of  Seminatrix  pygaea  and  males  of  Regina 
alleni  have  a  relatively  long  interspace,  a 
tendency  that  is  also  seen  in  males  of 
Nerodia  sipedon  and  Regina  septemvit- 
tata,  and  in  females  of  Nerodia  cyclopion, 
Tropidoclonion     lineatum,     and     three 


species  of  Regina  (no  data  available  for 
female  septemvittata).  The  interspace  is  re- 
latively short  in  females  of  Clonophis  kirt- 
landii. 

Kidney  Overlap.  —  The  greatest  degree 
of  kidney  overlap  occurs  in  Nerodia  and  a 
few     Thamnophis    {cyrtopsis,     male 


18   20  22  24  26  28 

30  32  34 

36  38 

c. 

kirtlandii 

cyclopion 

erythrogaster 

fasciata 

±A    1 

N. 

»!' , 

1 

-7\ 

rhombifera 
sipedon 

al leni 

1  -^'0  1 

R. 

1 

1  -f- 

6 

grahaniii 

-H-^ 

rigida 
septenivi  ttata 

pygaea 

dekayi 
occipitomaculata 

chrysocephalus 
couchii  A 

1  h 

—  4 

Se. 

T 

i 

'     1 

1     1 

St. 

-4'°       1 

Th. 

IV : 

i' 
1 

couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
godmani 

marcianus 

melanogaster 

nigronuchal is 

proximus 

radix 

rufipunctatus 

'  -1- 

4   1 
8   j 
3  1 

sauritus 

r*-^ 

scalaris 
sirtalis  A 
sirtalis  B 

lineatum  B 

striatula 

valeriae 

....'...^. 

I  

Tr. 

f-l-6        1 

V. 

~^'"      \ 

Figure  3.     Location  of  the  anterior  end  of  the  liver  in  female  thamnophiine  snakes  (expressed  as  a  %  of  total 
ventrals). 


No.  2 


Visceral  Topography  of  Snakes 


137 


marcianus,  melanogaster,  radix,  scalaris). 
The  least  amount  of  overlap  occurs  in 
Storeria  occipitomaculata  and  Virginia 
striatula.  Clonophis-Regina  again  separate 
completely  from  Nerodia. 

Liver-Gall  Bladder  Interspace.  — 
McDowell  (1979)  reported  that  the  most 
striking  visceral  feature  of  all  Acrochordus 
is  the  close  proximity  of  the  gall  bladder  to 
the  liver.  In  Acrochordus  granulatus  the 
gall  bladder  usually  lies  behind  the  liver, 
but  is  separated  from  it  by  less  than  one 
gall  bladder  length;  in  A.  arafurae  the  gall 
bladder  lies  immediately  behind  the  liver; 
and  in  A.  javanicus  the  gall  bladder  is 
usually  overlapped  by  the  posterior  end  of 
the  liver.  McDowell  stated  that  Acrochor- 
dus seems  to  be  the  only  snake  genus 
known  to  have  the  gall  bladder  so  near  the 
liver,  and  he  noted  that  having  the  gall 
bladder  displaced  far  behind  the  liver  is 
often  cited  as  a  distinctive  feature  of 
snakes. 

A  survey  of  Bergman's  many  studies 
(1950-1965)  on  the  visceral  topography  of 
a  wide  variety  of  snakes  reveals  that  the 
condition  described  by  McDowell  (1979)  is 
somewhat  more  widespread  than  he  had 
thought  and  that  this  feature  exhibits 
sexual  dimorphism  in  a  number  of  species. 
Bergman's  findings  can  be  summarized  as 
follows: 

1.  No  interspace,  liver  overlaps  gall 
bladder:  Colubridae,  Homalopsinae  —  fe- 
male Enhydris enhydris  (\955e),  Homa lop- 
sis  buccata  (1951),  male  Hypsirhina  {  =  En- 
hydris) alternans  (1960);  Acrochordidae  — 
Acrochordus  javanicus  (1958a). 

2.  Interspace  less  than  one  gall  bladder 
length:  Colubridae,  Homalopsinae  — 
male  Enhydris  enhydris  (1955e),  female 
Cerberus  rhynchops  (1955c),  Hypsirhina 
( =  Enhydris)  plumbea  (1960);  Acrochordi- 
dae —  Acrochordus  granulatus  (1958a); 
Elapidae  —  Enhydrina  schistosa  (1955d). 

3.  Interspace  one  to  two  times  gall  blad- 
der length:  Colubridae,  Natricinae  — 
male  Matrix  {- Sinonatrix)  trianguligera 
(1959b),  female  Matrix  (^Xenochrophis) 
vittata  (1950);  Colubridae,  Homalopsinae 
—  female  Hypsirhina  ( =  Enhydris)  alter- 


nans (1960),  male  Cerberus  rhynchops 
(1955c);  Elapidae  —  female  Hydrophis 
fascia tus  (1962a),  female  Thalassophis 
anomalus  (1954);  Viperidae  —  Ancistro- 
don  (  =  Calloselasma)  rhodostoma 
( 1 96 1  b) ,  Trimeresurus  gramineus  ( 1 96 1  b) . 

4,  Interspace  more  than  twice  gall  blad- 
der length:  Colubridae,  Natricinae  —  Ma- 
trix {^  Rhabdophis)  chrysarga  (1959a),  TV. 
i^Rhabdophis)  subminiata  (1956b), 
female  TV.  {  =  Sinonatrix)  trianguligera 
(1959b),  male  TV.  {  =  Xenochrophis)  vittata 
(1950);  Colubridae,  Homalopsinae  — 
Eordonia  leucobalia  (1960);  other  Colubri- 
bae  —  Ablabes  {  =  Gongylosoma) 
baliodeira  (1963),  Calamaria 
multipunctata  (1965),  Coluber  melanurus 
(  =  Elaphe  flavolineata)  (1961a),  C. 
(  =  Elaphe)  radiatus  (1961a),  Dendrophis 
( =  Dendrelaphis)  pictus  (1955b),  Dryophis 
(=Ahaetulla)  prasinus  (1956a),  Elapoides 
fuscus  (1956-58),  Ptyas  korros,  P.  mucosa 
(1952);  Aniliidae  —  Cylindrophis  rufus 
(1953);  Boidae  —  Xenopeltis  unicolor 
(1955a);  Elapidae  —  Bungarus  candidus, 
B.  fasciatus,  male  Hydrophis  fasciatus, 
Maja  tripudians  (1962b),  male 
Thalassophis  anomalus  (1954). 

We  found  the  Thamnophiini  to  be  highly 
variable  in  this  character  although  the 
majority  of  individuals  do  have  an  inter- 
space greater  than  one  gall  bladder  length 
(see  Table  VI  for  details).  Noteworthy 
exceptions  are  the  females  of  Thamnophis 
melanogaster  and  Virginia  valeriae,  in 
which  the  mean  values  are  0.9  and  0.2, 
respectively.  In  general,  the  interspace 
tends  to  be  relatively  short  in  most 
Merodia,  Storeria,  and  Virginia,  and  rela- 
ti'ely  long  in  Regina,  most  Thamnophis, 
and  Tropidoclonion.  By  far  the  greatest  in- 
terspace/gall bladder  values  occur  in 
Thamnophis proximus  and  T.  sauritus,  but 
this  reflects  unusually  short  gall  bladders 
rather  than  exceptionally  long  interspaces 
in  these  animals. 

Asymmetry  of  Kidney  Lengths 

In  only  11  taxa  are  the  differences  in 
length  between  the  right  and  left  kidneys 
statistically  significant.  The  left  kidney  is 
longer  than  the  right  in  male  Thamnophis 


138 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


c.  couchii  (difference  between  means  1 .6, 
significantly  different  at  p<.01)  and  female 
T.  nigronuchalisilA,  p<.02).  The  right 
kidney  is  longer  than  the  left  in  male 
Thamnophis  sauritus  (1.2,  p<.01)  and 
female  T.  cyrtopsis  {\  .1 ,  p  <.02),  T.  radix 
(1.9,  p<.01),  T.  sirtalis  fitchi  (2.0,  p<01), 
Nerodia  cyclopion  (1.4,  p<.01),  N. 
rhombifera  (1.7,  p<.05),  N.  sipedon  (1.2, 


p<.02),  Regina  alleni  (1.5,  p<.02),  and  R. 
grahamii  (1.2,  p<.05). 

Discriminant  Analysis 

In  an  effort  to  ascertain  which,  if  any, 
characters  could  be  used  taxonomically  to 
separate  genera  and  other  groups,  stepwise 
discriminant  analysis  was  performed  using 
the    Statistical    Package    for    the    Social 


JQ         42 


56        58 


N.  cyclopion 
erythrogaster 
fasciata 
rhombifera 
sipedon 
valida 

R.  grahamii 
rigida 
septemvittata 

St.  dekayi 

occipitomaculata 


Th, 


Tr, 

V. 


couchii  A 

couchii  B 

cyrtopsis 

elegans  A 

elegans  B 

eques 

marcianus 

melanogaster 

nigronuchalis 

proximus 

radix 

sauritus 

sirtalis  A 
sirtalis  B 

lineatum  B 

striatula 
valeriae 


Figure  4.     Location  of  the  posterior  end  of  the  liver  in  male  thamnophiine  snakes  (expressed  as  a  %  of  total 
ventrals). 


No.  2 


Visceral  Topography  of  Snakes 


139 


Sciences  (SPSS)  (Nie  et  al.,  1975;  Hull  and 
Nie,  1979).  Only  adult  male  specimens 
were  used  in  this  part  of  the  study  (see 
Materials  and  Methods).  Elsewhere  in  this 
paper  under  Materials  and  Methods  we 
have  discussed  the  details  of  how  the  speci- 
mens were  treated  for  the  discriminant 
analysis. 

In  stepwise  dicriminant  analysis,  the 
variable  that  best  discriminates  among  the 
groups  enters  the  model  first,  then  the  next 
best  discriminating  variable  enters,  etc. 
The  process  terminates  when  there  are  no 
more  variables  that  contribute  significantly 
to  discrimination  among  the  groups.  In 
this  analysis  four  variables  were  found  to 
discriminate  among  the  groups.  In  the 
rior  right  kidney,  (3)  posterior  heart,  and 
(4)  kidney  overlap.  In  the  four  variable 
model,  all  groups  but  Clonophis  and  Tro- 
pidoclonion  were  significantly  different 
(p<.05)  (see  Table  VII). 

Eleven  groups  were  used  in  the  dis- 
criminant analysis,  and  four  linear  discri- 
minant functions  were  computed.  How- 
ever, only  the  first  two  were  retained  as 


they  explain  83.21  %  of  the  relative  varia- 
tion (function  1  accounts  for  59.82%  and 
function  2  accounts  for  23.39%).  Function 
1  is  generally  a  right  kidney  anterior  di- 
mension. Function  2  is  a  kidney  overlap 
and  heart  posterior  dimension.  The  all- 
groups  scatterpoint  diagram  with  two  dis- 
criminant functions  appears  in  Fig.  20.  On 
dimension  1,  we  see  that  Seminatrix,  the 
Sauritus  group  of  Thamnophis,  and 
Virginia  are  separated  widely  from  the 
Elegans,  Sirtalis,  and  Radix  groups  of 
Thamnophis.  On  dimension  2,  we  see  that 
Nerodia  is  the  most  widely  separated  group 
from  Tropidoclonion  and  Storeria.  Ap- 
parently, as  the  right  kidney  anterior  mea- 
sure increases,  the  specimens  are  more 
likely  to  belong  to  Seminatrix,  the  Sauritus 
group  of  Thamnophis,  and  Virginia. 
Similarly,  as  right  kidney  anterior  de- 
creases, specimens  are  more  likely  to 
belong  to  the  Elegans,  Sirtalis,  and  Radix 
groups  of  Thamnophis.  Also,  as  kidney 
overlap  and  heart  posterior  measurements 
increase,  the  specimens  are  more  likely  to 
belong  to  Nerodia.  Similarly,  as  these  mea- 


Table  VII.   F  statistics  and  significance  between  pairs  of  taxa  in  the  four  variable  model  (df=4,  280). 


Taxon 

1  Clonophis 

2 

3 

4 

5 

6 

7 

8 

9 

10 

2  Nerodia 

16.557 
0.0000* 

3  Regina 

7.4392 
0.0000 

15.827 
0.0000 

4  Seminatrix 

11.479 
0.0000 

26.348 
0.0000 

9.4137 
0.0000 

5  Storeria 

6.7379 
0.0000 

52.616 
0.0000 

20.963 
0.0000 

20.288 
0.0000 

6  Thamnophis 

(Sauritus    group; 

6.5426 
0.0000 

27.357 
0.0000 

11.059 
0.0000 

5.1882 
0.0005 

14.861 
0 . 0000 

7  Thamnophis 
(Radix   group; 

8.6467 
0.0000 

38.890 
0.0000 

20.513 
0.0000 

37.208 
0.0000 

24.534 
0.0000 

37.326 
0.0000 

8  Thamnophis 
(Elegans    groupj 

24.343 
0.0000 

70.264 
0.0000 

40.509 
0.0000 

52.621 
0.0000 

58.937 
0.0000 

74.748 
0.0000 

24.394 
0.0000 

9  Thamnophis 

(Sirtalis   group,) 

16.569 
0.0000 

43.839 
0.0000 

27.789 
0.0000 

44.918 
0.0000 

42.816 
0.0000 

55.380 
0.0000 

7.5158 
0.0000 

4.7066 
0.0011 

LO  Tropidoclonion 

1.2101 
0.3067 

23.372 
0 . 0000 

11.002 
0.0000 

14.360 
0.0000 

8.1301 
0.0000 

11.953 
0.0000 

10.465 
0.0000 

22.652 
0.0000 

16.875 
0.0000 

LI  Virginia 

7.9233 
0.0000 

57.601 
0.0000 

15.848 
0.0000 

7.0630 
0.0000 

28.365 
0.0000 

16.768 
0.0000 

50.209 
0.0000 

76.698 
0.0000 

62.552 
0.0000 

6.9183 
0.0000 

*   significance  leve 

!l 

140 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


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No.  2 


Visceral  Topography  of  Snakes 


141 


surements  decrease,  the  specimens  are 
more  likely  to  belong  to  Tropidoclonion 
and  Storeria. 

The  model  was  used  to  classify  the  294 
original  specimens.  The  classification 
matrix  indicates  how  specimens  were  class- 
ified by  the  model  (see  Table  VIII).  Over 
66%  of  the  specimens  were  correctly  class- 
ified. The  Elegans  group  of  Thamnophis, 
which  had  the  largest  number  of  speci- 


mens, had  the  highest  prior  probability  of 
21.4%.  In  the  order  of  highest  percentage 
to  lowest,  Virginia  was  classified  correctly 
88.2%  of  the  time,  Nerodia  86.3%,  the 
Sauritus  group  of  Thamnophis  85.7%,  the 
Elegans  group  of  Thamnophis  76.2%, 
Seminatrix  75.0%,  Storeria  66.1  ^q,  the 
Radix  group  of  Thamnophis  63.6%, 
Tropidoclonion  60.0%,  Regina  58.3%,  the 
Sirtalis  group  of  Thamnophis  29.8%,  and 


36       38       40       42       44      46 


% 
48 


50       52       54       56       58       60 


N.     cyclop  ion 

erythrogaster 
fasciata 
rhombifera 
sipedon 
val ida 

R.  grahamii 
rigida 
septemvittata 

St.  dekayi 

occipitomaculata 

Th.  couchii  A 
couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
marcianus 
melanogaster 
nigronuchalis 
proximus 
radix 
sauritus 

sirtalis  A 
sirtalis  B 

Tr.  lineatum  B 

V.   striatula 
valeriae 


4-^ 

—  4 


Figure  5.     Location  of  the  posterior  end  of  the  liver  in  female  thamnophiine  snakes  (expressed  as  a  %  of 
total  ventrals). 


142 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


Clonophis  16.7<^o  (less  than  chance).  Vir- 
ginia has  the  highest  percentage  correctly 
classified,  the  Sauritus  group  of  Thamno- 
phis  the  third  highest,  and  Seminatrix  the 
fifth  highest,  a  notable  finding  inasmuch 
as  these  taxa  ranked  only  sixth,  eighth,  and 
ninth,  respectively,  in  terms  of  the  number 
of  specimens  per  group. 

The  discriminant  analysis  was  able  to 
distinguish  among  the  eight  genera  (as  well 
as  among  Ruthven's  four  species  groups  of 


Thamnophis)  at  the  0.05  level  ex'^ept  that 
Clonophis  and  Tropidoclonion  could  not 
be  distinguished  from  each  other.  All 
groups  except  Clonophis  could  be  classi- 
fied by  the  model  with  greater  success  than 
the  21%  prior  probability  obtained  by 
placing  them  all  in  the  Elegans  group  of 
Thamnophis,  the  numerically  largest 
sample.  Thus  the  visceral  topographic  data 
are  remarkably  concordant  with  the  other 
kinds   of  morphological  data  that  have 


4i      46       46       50      52      54 


% 
56 


N.     cyclopion 

erythrogaster 

fasciata 

rhombifera 

sipedon 

valida 

R.  grahamii 
rigida 

St.  dekayi 

occipitomaculata 

Th.  couchii  A 
couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
marcianus 
mel  an  og  aster 
nigronuchal is 
proximus 
radix 

sauritus 
sirtalis  A 
sirtalis  B 

Tr.  lineatum  B 

V.     striatula 
valeriae 


5  8      60       62      64       66       68 


Figure  6.     Location  of  the  posterior  end  of  the  pancreas  in  male  thamnophiine  snakes  (expressed  as  a  %  of 
total  ventrals). 


No.  2 


Visceral  Topography  of  Snakes 


143 


been     used     to     generate     the    existing 
classification  of  thamnophiine  snakes. 

Within  the  genus  Thamnophis,  all  of 
Ruthven's  species  groups  except  the  5/>- 
talis  group  are  distinguishable  from  each 
other.  In  the  classification  matrix  (Table 
VIII),  members  of  the  Sirtalis  group  are 
misclassified  as  members  of  the  Elegans 
group  much  more  frequently  (42. 6*^0)  than 
they  are  correctly  classified  (29.8%);  they 
also   are  often  misclassified   (21.3%)  as 


members  of  the  Radix  group.  One  of  the 
most  interesting  results  of  the  discriminant 
analysis  is  the  wide  separation  of  the 
Sauritus  group  from  the  other  three  groups 
(see  Fig.  20). 

Conclusions 

Visceral  topographic  data  alone  do  not 
clearly  delimit  thamnophiine  genera  nor 
establish  inter-  or  intrageneric  relation- 
ships.    Nevertheless,     some    trends    are 


% 


cyclop ion 

erythrogaster 

fasciata 

rhombifera 

sipedon 

valida 


R,  grahamii 
rigida 

St.  dekayi 

occipitomaculata 
Th.  couchii  A 

couchii  B 

cyrtopsis 

elegans  A 

elegans  B 

eques 

marcianus 

melanogaster 

nigronuchal is 

proximus 

radix 

sauritus 

sirtalis  A 

sirtalis  B 

Tr.  lineatum  B 

V.  striatula 
valeriae 


46   48   50   52   54   56   58   60   62   64   66 


Figure  7.     Location  of  the  posterior  end  of  the  pancreas  in  female  thamnophiine  snakes  (expressed  as  a  %  of 
total  ventrals). 


144 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


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No.  2 


Visceral  Topography  of  Snakes 


145 


apparent  that  serve  to  support  taxonomic 
conclusions  based  on  other  kinds  of  char- 
acters. 


Clonophis  kirtlandii  differs  from  all 
species  of  Nerodia  (in  which  genus  it  was 
formerly  placed;  see  Rossman,  1963b)  in 


C.     kirtlandii 

N.  cyclopion 

erythrogaster 

fasciata 
rhoinbifera 
sipedon 
valida 

R.  alleni 
grahamii 

rigida 
septemvittata 

Se.  pygaea 

St.  dekayi 

occipitomaculata 

Th.  brachystoma 
butleri 

chrysocephalus 
couchii  A 
couch ii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
godmani 
marcianus 
melanogaster 

nigronuchal is 

ordinoides 

proximus 

radix 

rufipunctatus 

sauritus 

scalaris 

sirtalis  A 
sirtalis  B 


Tr, 


lineatum  A 
lineatum  B 

striatula 
valeriae 


Figure  8.     Location  of  the  anterior  end  of  the  right  kidney  in  male  thamnophiine  snakes  (expressed  as  a  %  of 
total  ventrals). 


146 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


having  a  longer  liver,  shorter  kidneys  (the 
anterior  ends  have  been  displaced  poste- 
riorly), and  less  kidney  overlap.  Female 
Clonophis  can  also  be  distinguished  from 


female  Nerodia  by  having  a  shorter  heart- 
liver  interspace,  but  this  distinction  does 
not  apply  to  males. 
The  genus  Regina  has  also  been  included 


C.     kirtlandii 

N.     cyclopion 

erythrogaster 

fasciata 
rhombifera 
sipedon 
val ida 

R.  alleni 
grahamii 

rigida 
septemvittata 

Se.  pygaea 

St.  dekayi 

occipitomaculata 

Th.  brachystoma 
butleri 

chrysocephalus 
couchii  A 
couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
godniani 

marcianus 

melanogaster 

nigronuchal is 

ordinoides 

proximus 

radix 

rufipunctatus 

sauritus 

scalaris 

sirtalis  A 

sirtalis  B 

Tr.  lineatum  A 
lineatum  B 

V.   striatula 
valeriae 

Figure  9.     Location  of  the  anterior  end  of  the  right  kidney  in  female  thamnophiine  snakes  (expressed  as  a  % 
of  total  ventrals). 


No.  2 


Visceral  Topography  of  Snakes 


147 


in  Nerodia  in  the  past  (Smith  and  Huheey, 
1960;  Rossman,  1963b).  It  differs  from 
Nerodia  in  having  somewhat  shorter  kid- 
neys, less  kidney  overlap,  and  a  longer 


liver-gall  bladder  interspace.  Regina  alleni 
has  both  the  posterior  end  of  the  heart  and 
the  anterior  end  of  the  hver  situated  more 
posteriorly    than    in    the    other    crayfish 

% 


77     74  76  7  8  80  82  84  86  88  90  92  94 


C.  kirtlandii 

N.  cyclop  ion 

erythrogaster 
fasciata 
rhombifera 
sipedon 
val ida 

R.  alleni 
grahamii 
rigida 
septemvi ttata 

Se.  pygaea 
St.  dekayi 

occipitomaculata 

Th.  brachystoma 
butleri 

chrysocephalus 
couchii  A 
couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
godmani 
marcianus 
melanogaster 
nigronuchalis 
ordinoides 
proximus 
radix 

rufipunctatus 
sauritus 
scalaris 
sirtalis  A 
sirtalis  B 

Tr.  lineatum  A 
lineatum  B 

v.  striatula 
valeriae 


Figure  10.     Location  of  the  posterior  end  of  the  right  kidney  in  male  thamnophiine  snakes  (expressed  as  a  % 
of  total  ventrals). 


148 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


snakes.  Male  R.  alleni  have  the  longest 
heart-liver  interspace  of  any  thamnophiine 
in  our  study,  but  data  for  male  R.  grahamii 
and  R.   rigida  are  lacking.   In  terms  of 


positional  characters  (as  opposed  to  organ 
or  interspace  lengths),  the  organs  of  R. 
rigida  usually  have  the  anteriormost  posi- 
tions within  the  genus. 


7i       7b      78      80      82      84      86     88      90      92      94 


C.      kirtlandii 

N.     cyclopion 

erythrogaster 
fasciata 
rhombifera 
sipedon 
val ida 

R.  alleni 
grahamii 
rigida 
septemvi ttata 

Se.  pygaea 

St.  dekayi 

occipitomaculata 

Th.  brachystoma 
butleri 

chrysocephalus 
couchii  A 
couchii  B 
cyrtopsis 
elegans  A 

elegans  B 

eques 

godmani 

marcianus 

melanogaster 

nigronuchal is 

ordinoides 

proximus 

radix 

rufipunctatus 

sauritus 

scalaris 

sirtalis  A 
sirtalis  B 


Tr, 


V. 


lineatum  A 
1  ineatuni  B 

striatula 
valeriae 


Figure  1 1.     Location  of  the  posterior  end  ol  tlie  right  kidney  in  female  thamnophiine  snakes  (expressed  as  a 
%  of  total  ventrals). 


No.  2 


Visceral  Topography  of  Snakes 


149 


Within  the  genus  Nerodia,  where  there  is 
variation  from  the  generic  "norm,"  N. 
erythrogaster  or,  less  frequently,  N.  valida 
invariably  has  the  anteriormost  position. 


Nerodia  rhorribifera  shows  a  posterior  dis- 
placement of  the  heart  and  of  the  anterior 
end  of  the  liver  (but  only  slightly  more 
than  in  N.  cyclopion).  Organ  and  inter- 


kirtlandii 

cyclopion 

erythrogaster 

fasciata 

rhombifera 

sipedon 

valida 


R. 


alleni 
grahamii 
rigida 
septemvittata 

Se .  pygaea 

St.  dekayi 

occipitomaculata 

Th .  brachy stoma 
butlen 

chrysocephalus 
couchii  A 
couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
godmani 
marcianus 
melanogaster 

nigronuchalis 
ordinoides 
proximus 
radix 

rufipunctatus 
sauritus 
scalaris 
sirtalis  A 
sirtalis  B 
Tr.  lineatum  A 
1 ineatum  B 

V,  striatula 
valeriae 


Figure  12.     Location  of  the  anterior  end  of  the  left  kidney  in  male  thamnophiine  snakes  (expressed  as  a  %  of 
total  ventrals). 


150 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


space  lengths  show  no  consistent  intrage- 
neric  trends. 

The  only  external  feature  that  has  been 
used  consistently  to  distinguish  the  genera 


Nerodia  and  Thamnophis  is  the  presence 
of  an  undivided  anal  plate  in  the  latter 
(Conant,  1961),  but  Varkey  (1979)  has 
demonstrated    several    consistent    differ- 


% 

77     74     76     78     80     82     84     B6     88      90 


C.     kirtlandii 

N.  cyclopion 

erythrogaster 
fasciata 
rhombifera 
sipedon 
val ida 

R.  alleni 
grahamii 
r  i  g  i  d  a 
septemvittata 

Se.  pygaea 
St.  dekayi 

occipi tomaculata 

Th.  brachystoma 
butleri 

chrysocephalus 
couchii  A 
couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 

godmani 
marcianus 
melanogaster 
nigronuchal is 
ordinoides 
proximus 
radix 

rufipunctatus 
sauritus 
scalaris 
sirtalis  A 
sirtalis  B 
Tr.  lineatum  A 
lineatum  B 

V.   striatula 
valeriae 


Figure  13.     Location  of  the  anterior  end  of  the  left  kidney  in  female  thamnophiine  snakes  (expressed  as  a  % 
of  total  ventrals). 


No.  2 


Visceral  Topography  of  Snakes 


151 


ences  in  cranial  myology  between  the  two 
genera.  Our  data  on  visceral  topography 
do  not  provide  an  unequivocal  picture  of 


the  Nerodia-Thamnophis  relationship. 
Nevertheless,  if  one  were  to  remove  N. 
erythrogaster   and    N.    valida    from    the 


%  Females 

82     84     86     88     90     92    94     96     98 


%  Males 

82      84     86     88     90     92     94     96     98 


Se 


Th 


Tr, 


kirtlandii 

cyclopion 

erythrogaster 

fasciata 

rhombifera 

sipedon 

valida 


al leni 
grahamii 
rigida 
septemvittata 

pygaea 

St.  dekayi 

occipi tomaculata 

brachystoma 
butleri 

chrysocephalus 
couchii  A 
couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
godmani 

marcianus 

melanogaster 

nigronuchalis 

ordinoides 

proximus 

radix 

rufipunctatus 

sauritus 

scalaris 

sirtalis  A 

sirtalis  B 


lineatum  A 
lineatum  B 

striatula 
valeriae 


Figure  14.     Location  of  the  posterior  end  of  the  left  kidney  in  thamnophiine  snakes  (expressed  as  a  %  of 
total  ventrals). 


152 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


former  and  T.  proximus  and  T.  sauritus 
from  the  latter,  the  posterior  end  of  the 
liver  in  the  remaining  taxa  of  Thamnophis 
would  lie  anteriorly  to  its  relative  position 
in  the  remaining  Nerodia;  the  anterior  and 
posterior  ends  of  the  right  kidney  in  males 
show  a  similar  relationship.  As  a  matter  of 
fact,  the  anterior  end  of  the  right  kidney  in 
male  Thamnophis  (other  than  T.  proximus 
and  T.  sauritus)  lies  anteriorly  to  that 
position  in  all  other  thamnophiines  save  N. 
erythrogaster.  All  taxa  of  Thamnophis 
(except  T.  sauritus)  differ  from  all  species 
of  Nerodia  (except  N.  erythrogaster  and  TV. 


valida)  in  having  the  posterior  end  of  the 
right  kidney  of  females  lying  posteriorly  to 
that  of  males.  All  taxa  of  Thamnophis 
(except  female  T  melanogaster  and  T.  ni- 
gronuchalis)  have  a  liver-gall  bladder  inter- 
space more  than  twice  the  length  of  the  gall 
bladder;  in  all  species  of  Nerodia  (except 
N.  erythrogaster  and  N.  valida)  the  inter- 
space is  between  one  and  two  times  as  long 
as  the  gall  bladder.  Whether  the  frequent 
similarity  in  organ  positions  of  N.  erythro- 
gaster to  the  garter  snakes  reflects  phyletic 
affinities,  convergence  due  to  ecological 
similarities  (N.  erythrogaster  is  more  ter- 


N.     cyclopion 
erythrogaster 

fasciata 
rhombifera 
sipedon 
R.     grahamii 
rigida 
septemvi ttata 

Se.  pygaea 

St.   dekayi 

occipi tomaculata 

Th.  couchii   A 
couchii   B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
marcianus 
melanogaster 
nigronuchal is 
proximus 
radix 
sauritus 
sirtalis  A 
sirtalis  B 

Tr.   lineatum  B 
V    .   striatula 
valeriae 


Figure  15.     Liver  length  in  thamnophiine  snakes  (expressed  as  a  %  of  total  ventrals). 


No.  2 


Visceral  Topography  of  Snakes 


153 


restrial  than  its  congeners  and  has  a  larger 
anuran  component  in  its  diet  —  Mushin- 
sky  and  Hebrard,  1977;  Kofron,  1978),  or 
some  other  factors,  we  cannot  say. 


Rossman  (1963a)  noted  that  the  Sauritus 
group  of  Thamnophis  shows  no  close  affi- 
nity to  any  of  the  other  groups  established 
by  Ruthven  (1908),  and  our  study  confirms 


C.     kirtlandii 

N.  cyclopion 

erythrogaster 

fasciata 
rhombifera 
sipedon 
valida 

R.  alleni 
grahamii 
rigida 
septemvittata 

Se.  pygaea 
St.  dekayi 

occipi tomaculata 

Th.  brachystoma 
butleri 

chrysocephalus 
couchii  A 
couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
godmani 
marcianus 
melanogaster 
nigronuchalis 
ordinoides 
proximus 
radix 

ruf ipunctatus 
sauritus 
scalaris 
sirtalis  A 
sirtalis  B 

Tr.  lineatum  A 
lineatum  B 

V.  striatula 
valeriae 


Figure  16.     Right  icidney  length  in  thamnophiine  snakes  (expressed  as  a  %  of  total  ventrals). 


154 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


that  observation.  In  fact,  the  marked  dis- 
similarity of  the  ribbon  snakes  (T.  proxi- 
mus  and  T.  sauritus)  to  other  Thamnophis 
in  most  visceral  topographic  features  (see 
Table  IX)  proved  to  be  the  most  striking, 


C.     kirtlandii 

N.     cyclopion 

erythrogaster 

fasciata 

rhombifera 

sipedon 

valida 

R.     alleni 
grahamii 
rigida 
septemvittata 

Se.  pygaea 

St.  dekayi 

occipitomaculata 

Th.   brachystoma 
butleri 

chrysocephalus 
couchii  A 
couchii   B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
godmani 
marcianus 
melanogaster 
nigronuchal is 
ordinoides 
proximus 
radix 

rufipunctatus 
sauritus 
scalaris 
sirtalis  A 
sirtalis  B 

Tr.   lineatum  A 
lineatum  B 

V.     striatula 
valeriae 


and  unexpected,  discovery  revealed  by  our 
study.  In  almost  every  instance  the  organ 
positions  in  T.  proximus  and  T.  sauritus 
are  posterior  to  those  in  all  other  Thamno- 
phis. In  the  cases  of  the  posterior  end  of 


Figure  17.     Left  kidney  length  in  thamnophiine  snakes  (expressed  as  a  "/o  of  total  ventrals). 


No.  2 


Visceral  Topography  of  Snakes 


155 


the  heart  and  the  anterior  end  of  the  liver, 
the  ribbon  snakes  share  the  phenomenon 
of  posterior  displacement  with  T.  melano- 
gaster  and  T.  rufipunctatus,  but  in  all 
other  positional  characters  they  stand 
alone  within  the  genus  —  including  pos- 
session of  the  highest  liver-gall  bladder  in- 
terspace/gall bladder  length  values  of  any 
thamnophiine  (Table  VI).  They  also  differ 


from  their  congeners  in  having  a  relatively 
short  left  kidney.  That  the  relatively  long, 
slender-bodied  ribbon  snakes  should  be 
more  similar  to  the  stout-bodied  water 
snakes  (Nerodia),  and  especially  to  the 
short,  semifossorial  genera  (Clonophis, 
Seminatrix,  Storeria,  Tropidoclonion,  Vir- 
ginia), than  to  the  other  Thamnophis  poses 
a  real  enigma.  Whatever  the  cause  of  the 


%  Females 

4        6        8       10       12 


C. 
N. 


kirtlandii 
cyclop ion 
erythrogaster 


fasciata 

rhombifera 

sipedon 

R.  alleni 
grahami  i 
rigida 
septemvi ttata 

Se.  pygaea 
St.  dekayi 

occipitomaculata 

Th.  chrysocephalus 
couchii  A 
couchii  B 
cyrtopsis 
elegans  A 
elegans  B 
eques 
godmani 
marcianus 

melanogaster 

nigronuchalis 

proximus 

radix 

rufipunctatus 

sauritus 

scalaris 

sirtalis  A 

sirtalis  B 

Tr.  lineatum  B 

V.     striatula 
valeriae 


do 


15 


1+^    ! 


6    I 


■13       I 


-U-^    I 


Figure  18.     Heart-liver  interspace  in  thamnophiine  snakes  (expressed  as  a  %  of  total  ventrals). 


156 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


similarities,  it  certainly  does  not  seem  to  be 
due  to  either  phyletic  affinity  or  ecological 
convergence.  All  we  can  reasonably  con- 
clude is  that  T.  proximus  and  T.  sauritus 


are  unique  among  the  garter  snakes.  On 
the  basis  of  the  discriminant  analysis  and 
Student's  t-test  (Table  IX),  we  would  also 
conclude    that    the    other    three    species 


%  Females 

-2         0  2  i  6 


%  Males 

10      12     -4        2        0        2        4        6        8       10      12      14 


C.      kirtlandii 

N.     cyclopion 

erythrogaster 
fasciata 
rhombifera 
sipedon 
val ida 

R.  alleni 
grahamii 
rigida 
septemvittata 

Se.  pygaea 
St.  dekayi 

occipi tomaculata 

Th.  brachystoma 
butleri 
chrysocephalus 

couchii  A 

couchii  B 

cyrtopsis 

elegans  A 

elegans  B 

eques 

godmani 

marcianus 

melanogaster 

nigronuchalis 

ordinoides 

proximus 

radix 

rufipunctatus 

sauritus 

scalaris 

sirtalis  A 
sirtalis  B 

Tr.  lineatum  A 
lineatum  B 

V.     striatula 
valeriae 


Figure  19.     Kidney  overlap  in  thamnophiine  snakes  (expressed  as  a  <Vo  of  total  ventrals). 


No.  2 


Visceral  Tonography  of  Snakes 


157 


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158 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


groups  designated  by  Ruthven  (1908)  do 
not  appear  to  be  distinguishable  from  one 
another  solely  on  the  basis  of  visceral  topo- 
graphy. 

As  was  implied  above,  in  many  cases  the 
small,  semifossorial  thamnophiines  tend  to 
have  a  posterior  displacement  of  organs,  a 
condition  they  share  frequently  with  the 


ribbon  snakes  (Thamnophis  proximus,  T. 
sauritus)  and  occasionally  with  some 
species  of  Nerodia  and  Regina.  Posterior 
displacement  is  a  general  trend,  not  an  in- 
variable phenomenon,  and  both  inter-  and 
intrageneric  variation  occur  from  one 
character  to  the  next.  The  semifossorial 
genera  also  show  a  definite  trend  toward 


Table  IX.    Significance  of  Ruthven' s  Thamnophis   groups  compared  as  four  se  irate 
populations.   NS  indicates  difference  not  significant  at  p>.05. 


Character 

Sex 

Sauritus- 
Radix 

Sauritus - 
Elegans 

Sauritus - 
Sirtalis 

Radix- 
Elegans 

Radix- 
Sirtalis 

Elegans - 
Sirtalis 

post,  heart 

9 

.001^ 
.01 

.001 
.001 

.001 
.001 

NS 
NS 

NS 

NS 

NS 
NS 

ant.  liver 

d 
9 

.001 
.001 

.001 
.001 

.001 
.001 

.05 
.05 

.05 
.001 

NS 
NS 

post,  liver 

d 
9 

.001 
.001 

.001 
.001 

.001 
.001 

NS 
NS 

NS 
NS 

NS 
.05 

post,  gall  bladder 

d 
9 

.001 
.001 

.001 
.001 

.001 
.001 

NS 
NS 

NS 
NS 

NS 
NS 

post,  pancreas 

cf 

9 

.001 
.001 

.001 
.001 

.001 
.001 

NS 
NS 

NS 
NS 

NS 
NS 

ant.  r.  kidney 

d 
9 

.001 
.01 

.001 
.001 

.001 
.001 

.001 
.001 

.001 
.01 

.01 

NS 

post .  1.  kidney 

d 
9 

.001 
.001 

.001 
.001 

.001 
.001 

.001 
.001 

.PI 
.dOl 

NS 
NS 

ant.  1  kidney 

d 
9 

.001 
.001 

.001 
.001 

.001 
.001 

.001 
.001 

.00- 
.001 

.01 

NS 

post  1.  kidney 

d 
9 

.001 
.001 

.001 
.001 

.001 
.001 

.001 
.001 

.001 
.001 

.001 

NS 

liver  length 

d 
9 

NS 

NS 

.02 

NS 

.01 

NS 

NS 

NS 

NS 
.02 

NS 
NS 

r.  kidney  length 

d 
9 

.01 
.001 

.01 
.001 

.01 
.001 

NS 
.01 

NS 
NS 

NS 
NS 

1.  kidney  length 

d 
9 

.001 
.001 

.001 
.001 

.001 
.001 

NS 
NS 

NS 
NS 

NS 
NS 

heart-liver 
interspace 

cC 

9 

.02 

NS 

.01 

NS 

NS 

NS 

NS 
NS 

VS 
NS 

NS 
NS 

kidney  overlap 

d 

9 

NS 
.001 

NS 
.01 

NS 
.001 

NS 
NS 

NS 

K: 

NS 
NS 

Significance  levels  determined  using  2-tailed  Student's  t-test. 


No.  2 


Visceral  Topography  of  Snakes 


159 


having  relatively  short  kidneys,  but  the 
data  for  Storeria  are  equivocal  and  the 
characteristic  is  not  unique  to  those 
genera.  One  unique  feature  that  is  shared 
by  all  of  the  semifossorial  genera  is  the 
possession  of  a  relatively  long  liver.  We  do 
not  know  why  small  snakes  would  possess 
a  proportionally  longer  liver  than  large 
snakes,  but  perhaps  there  are  physiological 
constraints  that  prevent  the  mutual  reduc- 
tion of  body  and  of  liver  from  being 
directly  proportional  —  perhaps  a  mini- 
mum quantity  of  liver  tissue  is  required  for 
the  proper  functioning  of  that  organ. 

Acknowledgments 

For  the  loan  of  specimens  and  for  other 
courtesies,  we  are  indebted  to  the  follow- 
ing curators:  Charles  W.  Myers  and 
Richard  G.  Zweifel  (American  Museum  of 
Natural  History);  Harry  A.  Shankman 
(Arizona  State  University);  Douglas  C. 
Cox  (Brigham  Young  University);  C.J. 
McCoy,  Jr.  (Carnegie  Museum);  Hymen 
Marx  (Field  Museum  of  Natural  History); 
Walter  Auffenberg  and  Peter  Meylan 
(Florida  State  Museum);  Kenneth  CHffer 
and  Philip  J.  Regal  (James  Ford  Bell 
Museum  of  Natural  History);  Joseph  T. 
Collins  and  William  E.  Duellman  (Uni- 
versity of  Kansas  Museum  of  Natural 
History);  Gloria  Z.  Wurst  and  David  B. 
Wake  (Museum  of  Vertebrate  Zoology, 
University  of  California  at  Berkeley); 
Harold  A.  Dundee  (Tulane  University);  T. 
Paul  Maslin  (University  of  Colorado 
Museum  of  Natural  History);  Gary  Brei- 
tenbach  and  Arnold  G.  Kluge  (University 
of  Michigan  Museum  of  Zoology);  James 

F.  Jackson  (University  of  Southwestern 
Louisiana);  Jonathan  A.  Campbell  (Uni- 
versity of  Texas  at  Arlington);  and  Robert 

G.  Webb  (University  of  Texas  at  El  Paso). 
We  are  also  grateful  to  Darrel  R.  Frost, 
Mark  S.  Hafner,  Dominique  G.  Homber- 
ger,  and  Randy  H.  Vaeth  for  helpful  sug- 
gestions at  various  stages  in  the  develop- 
ment of  this  manuscript. 

Specimens  Examined' 

Clonophis    kirtlandii.     ILLINOIS,    Christian    Co.: 
LSUMZ  40065;  Cook  Co.:  FMNH  23166,  25437; 


/Gross  Pt.y:  FMNH  2989;  Will  Co.:  FMNH  55562, 
65902.  INDIANA,  Delaware  Co.:  FMNH  64670; 
Porter  Co.:  FMNH  42069;  /Orange  Co.?y  FMNH 
3060.  KENTUCKY,  Jefferson  Co.:  FMNH  25535. 
OHIO,  Hamilton  Co.:  LSUMZ  7445,  13539. 

Nerodia  cyclopion.  LOUISIANA,  Ascension  Par.: 
LSUMZ  13703;  Calcasieu  Par.:  LSUMZ  12150; 
Cameron  Par.:  LSUMZ  18671-2;  Iberville  Par.: 
LSUMZ  18286,  20703,  24669;  Jefferson  Par.: 
LSUMZ  8670,  13704;  Lafourche  Par.:  LSUMZ 
13557,  19183;  St.  Bernard  Par.:  LSUMZ  9280;  St. 
Charles  Par.:  LSUMZ  18757,  29355;  St.  James 
Par.:  LSUMZ  18293,  19174;  St.  Tammany  Par.: 
LSUMZ  34308;  Vermilion  Par.:  LSUMZ  24025, 
33939. 

Nerodia  erythrogaster.  LOUISIANA,  Acadia  Par.: 
LSUMZ  20310;  Cameron  Par.:  LSUMZ  20344; 
East  Baton  Rouge  Par.:  LSUMZ  17321,  17702, 
19175,  20312,  20723,  22909,  24028;  Iberville  Par.: 
LSUMZ  18287,  22558-9;  Jefferson  Par.:  LSUMZ 
18716;  Livingston  Par.:  LSUMZ  28812;  St.  Ber- 
nard Par.:  LSUMZ  8992;  St.  John  the  Baptist  Par.: 
LSUMZ  23864;  St.  Tammany  Par.:  LSUMZ 
12983,  20279;  Vermilion  Par.:  LSUMZ  34295; 
Washington  Par.:  LSUMZ  12540;  West  Baton 
Rouge  Par.:  LSUMZ  11887;  West  FeHciana  Par.: 
LSUMZ  18758. 

Nerodia  fasciata.  LOUISIANA,  Ascension  Par.: 
LSUMZ  17698;  Cameron  Par.:  LSUMZ  12731, 
17315,  20281,  28666;  Jefferson  Par.:  LSUMZ  8947, 
8953;  Natchitoches  Par.:  LSUMZ  30410;  Plaque- 
mines Par.:  LSUMZ  8653;  Pointe  Coupee  Par.: 
LSUMZ  20274;  St.  Charles  Par.:  LSUMZ  7142, 
7527;  St.  Landry  Par.:  LSUMZ  18113,  18122;  St. 
Martin  Par.:  LSUMZ  19171,  19173. 

Nerodia  rhombifera.  LOUISIANA,  East  Baton 
Rouge  Par.:  LSUMZ  17687,  17794,  17945,  20799, 
23662,  28008-10;  Iberville  Par.:  LSUMZ  13756;  St. 
Charles  Par.:  LSUMZ  9216. 

Nerodia  sipedon.  ALABAMA,  Jackson  Co.: 
LSUMZ  36375;  Pickens  Co.:  LSUMZ  36399, 
36400.  ILLINOIS,  Jackson  Co.:  LSUMZ  27610; 
Pope  Co.:  LSUMZ  27599.  MISSISSIPPI,  Greene 
Co.:  LSUMZ  36379,  36381-3,  36385,  36387, 
36390-3,  36396-7;  Lauderdale  Co.:  LSUMZ 
36403-4;  Wilkinson  Co.:  LSUMZ  28712. 
MISSOURI,  Lawrence  Co.:  LSUMZ  9107. 

Nerodia  valida.  MEXICO,  Colima:  LSUMZ  7876; 
Nayarit:  LSUMZ  33099,  36266,  36268;  Sinaloa: 
AMNH  36269,  84077,  84080-2,  87575,  87577, 
88889-90,  88892;  Sonora:  AMNH  84074-6. 

Regina  alleni.  FLORIDA,  Alachua  Co.:  FSM  2476, 
2498,  6634,  6637,  7171,  9096,  LSUMZ  13618-9; 
Collier  Co.:  LSUMZ  28992;  Dade  Co.:  FSM 
42527;  Dixie  Co.:  LSUMZ  7473;  Hillsborough  Co.: 
FSM   42529;    Indian    River   Co.:    FSM   42524-6, 


160 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


42530;  Polk  Co.:  FSM  1868;  Sumter  Co.:  FSM 
11157. 
Regina  grahamii.  LOUISIANA,  East  Baton  Rouge 
Par.:  LSUMZ  17947,  33460,  USL  7623;  Iberville 
Par.:  LSUMZ  20271;  Lafayette  Par.:  USL  20945; 
St.  Landry  Par.:  LSUMZ  28665,  USL  15936, 
23236,  23414,  23427;  St.  Martin  Par.:  USL  22953, 
24432;  Terrebonne  Par.:  LSUMZ  36484-7;  Vermi- 
lion Par.:  USL  10687,  17353.  TEXAS,  Chambers 
Co.:  LSUMZ  33462. 

Regina  rigida.  NO  DATA:  USL  6067,  8820.  LOUISI- 
ANA, Iberville  Par.:  LSUMZ  22556;  Lafayette 
Par.:  USL  24245;  Natchitoches  Par.:  LSUMZ 
12988;  Orleans  Par.:  LSUMZ  8982-3;  Sabine  Par.: 
USL  24453;  St.  Charles  Par.:  LSUMZ  8680;  St. 
Landry  Par.:  USL  15930,  17620;  St.  Martin  Par.: 
USL  14365,  19471,  22425,  24433;  Terrebonne  Par.: 
LSUMZ  36483. 

Regina  septemvittata.  ALABAMA,  Baldwin  Co.: 
LSUMZ  15783.  NORTH  CAROLINA,  Orange 
Co.:  LSUMZ  14353^.  OHIO,  Montgomery  Co.: 
LSUMZ  24476,  30184-5.  TENNESSEE,  Clay  Co.: 
LSUMZ  34795;  Jackson  Co.:  LSUMZ  34798. 

Seminatrix  pygaea.  FLORIDA,  Alachua  Co.:  FSM 
9813  (-6,-12),  14146  (-4),  14147  (-1,-7),  14215  (-4), 
14216  (-2,-4,-9),  14217  (-3,-5,-7),  14218  (-4,-6); 
Dade  Co.:  LSUMZ  6530,  7405,  24582. 

Storeria  dekayi.  LOUISIANA,  Ascension  Par.: 
LSUMZ  18776;  Cameron  Par.:  LSUMZ  2764, 
12196,  18168-70,  24038,  28819-20,  28822,  29977, 
32649;  Iberia  Par.:  LSUMZ  2771;  Iberville  Par.: 
LSUMZ  12229,  23877;  St.  Landry  Par.:  LSUMZ 
18665,  20074;  Vermilion  Par.:  LSUMZ  24733. 

Storeria  occipitomaculata.  LOUISIANA,  Claiborne 
Par.:  LSUMZ  24658;  East  Feliciana  Par.:  LSUMZ 
16686;  Natchitoches  Par.:  LSUMZ  24745,  33077-8; 
West  Feliciana  Par.:  LSUMZ  12602,  17898. 

Thamnophis  brachystoma.  PENNSYLVANIA,  Clar- 
ion Co.:  CM  28292-3,  28295,  28297-9,  28302-3, 
28306-9,  28311,  28313,  28317-8,  28320-1. 

Thamnophis  butleri.  CANADA,  Ontario:  UMMZ 
90193.  INDIANA,  Noble  Co.:  UMMZ  132822. 
OHIO,  Lucas  Co.:  UMMZ  68864,  99627(3). 
MICHIGAN,  Sanilac  Co.:  UMMZ  98774;  Wash- 
tenaw Co.:  UMMZ  465234;  Wayne  Co.:  UMMZ 
89519.  WISCONSIN,  Waukesha  Co.:  AMNH 
76178-80. 

Thamnophis  chrysocephalus.  MEXICO,  Guerrero: 
AMNH  72500-1,  72503;  Oaxaca,  AMNH  91094-5, 
93235,  97855-6,  97865-6,  97868-9,  97871. 

Thamnophis  couchii  couchii.  CALIFORNIA,  Ama- 
dor Co.:  LSUMZ  16530,  16544;  Kern  Co.:  LSUMZ 
16549;  Shasta  Co.:  LSUMZ  22938,  34587-8,  34590, 
MVZ  18824-5;  Tehama  Co.:  LSUMZ  16550; 
Tulare  Co.:  LSUMZ  16547;  Tuolumne  Co.: 
LSUMZ  34585. 


Thamnophis  couchii  hydrophilus.  CALIFORNIA, 
Humboldt  Co.:  LSUMZ  34578;  Shasta  Co.: 
LSUMZ  1655M;  Trinity  Co.:  LSUMZ  34594-5. 
OREGON,  Jackson  Co.:  LSUMZ  16560-4,  16567. 

Thamnophis  cyrtopsis.  ARIZONA,  Coconino  Co.: 
LSUMZ  29940,  30062,  30083,  30088;  Gila  Co.: 
LSUMZ  30061;  Maricopa  Co.:  LSUMZ  30063, 
30081;  Pima  Co.:  LSUMZ  30066,  30090;  Santa 
Cruz  Co.:  LSUMZ  10035,  30072,  30076-7;  Yavapai 
Co.:  LSUMZ  29943,  29945-6,  29948,  30064-5, 
30067-8. 

Thamnophis  eiegans  terrestris.  CALIFORNIA, 
Mendocino  Co.:  LSUMZ  34378,  34380;  San  Mateo 
Co.:  LSUMZ  7922,  16502-3,  16507,  34371,  34373; 
Sonoma  Co.:  LSUMZ  34368-9,  34374-5;  Sonoma- 
Mendocino  Co.:  LSUMZ  34367. 

Thamnophis  eiegans  vagrans.  NO  DATA:  LSUMZ 
20747-50.  ARIZONA,  Coconino  Co.:  LSUMZ 
29957,  29959-62.  COLORADO,  Conejos  Co.: 
LSUMZ  11571,  11609,  11611,  11615,  30051,  30055; 
Costilla  Co.:  LSUMZ  7985,  11603-5,  11607,  11614, 
11618,  13929,  13931-2,  30050;  Rio  Grande  Co.: 
LSUMZ  30056. 

Thamnophis  eques  megalops.  MEXICO,  Chihuahua: 
AMNH  104471,  104772,  BYU  22701;  San  Luis 
PotosK  LSUMZ  4374-5,  4879. 

Thamnophis  eques  virgatenuis.  MEXICO,  Durango: 
AMNH  102521,  LSUMZ  16424-6,  16429-30. 

Thamnophis  godmani.  MEXICO,  Oaxaca:  AMNH 
89604,  91101-2,  91105,  97853,  97873-4,  97884, 
97888,  103090,  103092-5,  103101,  103103,  103105, 
103113,  104394,  106993,  106995-8,  107002-5, 
718170. 

Thamnophis    marcianus.     TEXAS,     Bexar    Co. 
LSUMZ  10315;  Duval  Co.:  LSUMZ  23239,  23243 
Hartley   Co.:    LSUMZ    10407;    Jeff   Davis   Co. 
LSUMZ  29608;  McMuUen  Co.:   LSUMZ  23248 
Moore  Co.:  LSUMZ  10365;  Presidio  Co.:  LSUMZ 
23255;  San  Patricio  Co.:  LSUMZ  23249,  23252; 
Webb  Co.:  LSUMZ  30929;  Zavala  Co.:  LSUMZ 
23254. 

Thamnophis  melanogaster.  MEXICO,  Jalisco: 
LSUMZ  16434;  Michoaca'n:  LSUMZ  14489-90, 
14492-3,  16435,  34346,  36277,  36279-80,  36282-6. 

Thamnophis  nigronuchalis.  MEXICO,  Durango: 
LSUMZ  11637,  16448,  16450-5,  16459-60,  UTEP 
3386-7. 

Thamnophis  ordinoides.  CALIFORNIA,  Del  Norte 
Co.:  MVZ  30276-7,  30279.  OREGON,  Clatsop 
Co.:  MVZ  34265-8,  36848;  Polk  Co.:  MVZ  24808; 
TUlamook  Co.:  MVZ  47856.  WASHINGTON, 
Clark  Co.:  MVZ  34259;  King  Co.:  MVZ  38653, 
38655,  38657,  38670,  38674;  Lewis  Co.:  MVZ 
70366;  Pacific  Co.:  MVZ  34262. 

Thamnophis  proximus.  LOUISIANA,  Acadia  Par.: 
LSUMZ   17899;   Cameron  Par.:   LSUMZ  33964; 


No.  2 


Visceral  Topography  of  Snakes 


161 


Claiborne  Par.:  LSUMZ  33966;  East  Baton  Rouge 
Par.:  LSUMZ  16912,  18714,  20254;  Iberia  Par.: 
LSUMZ  18077;  Iberville  Par.:  LSUMZ  20255, 
20316,  22548;  Livingston  Par.:  LSUMZ  7960, 
18974;  Pointe  Coupee  Par.:  LSUMZ  20220;  St. 
Tammany  Par.;  LSUMZ  7934;  Vermilion  Fax.: 
LSUMZ  24052.  TEXAS,  Hidalgo  Co.:  LSUMZ 
18621-3. 

Thamnophis  radix.  NO  DATA:  LSUMZ  20735^0, 
20742-5.  COLORADO,  Denver  Co.:  LSUMZ 
7465;  Larimer  Co.:  UC  31837^0,  31842-3,  31847, 
31851,  31873,  31888.  ILLINOIS,  Iroquois  Co.: 
LSUMZ  8126.  NEW  MEXICO,  San  Miguel  Co.: 
LSUMZ  7942,  7944,  7972. 

Thamnophis  rufipunctatus.  ARIZONA,  Coconino 
Co.:  LSUMZ  uncatalogued,  LSUMZ  36815. 
MEXICO,  Chihuahua:  AMNH  4342,  68286,  ASU 
17042,  5304-5,  5335,  UTEP  2043,  2262,  3657. 

Thamnophis  sauritus.  FLORIDA,  Alachua  Co.:  FSM 
14183,  14550  (-1),  14550  (-2),  14550  (-3),  14550  (-4), 
14550  (-7),  14550  (-8),  14550  (-9),  14551  (-2),  14551 
{,-4),  39197;  Collier  Co.:  FSM  39198,  39200-2;  Dade 
Co.:  FSM  22874,  39204-5;  Franklin  Co.:  LSUMZ 
21805-6,  21810;  Pasco  Co.:  LSUMZ  22003.  LOUI- 
SIANA, St.  Tammany  Par.:  LSUMZ  8302,  23770. 

Thamnophis  scalaris.  MEXICO,  Distrito  Federal: 
AMNH  75934;  JaUsco:  UTA  R-4932,  R^949, 
5991,  5993;  Mexico:  AMNH  71315  (2),  94714; 
Michoacan:  AMNH  88724. 

Thamnophis  sirtalis  fitchi.  CALIFORNIA,  Amador 
Co.:  LSUMZ  16486-8,  16489-92;  Mendocino  Co.: 
LSUMZ  16493;  Modoc  Co.:  LSUMZ  8215;  Plumas 
Co.:  LSUMZ  16477-8,  16481-2;  Shasta  Co.: 
LSUMZ  16496-8. 

Thamnophis  sirtalis  sirtalis.  INDIANA,  Allen  Co.: 
LSUMZ  7988.  MINNESOTA,  Carlton  Co.:  JFBM 
1115,  Cass  Co.:  LSUMZ  7991,  7996;  Clearwater 
Co.:  JFBM  2644-5,  2651-2,  2657,  2659;  Isanti  Co.: 
LSUMZ  23229,  23232,  23234,  24461-2;  Pine  Co.: 
LSUMZ  23230. 

Tropidoclonion  lineatum.  NEBRASKA,  Jefferson 
Co.:  KU  45252-65,  45267-8;  Richardson  Co.:  KU 
52228.  NEW  MEXICO,  San  Miguel  Co.:  LSUMZ 
29998-9,  30096-7.  TEXAS,  Travis  Co.:  LSUMZ 
20078-9. 

Virginia  striatula.  NO  DATA:  USL  5395,  15841. 
LOUISIANA,  Acadia  Par.:  LSUMZ  12091;  Ascen- 
sion Par.:  LSUMZ  12087,  18777;  Caddo  Par.: 
LSUMZ  20210;  East  Baton  Rouge  Par.:  LSUMZ 
1598,  1604-5,  2786,  17348,  18712,  23536,  23745; 
East  Feliciana  Par.:  LSUMZ  2779;  Lafayette  Par.: 
USL  11179,  22890;  Livingston  Par.:  LSUMZ 
12126;  Sabine  Par.:  LSUMZ  20193;  St.  Helena 
Par.:  LSUMZ  18360;  St.  Landry  Par.:  USL  18277; 
St.  Tammany  Par.:  LSUMZ  2773. 


Virginia  valeriae.  FLORIDA,  Alachua  Co.:  FSM 
42545;  Leon  Co.:  FSM  1942,  34858;  Liberty  Co.: 
FSM  42531-2,  42534-5;  Wakulla  Co.:  FSM  32991. 
GEORGIA,  Chattahoochee  Co.:  FSM  42546. 
LOUISIANA,  Bossier  Par.:  LSUMZ  24656;  Caddo 
Par.:  LSUMZ  12094;  East  Baton  Rouge  Par.: 
LSUMZ  12147,  17671;  East  Feliciana  Par.: 
LSUMZ  15536;  Livingston  Par.:  LSUMZ  20256; 
St.  Helena  Par.:  TU  5957;  St.  Tammany  Par.:  TU 
1 1844,  14238,  18395;  Webster  Par.:  LSUMZ  12142; 
West  Feliciana  Par.:  LSUMZ  17901.  MISSISSIP- 
PI, Hancock  Co.:  TU  14304,  15056,  17681. 

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Appendix  A 
Comparative  data  on  non-thamnophiine 
snakes  obtained  from  the  literature  fell 
within  the  outer  parameters  of  the  thamno- 
phiine  data  sets  generated  by  our  study 
except  for  the  following  taxa  whose  organ 


positions  lie  more  posteriorly  or  which 
have  longer  organs  or  interspaces. 
Posterior  end   of  heart  —  non-natricine 
and    non-homalopsine    Colubridae: 
Boiga,^      Chamaetortus,^    Coluber 
(  =  Gonyosoma)  oxycephalus,  ^ 

Dipsadoboa, '  and  male  Zamenis 
rhodorhacis;^  Acrochordidae:  Acro- 
chordus  arafurae,^  A.  granulatus,^  A. 
javanicus;^  Boidae:  male  Bolyeria/  male 
Corallus,*  rnale  Eunectes,*  mdXt  Licha- 
nura,*  male  Loxocemus,* male  Xenc- 
peltis;*  Viperidae: C<2W5W5  rhombeatus.^ 

Anterior  end  of  right  kidney  —  non-natri- 
cine and  non-homalopsine  Colubridae: 
Coluber  (  =  Gonyosoma)  oxycephalus,^ 
female  Philothamnus  semivariegatus,^ 
male  Psammophis  sibilans,^  male 
Zamenis  florulentus,^  Z.  rhodorhacis? 
Posterior  end  of  right  kidney  —  non-natri- 
cine and  non-homalopsine  Colubridae: 
Coluber  ( =  Gonyosoma)  oxycephalus,  ^ 
male  Leptodira  (  =  Crotaphopeltis) 
hotamboeia,^  female  Philothamnus 
semivariegatus,^  male  Psammophis  sibi- 
lans;^  Viperidae:  male  Causus 
rhombeatus.^ 
Anterior  end  of  left  kidney  —  non-natri- 
cine and  non-homalopsine  Colubridae: 
male  Coluber  {  =  Gonyosoma)  oxyce- 
phalus,^ female  Philothamnus  semi- 
variegatus,^  male  Psammophis  sibilans,^ 
male  Zamenis  florulentus.^ 
Posterior  end  of  left  kidney  —  non-natri- 
cine and  non-homalopsine  Colubridae: 
male  Coluber  {  =  Gonyosoma)  oxyce- 
phalus,^ female  Philothamnus  semiva- 
riegatus,^  male  Psammophis  sibilans.^ 
Right  kidney  length  —  Viperidae:  Causus 

rhombeatus.^ 
Heart-liver  interspace  —  The  following 
taxa  had  an  overlap  —  Tropidophiidae: 
Trachyboa  gularis,^  Tropidophis;^ 
Viperidae:  Causus  rhombeatus.^ 
Kidney  overlap  —  all  taxa  reported  in  the 
literature  have  an  overlap,  but  Causus 


164 


Tulane  Studies  in  Zoology  and  Botany 


Vol.  23 


rhombeatus^  (Viperidae)  is  the  only  one 
to  have  a  greater  overlap  than  any  of  the 
Thamnophiini. 

The  following  taxa  have  an  organ  posi- 
tion lying  more  anteriorly  or  have  shorter 
organs  than  any  of  the  Thamnophiini. 
Posterior  end  of  left  kidney  —  Tropido- 

phiidae:  female  Exiliboa  placata.' 
Liver  length  —  non-natricine  and  non- 
homalopsine     Colubridae:     female 
Philothamnus  semivariegatus,^ 
Heart-liver  interspace  —  Colubridae,  Na- 
tricinae:    male   Natrix  (  =  Amphiesma) 
vibakari^  from  Japan. 
Kidney  asymmetry  —  In  the  present  study 
males  in  64°7o  of  the  taxa  have  the  left 
kidney  longer  than  the  right  but  the  dif- 
ference is  significant  in  only  5"7o.  How- 
ever, the  literature  reveals  that  in  the 


'Rasmussen  (1979) 
Thompson  (1914) 
'McDowell  (1979) 
'Underwood  (1976) 
'Thompson  (1913b) 
'Brongersma  (1951) 
'Bogert  (1968) 
'Bergman  (1959b) 
'Bergman  (1955e) 
'"Bergman  (1956-58) 
"Bergman  (1960) 


males  of  most  taxa  the  right  kidney  is 
longer  than  the  left.  The  following  are 
the  taxa  in  which  the  left  kidney  is 
longer:  Colubridae,  Natricinae  —  Natrix 
( =  Sinonatrix)  trianguligera;*  Colu- 
bridae, Homalopsinae:  Enhydris  enhy- 
dris;"  other  Colubridae:  Coluber 
(  =  Gonyosoma)  oxycephalus,^ 

Elapoides  fuscus.'"  Females  in  76%  of 
the  thamnophiines  have  the  right  kidney 
longer  than  the  left  (28%  significantly 
different)  as  do  the  females  of  all  taxa 
reported  in  the  literature  except: 
Colubridae,  Natricinae  —  Natrix 
(=Amphiesma)  vibakari;^  Colubridae, 
Homalopsinae:  Hypsirhina  {  =  Enhy- 
dris) plumbea;''  other  Colubridae: 
Elapoides  fuscus. ' " 


December  15,  1982 


087