(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Biodiversity Heritage Library | Children's Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Tulane studies in zoology and botany"

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 
in CBE Style Manual, 4th ed., 1978. Manuscripts should be typewritten and double 
spaced. Two additional copies should accompany the original to expedite editing and 
publication. Legends for figures should appear on a separate page and in sequence. 
Illustrations should be proportioned for one or two column width corresponding to our 
printed page size, and should allow for insertion of the legend if occupying a whole 
page. Guidelines for letter and other extraneous markings should be done with a 
non-photo blue pencil such as Eagle Prismacolor. Photographs should be on glossy 
paper. 

Many tables, if carefully prepared with a carbon ribbon and electric typewriter, can be 
photographically reproduced, thus helping to reduce publication costs. Lettering in any 
illustrative or tabular material should be of such a size that no letter will be less than 1 Vi 
mm high when reduced for publication. 

An abstract not exceeding three percent of the length of the article must accompany the 
manuscript. 

Separates of published articles are available to authors at a nominal cost. 

Page charges, calculated at $45/page, are solicited from authors who have funds for this 
purpose through their institutions or grants. Acceptance of papers is not dependent on 
ability to underwrite costs but excessive illustrations and tabular matter may be charged 
to the author. 

EXCHANGES, SUBSCRIPTIONS, ORDERS FOR INDIVIDUAL COPIES: Ex- 
changes are invited from institutions publishing comparable series. Subscriptions are 
billed in advance. A price list of back issues is available on request. Individuals should 
send th^ir remittance, preferably money order, along with their orders. Remittances 
should be made payable to "Tulane University." Subscription rates: 

Volume 23. $8.50 domestic, $9.50 foreign. 

Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the 
mails, will be replaced if the editorial offices are notified before the second subsequent 
issue is released. 

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 



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 
in CBE Style Manual, 4th ed., 1978. Manuscripts should be typewritten and double 
spaced. Two additional copies should accompany the original to expedite editing and 
publication. Legends for figures should appear on a separate page and in sequence. 
Illustrations should be proportioned for one or two column width corresponding to our 
printed page size, and should allow for insertion of the legend if occupying a whole 
page. Guidelines for letter and other extraneous markings should be done with a 
non-photo blue pencil such as Eagle Prismacolor. Photographs should be on glossy 
paper. 

Many tables, if carefully prepared with a carbon ribbon and electric typewriter, can be 
photographically reproduced, thus helping to reduce publication costs. Lettering in any 
illustrative or tabular material should be of such a size that no letter will be less than 1 Vi 
mm high when reduced for publication. 

An abstract not exceeding three percent of the length of the article must accompany the 
manuscript. 

Separates of published articles are available to authors at a nominal cost. 

Page charges, calculated at $45/page, are solicited from authors who have funds for this 
purpose through their institutions or grants. Acceptance of papers is not dependent on 
ability to underwrite costs but excessive illustrations and tabular matter may be charged 
to the author. 

EXCHANGES, SUBSCRIPTIONS, ORDERS FOR INDIVIDUAL COPIES: Ex- 
changes are invited from institutions publishing comparable series. Subscriptions are 
billed in advance. A price list of back issues is available on request. Individuals should 
send their remittance, preferably money order, along with their orders. Remittances 
should be made payable to "Tulane University." Subscription rates: 

Volume 23. $8.50 domestic, $9.50 foreign. 

Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the 
mails, will be replaced if the editorial offices are notified before the second subsequent 
issue is released. 

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 



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- 



12 



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 



^ 








."s* 








"•*3 








3 




g 




c 




o 




TS 




a> 




CO 




cd 




X> 




CO 




(U 




J3 




CJ 




•<-> 




^ 








"o 




^.^ 




cd 




3 




C 




C 




cd 


• 


Cm 


c 


o 


cd 


(L> 


a 




^ 

o^ 


3 




C 




T3 


CO 


^ 


kH 


6 






a 


CO 


aj 


w 


(U 




00 


^ 


c 


.1 




•^ 




-^ 


*S 


s; 


J3 


o 


o 


5 


3 


5J 
to 


U 


O 


• 


s: 


CO 


^ 


^ 






Lh 




O 


O 


Cm 


C4-C 


c 


73 


o 


!U 




00 


5 




3 


"c 


a 


a> 


o 


C4-1 



>» 


CO 


x> 


■t-> 




0) 


cS 


CO 


-!-> 




c« 


IZ) 


T3 


_D 


(L» 


a 


> 


CTJ 


%-> 


a> 


O 


-<-> 


3 


^ 


-a 




o 


c« 


u> 


l-l 


O. 





u 


0, 


oi 










<N 






C4-I 


(U 









X> 


CO 


cd 




H 


CO 



« 3 O 



01 
01 



■g O _ CO 

^ Q> 3 ^ 

E ^ n ii 

tS £ E 3 

w 2 ''" 



« -3 






o « 



3 on 
^ exi 

O 



^ 












u 


ba 


3 


R 


^^ 


O) 


,^ 


>> 


CO 




_aj 


w^ 






"« 


V 


E 




C/3 





3 
00 

3 

< 

03 






ON 









00 fs 



O OS 
r- 00 



3 



03 



cd 



OS 



>% 


3 






c^. 


c^- 










00 










1> 






(>• 3 


3 
< 




c^- 






C 






'—1 






_>. 


_>^ 


3 




(U 


>.c"- 




(U 


3 


3 


"-5 




c 


03 


c^- 




C 


■— ) 


'— > 


>, 




3 




c 

3 
I—) 




3 


c3 


03 


03 




— ) 


fS 


'^ 






vO 


ro 


^ 






^ 


r-" 






r-" 


00 


r~-' 






^— < 


1-^ 






^^ 


^^ 


^^ 
























' >< s 


X 


•r^ 


1 


^ w 


X vo 


X OS 


1 




















rsi ^^ 


On 






QO ^^ 


^^ 


00 






OS 


d 






d 


Os" 


_^ 






<N 


m 






m 


<N 


m 







+ + 



00 



(N 






ra 



O rn ^ 



(N rsi ^ 

^' ^ ^ 






so 



+ 






fC^ <N "^ en 



CsJ 



m 









so 



OS 



Os — ' ' _; d 

-^ <N (N <N 








Qi 


Z X 






AGUA] 

STGO 

MEZQ 






G 


S 


S u 


Oh 


N 




CO 


H 


oi Z 


< 


< 


< 


PQ 


C/D 


u u 


Dl- 


z 


00 


CT) 



W 




Q 


ai 


OC 


w 


w 


D 


> 


Q 






m 



4 



4 



N 
H 
< 
CU 






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). 

Literature Cited 

AGASSIZ, L. 1857. Contributions to the natural 
history of the United States of America. Vol. 1 



(Part 1) Essay on Classification, pp. 3-232; (Part 
II) North American Testudinata, pp. 233-450. 
Vol. 2 (Part III) Embryology of the turtles, pp. 
451-643 and plates 1-34. Little, Brown and Co., 
Boston. 

ALBRITTON, C.C., JR. 1958. Quaternary strati- 
graphy of the Guadiana Valley, Durango, Mexico. 
Bull. Geol. Soc. Am. 69:1197-1216. 

ALTINI, G. 1942. I rettili dei Laghi Chapala, Patz- 
cuaro e Peten raccolti nel 1932 dal Prof. Alessan- 
dro Ghigi e dal Prof. Alula Taibel. Atti Soc. Ital. 
Sci. Nat., Milan 81(3-4):153-195. 

ALVAREZ, J. 1949. Contribucidn al conocimiento 
de los peces de la region de los Llanos, Estado de 
Puebla (Mexico). An Esc. Nac. Cienc. Biol., Mex. 
6:81-107. 

. 1963. Ictiologfa Michoacana III. Los 

peces de San Juanico y de Tocumbo, Mich. An. 
Esc. Nac. Cienc. Biol., Mex. 12:111-138. 

. 1972. Ictiologfa Michoacana V. Origen y 



distribucion de la ictiofauna dulceacuicola de 

Michoacan. An. Esc. Nac. Cienc. Biol., Mex. 

19:155-161. 
AMADON, D. 1966. The super species concept. 

Syst. Zool. 15:246-249. 
. 1968. Further remarks on the super 

species concept. Syst. Zool. 17:345-346. 
and LL. SHORT. 1976. Treatment of sub- 



species approaching species status. Syst. Zool. 
25:161-167. 

ARELLANO, A.R.V. 1953. Estratigrafia de la 
Cuenca de Mexico. Mem. Cong. Cien. de Mex. 
3:172-185. 

ASHTON, R.E., S.R. EDWARDS, and G.R. PIS- 
ANI. 1976. Endangered and threatened amphibi- 
ans and reptiles in the United States. SSAR, Misc. 
Publ. Herpet. Circ. 5:1-65. 

ATCHLEY, W.R., C.T. GASKINS, and D. AN- 
DERSON. 1975. Statistical properties of ratios in 
biological data. Amer. Zool. 15:829. 

. 1976. Statistical properties of ratios. I. 

Empirical results. Syst. Zool. 25:137-148. 

AXTELL, R.W. 1978. Ancient playas and their in- 
fluence on the recent herpetofauna of the north- 
ern Chihuahuan desert. U.S. D.I. Natl. Park Serv. 
Trans. Proc. 3:493-512. 

BAIRD, S.F. 1859. Reptiles of the boundary, with 
notes by the naturalists of the survey. U.S. -Mex. 
Boundary Survey (Emory) 3(2): 1-35. 

BARBOUR, CD. 1973. A biogeographical history 
of Chirostoma (Pisces: Atherinidae): A species 
flock from the Mexican Plateau. Copeia 1973: 
533-556. 

BELTZ, R.E. 1954. Notes on the winter behavior of 
captive nonindigenous Chelonia in southern Cali- 
fornia. Herpetologica 10:124. 



66 



Tulune Studies in Zoology and Botany 



Vol.23 



BERNAL, 1. 1959. Tcnochlillan en una isla. Mex. 
Insl. Nac. Antrop. E. Hist. Ser. Historica 2: 1-147. 

BERRY. J.F. 1977. A model for plastral reduction 
in kinosternid turtles (abstract). Program, 57th 
meeting Amer. Soc. Icthyol. and Herpetoi. 

. 1978. Variation and systematics in the 

Kitwsternon scorpioides and A', leucosiomum 
complexes (Reptilia: Testudines: Kinosternidae) 
of Mexico and Central America. Ph.D. disserta- 
tion. Univ. of Utah, Salt Lake City. 

and J.M. LEGLER. 1980. A new mud 



turtle, genus Kinosternon, from Sonora, Mexico. 
Contrib. Sci. Los Angeles Co. Mus. Nat. Hist. 
325:1-12. 
BERRY, J.F. and R. SHINE. 1980. Sexual size di- 
morphism and sexual selection in turtles (Order 
Testudines). Oecologia 44:185-191. 
BLASQUES LOPEZ, L. 1959. Hidrogeologfa de las 
regiones deserticas de Mexico. An Inst. Geol. 
15:1-172. 
BOCOURT, M.-F. 1876. Note sur quelques reptiles 
de I'isthme de Tehuantepec (Mexique) donne's par 
M. Sumichrast au Museum. J. Zool., Paris, 
5(5/6): 386-411. 
BOGERT, CM. and J. A. OLIVER. 1945. A pre- 
liminary analysis of the herpetofauna of Sonora. 
Bull. Amer. Mus. Nat. Hist. 83:297-426. 
BOULENGER, G.A. 1889. Catalogue of the chel- 
onians, rhynchocephalians, and crocodiles in the 
British Museum (Nat. Hist.). Taylor and Francis, 
London. 
BOWLER, J.K. 1977. Longevity of reptiles and 
amphibians in North American collections as of 
November, 1975. SSAR, Misc. Publ. Herpet. 
Circ. 6:1-32. 
BRADBURY, J. P. 1971. Paleoliminology of Lake 
Texcoco, Mexico. Evidence from Diatoms. Lim- 
nology and Oceanography 16:180-200. 
BRAVO-HOLLIS, M. and J. CABALLERO DE- 
LOYA. 1973. Catalogo de la colecci6n helminto- 
logica del Instituto de Biologta. Inst. biol. Univ. 
Nac. Auton. Me'x. Publ. Esp. 2:1-138. 
BRIBIESCA CASTREJON, J.L. 1960. Hidrologfa 
historica del Valle de Me'xico Ing. Hidraul. Mdx. 
14(3):43-62. 
BROWN, B.C. 1950. An annotated check list of the 
reptiles and amphibians of Texas. Baylor Univ. 
Stud., Baylor Univ. Press, Waco, Texas. 
BRYAN, K. 1946. Comentario e intento de corre- 
lacion con la cronologia glacial. Mem. Segundo 
Cong. Me'x. Cienc. Sociales 5:220-225. 

. 1948. Los Suelos complejos y fosiles de la 

altiplanicie de Me'xico, en relacion con cambios 
climalicos. Bol. Soc. Geol. Mex. 13:1-20. 
BUSHNELL, J.H. 1971. Porifera and Ectoprocta 
in Mexico: Architecture and environment of 



Carieriu.s lalilenlus (Spongillidae) and I rederi- 
cella ausiratiensis (Fredericcllidae). Southwestern 
Nat. 15:331-346. 

CABALLERO Y CABALLERO, E. 1938a. Algunos 
trematodos de reptiles de Me'xico. An. Int. Biol. 
Univ. Mex. 9:103-120. 

. 1938b. A new species of Camullanus from 

the stomach of Kinosternon hiriipes, IV. Parasit- 
ology 31:448-450. 

. 1939. Nematodos de los reptiles de Mex- 



ico. V. An. Inst. Biol. Univ. Mex. 10:275-282. 
. 1940a. Revisfon de las especies que actual- 



mente forman el genero Heronimus MacCallum, 
1902 (Trematoda: Heronimidae Ward, 1917). An. 
Inst. Biol. Univ. Mex. 11:225-230. 

.. 1940b. Trematodos de las tortugas de 



Mexico. An. Inst. Biol. Univ. Mex. 11:559-572. 
and M.C. CERECERO (= Zerecero y 



D.). 1943. Nematodos de los reptiles de Mexico. 
VIII. Descripcion de tres nuevas especies. An. 
Inst. Biol. Univ. Mex. 14:527-539. 

CAGLE, F.R. 1957. Reptiles, p. 273-358. In W.F. 
Blair, P. Brodkorb, F.R. Cagle, and G.A. Moore 
(eds.). Vertebrates of the United States. McGraw 
Hill, New York. 

CAMACHO, H. 1925. Las aguas subterraneas del 
Valle de Morelia, Estado de Michoacan. An. Inst. 
Geol. Mex. 2:5-15. 

CAMPBELL, B. 1934. Report on a collection of 
reptiles and amphibians made in Arizona during 
the summer of 1933. Occ. Pap. Mus. Zool. Univ. 
Michigan 289:1-10. 

CARR, A.F. 1952. Handbook of Turtles: the turtles 
of the United States, Canada, and Baja Californ- 
ia. Cornell Univ. Press, Ithaca, NY. 

CASAS ANDREU, G. 1965. Estudio preliminar 
sobre las tortugas de agua dulce en Me'xico. An. 
Inst. Nac. Inves. Biol.-Pesq. 1:365-401. 

1967. ContribuciCn al conocimiento de 

las tortugas dulceacuicolas de Mexico. Mexico, D. 
F., Univ. Nac. Auton. Me'xico, Fac. Ciencias, 
Dept. Biol. 96 pp. 

COCHRAN, D.M. 1961. Type specimens of reptiles 
and amphibians in the United Stales National 
Museum. Bull. U.S. Natl. Mus. 220:1-291. 

and C.J. COIN. 1970. The new field book 

of reptiles and amphibians. C.P. Putnam's Sons, 
New York. 

COLE, G.A. 1963. The American southwest and 
Middle America, p. 393-434. In D.G. Frey (ed). 
Limnology in North America, Univ. of Wisconsin 
Press, Madison. 

and M.C. WHITESIDE. 1965. An ecolog- 
ical reconnaissance of Quitobaquito Spring, 
Arizona. J. Arizona Acad. Sci. 3:1-6. 

CONANT, R. 1963. Semiaquatic snakes of the 



No. 



Kinosternon Biosystematics 



67 



genus Thamnophis from the isolated drainage 
system of the Rio Nazas and adjacent areas in 
Mexico. Copeia 1963:437-499. 
. 1969. A review of the water snakes of the 



genus Natrix in Mexico. Bull. Amer. Mus. Nat. 
Hist. 142:1-140. 
. 1975. A field guide to reptiles and amph- 



ibians of eastern and central North America. 
Second edition. Houghton Mifflin Co., Boston. 
. 1978. Semiaquatic reptiles and amphibi- 



ans of the Chihuahuan Desert and their relation- 
ships to drainage patterns of the Region. U. S.D.I. 
Natl. Park Serv., Trans. Proc. 3:455-492. 
and J.F. BERRY. 1978. Turtles of the 



family Kinosternidae in the southwestern United 
States and adjacent Mexico: identification and 
distribution. Amer. Mus. Novitates 2642:1-18. 

CONTRERAS-BALDERAS, S. 1974. Cambios de 
composicidn de comunidades de peces en zonas 
semiaridas de Me'xico. Publ. Biol. Inst. Inv. 
Cient., UANL. Mex. 1:181-194. 

. 1975. Zoogeography and evolution of 

Notropis lutrensis and "Notropis" ornatus in the 
Rio Grande basin and range, Mexico and United 
States (Pisces:Cyprinidae). Ph.D. dissertation. 
Tulane University, New Orleans. 

COOPER, J.G. 1870. The fauna of California and 
its geographical distribution. Proc. California 
Acad. Sci. 4(2):61-70. 

COPE, E.D. 1875. Check-list of North American 
Batrachia and Reptilia; with a systematic list of 
the higher groups, and an essay on geographical 
distribution. Based on the specimens contained in 
the U.S. National Museum. Bull. U.S. Natl. Mus. 
1:1-104. 

. 1880. On the zoological position of Tex- 
as. Bull. U.S. Natl. Mus. 17:1-51. 

. 1885. A contribution to the herpetology 



of Mexico. Proc. Amer. Phil. Soc. 22:379-404. 
. 1887. Catalogue of batrachians and rep- 



tiles of Central America and Mexico. Bull. U.S. 
Natl. Mus. 32:1-98. 
. 1896. The geographical distribution of 



Batrachia and Reptilia in North America. Amer. 
Natur. 30:886-902, 1003-1026. 
. 1900. The crocodilians, lizards, and 



snakes of North America. Rept. U.S. Natl. Mus. 
1898:153-1270. 

CORRUCCINI, R.S. 1977. Correlation properties 
of morphometric ratios. Syst. Zool. 26:211-214. 

COUES, E. 1875. Synopsis of the reptiles and bat- 
rachians of Arizona, with critical and field notes 
and an extensive synonym. Rept. Geogr. Geol. 
Expl. Surv. W. of 100th Merid., Wheeler 5:585- 
633. 



CROULET. C.H. 1963. A taste of the tropics. J. 

Philadelphia Herpetol. Soc, 11:1-5. 
DeBUEN, F. 1943. Los Lagos Michoacanos. I. 

Caracteres generales. El Lago de Zirahuen. Rev. 

Soc. Mex. Hist. Nat. 4:211-232. 
. 1944. Limnobiologi'a de Pa'tzcuaro. An. 

Inst. Biol. Mex. 15:261-312. 
. 1945. Resultados de una campana limno- 



Idgica en Chapala y observaciones sobre otras 
aguas explorados. Rev. Soc. Mex. Hist. Nat. 
6:129-144. 
DEEVEY, E.S. 1957. Limnologic studies in Middle 
America with a chapter on Aztec limnology. 
Trans. Connecticut Acad. Arts Sci. 39:213-328. 
DEGENHARDT, W.G. and J.L. CHRISTIANSEN. 
1974. Distribution and habitats of turtles in New 
Mexico. Southwestern Nat. 19:21-46. 
DE TERRA, H., J. ROMERO and T.D. STEW- 
ART. 1949. Tepexpan man. Viking Fund Publ. 
Anthro. No. 11. 
DITMARS, R.L. 1907. The reptile book. Double- 
day, Page and Co., New York. 

. 1936. The reptiles of North America. A 

review of the crocodiles, lizards, snakes, turtles 
and tortoises inhabiting the United States and 
northern Mexico. Doubleday Doran, New York. 
DIXON, J.R., C.A., KETCHERSID and C.S. 
LIEB. 1972. The herpetofauna of Queretaro, 
Mexico, with remarks on taxonomic problems. 
Southwestern Nat. 16:225-237. 
DIXON, W.J. 1973. BMD: Biomedical computer 
programs. Univ. California Press, Berkeley. 

. 1977. BMDP-77; Biomedical computer 

Programs. Univ. California Press, Berkeley. 
DODSON, P. 1978. On the use of ratios in growth 

studies. Syst. Zool. 27:62-67. 
DUELLMAN, W.E. 1955. Notes on reptiles and 
amphibians from Arizona. Occ. Pap. Mus. Zool. 
Univ. Michigan 569:1-14. 

. 1961. The amphibians and reptiles of 

Michoacan, Mexico. Univ. Kansas Publ. Mus. 
Nat. Hist. 15:1-148. 
. 1965. A biogeographic account of the 



herpetofauna of Michoacan, Mexico. Univ. Kan- 
sas Publ. Mus. Nat. Hist. 15:627-709. 
, T. FRITTS, and A.E. LEVITON. 1978. 



Museum acronyms. Herpetol. Review 9:5-9. 
DUCES, A.A.D. 1869. Catalogo de animales verte- 

brados observados en la republica Mexicana. 

Naturaleza 1:137-145, 414. 
. 1888. Erpetologi'a del Valle de Me'xico. La 

Naturaleza. 2(1):97-146. 
. 1895. Fauna del estado de Guanajuato. 



In Memoria sobre la administracidn publica del 
estado de Guanajuato presentada al Congreso del 



68 



Tulane Studies in Zoology and Botany 



Vol. 23 



mismo por el c. Governador Constitucional Lie. 
Joaquin Obregon Gonzales, el 1 de Abril de 1895, 
Morelia. 
. 1896a. Sobre la distributio'n geografica 



de los animales. Acl. Soc. Scient. Chili, 6:liv-lvi. 
. 1896b. Comparaci6n entre el esqueleto de 



la ave y el de la tortuga. Mems. Revta. Soc. Cient. 
'Antonio Alzate' 9:329-331. 
. 1896c. Reptiles y batracios de los Estados 



Unidos Mexicanos. Naturaleza, 2(2);479-485. 
. 1898. El Caracter en los animales. Natu- 



raleza 2(3):39-42. 

DUMERIL, A.M.C. and G. BIBRON. 1834. Erpet- 
ologie generale ou histoire naturelle complete des 
reptiles. Vol. 1. Librairie Encyclopedique de 
Roret, Paris. 

and A.H.A. DUMERIL. 1851. Catalogue 

methodique de la collection des reptiles du 
Museum d'Histoire Naturelle. Gide & Boudry, 
Paris. 

DUMERIL, A.H.A. 1870. Etu'cies sur les reptiles. 
Mission scientifique au Mexique et dans I'Amefique 
Centrale . . . recherches zoologiques. 3d part. 
Imprimerie Imp^riale, Paris. 

DUNN, E.R. 1936. The amphibians and reptiles of 
the Mexican expedition of 1934. Proc. Acad. Nat. 
Sci. Philadelphia 88:471-477. 

ERNST. C.H. and R.W. BARBOUR. 1972. Turtles 
of the United States. Univ. Press Kentucky, Lex- 
ington. 

, R.W. BARBOUR, E.M. ERNST, and 

J.R. BUTLER. 1974. SubspeciTic variation and 
intergradation in Florida Kinoslernon subrubrum. 
Herpetologica 3:317-320. 

FARRIS, J.S. 1966. Estimation of conservatism of 
characters by constancy within biological popula- 
tions. Evolution 20:587-591. 

FITZINGER, L.J. F.J. 1835. Entwurf einer syste- 
matischen Anordnung der Schildkroten nach den 
Grundsatzen der naturlichen Methode. Annln. 
Naturh. Mus. Wien 1:103-128. 

FOREMAN, F. 1955. Palynology in southern North 
America, part II: study of two cores from lake 
sediments of the Mexico City basin. Bull. Geol. 
Soc. Amer. 66:475-5 lOcion . 

Museo Nacional, Mexico, D.F. 

FUNK, R.S. 1974. Geographic distribution: Kino- 
slernon sononense. Herpetol. Rev. 5:20. 

GADOW, H.F. 1905. The distribution of Mexican 
amphibians and reptiles. Proc. Zool. Soc. London 
1905:191-245. 

. 1908. Through southern Mexico, being an 

account of the travels of a naturalist. London, 
Witherby & Co. 

. 1930. Jorullo. The history of the volcano 



GARCIA CUBAS, A. 1884. Cuadro geografico, 

estadi'stico, descriptivo e histdrico de los Estados 

Unidos Mexicanos. Seer. Fomento. Mexico, D.F. 

GARMAN, S. 1884. The North American reptiles 

and batrachians. Bull. Essex Inst. 16:1-46. 
. 1887. Reptiles and batrachians from Tex- 
as and Mexico. Bull. Essex Inst. 19:119-138 (In 
separate, pp. 1-20). 

GATES, G.O. 1957. A study of the herpetofauna 
in the vicinity of Wickenburg, Maricopa Co., Ari- 
zona. Trans. Kansas Acad. Sci. 60:403-418. 
GIJZEN, A. and H. WERMUTH. 1958. Schildkroten- 
Pflege in offentilichen Schau-Aquarien nach bio- 
logischen Gesichtspunkten. Bull Soc. R. Zool. 
Anvers 6:1-65. 
GLASS, B. and N. HARTWEG. 1951. Kinoslernon 
murrayi, a new musk turtle of the hirlipes group 
from Texas. Copeia 1951:50-52. 
GMELIN, J.-F. 1788. CaroH a Linne, Systema 
naturae per regna tria natural, secondum classes, 
ordines, genera, species, cum characteribus differ- 
entiis, synonymis, locis. 1(3): 1038-1516. 
GOLDMAN, E.A. 1951. Biological investigations in 

Mexico, Smithsonian Misc. Coll. 115:1-476. 
GOLOMB, B. 1965. Paleogeography of the Basin of 
Mexico. Ph.D. dissertation. Univ. of California, 
Los Angeles. 
GOULD, S.J. and R.F. JOHNSTON. 1972. Geo- 
graphic variation. Ann. Rev. Ecol. Syst. 3:457- 
498. 
GRAY, J.E. 1844. Catalogue of the tortoises, croco- 
diles, and amphisbaenians, in the collection of the 
British Museum. Edward Newman, London. 

. 1855. Catalogue of the shield reptiles in 

the collection of the British Museum. Part I. 
Testudinata (tortoises). Taylor and Francis, 
London. 
. 1869. Notes on the families and genera of 



tortoises (Testudinata) and on the characters 
afforded by the study of their skulls. Proc. Zool. 
Soc. London 1869:165-225. 
. 1870. Supplement to the catalogue of 



shield reptiles in the collection of the British 
Museum. Part I. Testudinata (Tortoises). Taylor 
and Francis, London. 
. 1873. Notes on the tortoises of the zool- 



of Jorullo and the reclamation of the devastated 
district by animals and plants. Cambridge. 



ogy of Mexico of Mm. A Dumeril and Bocourt. 
Ann. Mag. Nat. Hist. (4)12:109-114. 

GREENE, H.W. 1972. Mexican reptiles in the 
Senckenberg Museum. Pittsburgh. Pennsylvania, 
Carnegie Mus.: 1-15. 

GRINNELL, J. and C.L. CAMP. 1917. A distribu- 
tional list of the amphibians and reptiles of Cali- 
fornia. Univ. California Publ. Zool. 17:127-208. 

GUNTHER, A.C.L.G. 1885-1902. Biologia Centrali- 
Americana. Reptilia and Batrachia. Porter, Lon- 



No. 1 



Kinosternon Biosystematics 



69 



don. 
HAMBRICK, P.S. 1976. Additions to the Texas 

herpetofauna, with notes on peripheral range 

extensions and new records of Texas amphibians 

and reptiles. Texas J. Sci. 27:291-299. 
HARDY, L.M., and R.W. MCDIARMID. 1969. 

The amphibians and reptiles of Sinaloa, Mexico. 

Univ. of Kansas Publ. Mus. Nat. Hist. 18:39-252. 
HENRICKSON, J. 1978. Saline habitats and halo- 

phytic vegetation of the Chihuahuan Desert 

region. USDI. Natl. Park Serv. Trans. Proc. Serv. 

3:289-314. 
HERINGHI, H.L. 1969. An ecological survey of 

the herpetofauna of Alamos, Sonora, Mexico. 

M.S. thesis. Arizona State University, Tempe. 
HERRERA, A.L. 1890. Notas acerca de los verte- 

brados del Valle de Mexico. Naturaleza 2{1):299- 

342. 
. 1891. El clima de Valle de Mexico y la 

biologia de los vertebrados. (Part 1.) Naturaleza 

2(2)1-2:38-86. 
. 1893. El clima de Valle de Mexico y la 



biologia de los vertebrados.(Part 2.) Naturaleza 
2(2):324-358. 

1899. Sinonimia vulgar y cientifica de los 



principals vertebrados me'xicanos;.. Mexico. Seer. 
Fomento.: 1-31. 

.. 1904. Catalogo de la coleccion de reptiles 



y batracios del Museo Nacional. Segunda edicidn. 
Museo Nacional, Mexico, D.F. 
and D. V. LOPE. 1899. La vie sur les hauts 



plateaux. Influence de la pression barometrique 
sur la pression barometrique sur la constitution et 
le development des etres organises. Traitment 
climaterique de la tuberculose. I. Escalante, Mex- 
ico. 

HEYER, W.R. 1978. Systematics of the Fuscus 
group of the frog genus Leptodactylus (Amphib- 
ia, Leptodactylidae). Sci. Bull. Los Angeles Co. 
Nat. Hist. Mus. 29:1-85. 

HIBBARD, C.W. 1955. Pleistocene vertebrates 
from the Upper Becerra (Becerra Superior) form- 
ation. Valley of Tequixquiac, Mexico, with notes 
on other Pleistocene forms. Contrib. Mus. Pale- 
ontology, Univ. Mich. 12:47-96. 

HUBBS, C.L. and R.R. MILLER. 1948. Correla- 
tion between fish distribution and hydrographic 
history in the desert basins of western United 
States. In The Great Basin, with emphasis on 
glacial and postglacial times. Bull. Univ. Utah 
38(20): 17-166. 

and V.G. SPRINGER. 1957. A revision 

of the Gambusia nobilis species group, with de- 
scriptions of three new species, and notes on their 
variation, ecology, and evolution. Texas J. Sci. 
9:279-327. 

HULSE, A.C. 1974. An autecological study of 



Kinosternon sonoriense LeConte (Chelonia: Kino- 
sternidae). Ph.D. dissertation. Arizona State 
Univ., Tempe. 
. 1976. Growth and morphometries of 



Kinosternon sonoriense (Reptilia, Testudines, 
Kinosternidae). J. Herpet. 10:341-348. 

HUNTINGTON, E. 1914. The climatic factor as il- 
lustrated in arid America. Carnegie Institute 
Washington 192:1-262. 

INGER, R.F. 1961. Problems in the application of 
the subspecies concept in vertebrate taxonomy, 
pp. 262-285. In W.F. Blair (ed), Vertebrate speci- 
ation. Univ. Texas Press, Austin. 

IVERSON, J.B. 1976. Kinosternon sonoriense. Cat. 
Amer. Amph. Rept. 176:1-2. 

1977a. Geographic variation in the musk 

turtle Sternotherus minor. Copeia 1977:502-517. 

. 1977b. Kinosternon subrubrum. Cat. 



Amer. Amph. Rept. 193:1-4. 
1978. Distributional problems of the 



genus Kinosternon in the American southwest. 
Copeia 1978:476-479. 
. 1979. A taxonomic reappraisal of the 



yellow mud turtle, Kinosternon flavescens (Testu- 
dines: Kinosternidae). Copeia, 1979:212-225. 
. (in press) Kinosternon hirtipes. Cat. 



Amer. Amph. Rept. :l-4. 
. MS 1 . Scaling of skeletal tissues in turtles: 



Responses to predation. Evolution. 
MS 2. Geographic variation in sexual di- 



morphism in the mud turtles, Kinosternon hir- 
tipes. 
and J.F. BERRY. 1979. The genus Kino- 



sternon in northeastern Mexico (Testudines:Kino- 
sternidae). Herpetologica 35:318-324. 
and F.O. WEYMAN. MS . Sexual di- 



morphism in kinosternid turtle skeletal compon- 
ents. 

IVES, R.L. 1936. Desert floods in the Sonoyta 
Valley. Amer. J. Sci. 32:349-360. 

JOHNSON, C.A. 1969. A redescription of Myxidi- 
um chelonarum Johnson, 1969 (Cnidospora:Myx- 
idiidae) from various North American turtles. J. 
Protozool. 16:700-702. 

KAUFFELD, C.F. 1943. Field notes on some Ari- 
zona reptiles and amphibians. Amer. Midi. Nat. 
29:342-359. 

KLUGE, A.G. and J.S. FARRIS. 1969. Quantita- 
tive phyletics and the evolution of anurans. Syst. 
Zool. 18:1-32. 

KRANZ, F.M., H.M. SMITH and R.B. SMITH. 
1971. Interpretive essay on the eleventh book of 
the history of Sahagun. Bull. Philadelphia Her- 
pet. Soc. 18:11-24. 

LACEPEDE, B. 1788. Histoire naturelle des quad- 
rupedes ovipares et des serpens Vol. 1. Hotel de 



70 



Tulane Studies in Zoology and Botany 



Vol. 23 



Thou, Paris. 
LAMPE, E. 1901. Catalog der Reptilien-Sammlung 
(Schildkrolen, Crocodile, Eidechsen und Cha- 
maeleons) des Nalurhistorischen Museums zu 
Wiesbaden. Jahrb. Nassau. Ver. Naturk. 
54:177-222. 
LaRlVERS, 1. 1942. Some new amphibians and rep- 
tiles records from Nevada. J. Entomol. Zool. 
34:52-68. 
LeCONTE, J. 1854. Description of four new species 
of Kinoslernum. Proc. Acad. Nat. Sci. Philadel- 
phia 7:180-190. 

. 1859. Description of two new species of 

tortoises. Proc. Acad. Nat. Sci. Philadelphia 
11:4-7. 
LEGLER, J.M. 1960. Remarks on the natural his- 
tory of the Big Bend slider, Pseudeinys scripta 
gaigeae Hartweg. Herpetologica 16:139. 

and R.G. WEBB. 1970. A new slider turtle 

(Pseudeinys scripta) from Sonora, Mexico. Herpe- 
tologica 26:157-168. 

LICHTENSTEIN, H. 1856. Nomenclator reptilium 
el amphibiorum musei zoologici berolinensis. 
Berlin. 
LINER, E.A. 1954. The herpetofauna of Lafayette, 
Terrebonne, and Vermilion Parishes, Louisiana. 
Proc. Louisiana Acad. Sci. 17:65-85. 

. 1964. Notes on four small herpetological 

collections from Mexico. 1. Introduction, turtle 
and snakes. Southwestern. Nat. 8:221-227. 
LITTLE, E.L. 1940. Amphibians and reptiles of the 
Roosevelt Reservoir Area, Arizona. Copeia 
1940:260-265. 
LOPEZ, E. 1975. Effects of low-sodium salines 
upon the atrio-ventricular propagation of the 
turtle heart. Jap. J. Physiol. 25:1-15. 
LORENZO, J.L. 1958. Una hipotesis paleoclimatica 
para la cuenca de Mexico. Misc. Paul. Rivet, 
Octogenario Dicata:579-584. 
MCARTHUR, R. and E.O. WILSON. 1967. The 
theory of island biogeography. Monogr. Pop. 
Biol. 1:1-203. Princeton Univ. Press, Princeton. 
MALDONADO-KOERDELL, M. 1955. La historia 
geohidrologiza de la cuenca de Mexico. Revta, 
Mex. Estud. Antrop. 14:15-21. 
MALKIN, B. 1958. Cora ethnozoology, herpeto- 
logical knowlege; a bioecologica! and cross 
cultural approach. Anthrop. Q. 31:73-90. 
MALNATE, E.V. 1971. A catalogue of primary 
types in the herpetological collections of the 
Academy of Natural Science of Philadelphia. 
Proc. Acad. Nat. Sci. Philadelphia 123(9):345- 
375. 
MARSHALL, J.T., JR. 1957. Birds of the pine-oak 
woodland in southern Arizona and adjacent Mex- 
ico. Pacific Coast Avifauna 32:1-125. 



MARTIN DEL CAMPO, R. 1937. Contribucion al 
conocimienio de los balracios y reptiles del Valle 
des Mesquital, Hgo. An. Inst. Biol. Univ. Mex. 
8:259-266. 

. 1938. Ensayo de interprelacion del libro 

undecimo de la Historia de Sahagun. An. Inst. 
Biol. Univ. Mex. 9:379-391. 

. 1940. Vertebrados de Patzcuaro. An. 



Inst. Biol. Univ. Mex. 1 1(2):481-492. 

.. 1955. Productos biologicas del Valle de 



Mexico. Revta. Mex. Estud. Antrop. 14:53-77. 

MAYR, E. 1970. Populations, species, and evolu- 
tion. Belknap Press, Cambridge, Massachusetts. 

MCNATT, R.M. 1978. Fishes and habitats of the 
San Pedro River basin, southeastern Arizona. 
Program 58th meeting Amer. Soc. Ichthyol. and 
Herpetol. 

MEARNS, E.A. 1907. Mammals of the Mexican 
boundary of the United States. A descriptive 
catalog of the species of mammals occurring in 
that region; with a general summary of the natural 
history, and a list of trees. Bull. U.S. Nat'l. Mus. 
56:1-530. 

MEEK, S.E. 1904. The freshwater fishes of Mexico 
north of the Isthmus of Tehuantepec. Field Col. 
Mus. Pub. 93 (Zool.) 5:1-252. 

MERTENS, R. and H. WERMUTH. 1955. Die 
rezenten SchildkrOten, Krokodile und Brucken- 
echsen. Eine kritische Liste der heute lebenden 
Arten und Rassen. Zool. Jb., Abt. Allg. Zool. 
83:323-440. 

MILLER, R.R. 1958. Origin and affinities of the 
freshwater fish fauna of western North America, 
p. 187-222. In C.L. Hubbs (ed.), Zoogeography, 
Publ. Amer. Assoc. Adv. Sci. #51. 

. 1961. Man and the changing fish fauna of 

the American southwest. Pap. Michigan Acad. 
Sci. Arts, Lett. 46:365-404. 

_. 1968. A drainage map of Mexico. Syst. 



Zool. 17:174-175. Minckley, W.L. and R.K. Koehn. 
1965. Re-discovery of the fish fauna of the Sauz 
Basin, northern Chihuahua, Mexico. Southwest- 
ern. Nat. 10:313-315. 

MITTERMEIER, R.A. 1971. Status — the market 
in So.E. Mexico. Int. Turtle Tortoise Soc. J. 
5(3):15-19. 

MOLL, E.O. and J.M. LEGLER. 1971 . The life his- 
tory of a neotropical slider turtle, Pseudemys 
scripta (SchoepfO, in Panama. Sci. Bull. Los 
Angeles Co. Mus. Nat. Hist. 11:1-102. 

and K.L. WILLIAMS. 1963. The musk 

turtle Sternothaerus odoratus from Mexico. 
Copeia 1963:157. 

MONTEVECCHI, W.A. and J. BURGER. 1975. 
Aspects of the reproductive biology of the north- 
ern diamondback terrapin, Malaclemys terrapin 



No. 1 



Kinosternon Biosystematics 



71 



terrapin. Amer. Midi. Nat. 94:166-178. 

MOOSER, F. 1957. Los ciclos de vulcanismo que 
formaron la cuenca de Mexico. XX Cong. Geol. 
Internac. 1:337-348. 

. 1963. La cuenca lacustre del Valle de 

Me'xico, p. 1-48. In Mesas Redondas sobre Prob- 
lemas del Valle de Mexico, 12-16 Nov. 1962. Inst. 
Mex. Recursos Nat. Renov., A.C., Me'xico. 

MOOSER, F., S.E. WHITE, and J.L. LORENZO. 
1956. La cuenca de Me'xico: consideraciones geo- 
logicas y arqueologicas. Inst. Nac. Antropol. 
Hist., Publ. No. 2. 

MORAFKA, D.J. 1977. A biogeographical analysis 
of the Chihuahuan Desert through its herpeto- 
fauna. Biogeographica 9:1-320. 

MULLER, F. 1885. Vierter Nachtrag zum Katalog 
der herpetologischen Sammlung des Easier Muse- 
ums. Verh. Naturf. Ges. Basel 7:668-717. 

MOLLER, J.W. 1865. Reisen in den Vereinigten 
Staaten, Canada und Mexiko. III. Beitrage zur 
Geschichte, Statistik und Zoologie von Mexiko. 
Dritte Abtheilung. Die Wirbelthiere Mexikos. III. 
Amphibia. Leipzig, Brockhaus. 

NICKERSON, M.A. and C.E. MAYS. 1970. A pre- 
liminary herpetofaunal analysis of the Graham 
(Pinaleno) Mountain Region, Graham Co., Ari- 
zona, with ecological comments. Trans. Kansas 
Acad. Sci. 72:492-505. 

NIEDERBERGER, C. 1979. Early sedentary econ- 
omy in the Basin of Mexico. Science 203:131-142. 

NILES, D.M. 1962. Records for the Sonora mud 
turtle, Kinosternon sonoriense, in New Mexico. 
Herpetologica 18:205-206. 

NUSSBAUM, R.A. 1976. Geographic variation and 
systematics of salamanders of the genus Dicamp- 
todon Strauch (Ambystomatidae). Misc. Publ. 
Mus. Zool. Univ. Michigan 149:1-94. 

OHMART, R.D., W.O. DEASON and S.J. FREE- 
LAND. 1975. Dynamics of marsh land formation 
and succession along the Lower Colorado River 
and their importance and management problems 
as related to wildhfe in the Arid Southwest. Trans. 
40th North Amer. Wildlife and Nat. Res. Conf.: 
240-251. 

PARSONS, T.S. 1968. Variation in the choanal 
structure of Recent turtles. Canadian J. Zool. 46: 
1235-1263. 

PEREZ VILLEGAS, G. and T. REYNA TRUJIL- 
LO. 1978. Regidnes faunisticas y el medio geo- 
grafrico del Valle de Mexico. Congreso Nac. 
Zool, 1:211-218. 

PETERS, J. A. 1952. Catalogue of type specimens 

in the herpetological collections of the University 

of Michigan Museum of Zoology. Occ. Pap. Mus. 

Zool. Univ. Michigan 529:1-55. 

PICKWELL, G.B. 1947. Amphibians and reptiles 



of the Pacific states. Stanford Univ. Press, Palo 
Alto. 
PRATT, H.S. 1923. A manual of land and fresh- 
water vertebrate animals of the United States. P. 
Blakiston's Sons, Philadelphia. 
PRITCHARD, P.C.H. 1967. Living turtles of the 
world. T.F.H. Publ. Co., Jersey City, NJ. 

. 1979. Encyclopedia of turtles. T.F.H. 

Publ. Co., Neptune, New Jersey. 
ROHLF, F.J. and J. KISPAUGH. 1972. Numerical 
taxonomy system of multivariate statistical pro- 
grams. State Univ. Stony Brook, New York. 
ROMERO, H. 1965. Catalogo sistematica de los 
peces del Alto Lerma con descripcion de una 
nueva especie. An. Esc. Nac. Cien. Biol., Mex. 
14:47-80. 
RUST, H.-T. 1934. Systematische Liste der leben- 
den Schildkroten. Blatt. f. Aquar.-Terrk. 45:59- 
67. 

. 1938. Interessante schildkroten. V. Die 

Schildkrotengatung Kinosternon Spix. Wschr. 
Aquar. Terrk. 35:22-24. 
RUTHVEN, A.G. 1907. A collection of reptiles and 
amphibians from southern New Mexico and Ari- 
zona. Bull. Amer. Mus. Nat. Hist. 23:483-597. 
SCHMIDT, K.P. 1922. The amphibians and reptiles 
of the Lower California and the neighboring is- 
lands. Bull. Amer. Mus. Nat. Hist. 46:607-707. 

. 1953. A check list of North American 

amphibians and reptiles. Sixth edition. Amer. 
Soc. Ichthyol. Herpetol, Univ. Chicago Press, 
Chicago. 
SEARS, P.B. and K.H. CLISBY. 1955. Palynology 
in southern North America part IV: Pleistocene 
climate in Mexico. Bull. Geol. Soc. Amer. 66:521- 
530. 
SEMMLER, R., M.E. SEIDEL and S. WILLIAMS. 
1977. Kinosternon from Mexico. New Mexico 
Herpetol. Soc. Newsl. 14:18. 
SERVICE, J. 1972. A user's guide to the statistical 
analysis system. North Carolina State University 
Supply, Raleigh. 
SIEBENROCK, F. 1905. Chelonologische Notizen. 
Zool. Anz. 28:460-468. 

1906. Schildkroten aus Sudmexiko. Zool. 

Anz. 30:94-102. 

. 1907. Die Schildkrotenfamilie Cinoster- 

nidae. Akad. Wiss. Wien. 116:527-599. 

. 1909. Synopsis der rezenten Schildkroten. 

Zool. Jahrb., Suppl. 10:427-618. 
SMITH, H. M. and H. K. BUECHNER. 1947. The 
influence of the Balcones escarpment on the distri- 
bution of amphibians and reptiles in Texas. Bull. 
Chicago Acad. Sci. 8:1-16. 
SMITH, H. M. and R. B. SMITH 1975. The herpe- 
tological names of Herrera (1899) and their 



72 



Tulane Studies in Zoology and Botany 



Vol.23 



status. Trans. Kansas Acad. Sci. 78:85-87. 

_. 1980. Synopsis of the herpetofauna of 



tersbourg (2)8(1 3): 1-207. 
. 1890. Bemerkungen uber die Schildkrd- 



Mexico. Vol. 6. Guide to the Mexican turtles. 
John Johnson, North Bennington, Vermont. 
. and E. H. TAYLOR. 1950a. An anno- 



tated checklist and key to the reptiles of Mexico 
exclusive of the snakes. Bull. U.S. Natl. Mus. 
199:1-253. 

.. 1950b. Type localities of Mexican reptiles 



and amphibians. Univ. Kansas Sci. Bull. 33(8):313- 
380. 
K. L. WILLIAMS and E. O. MOLL. 



1963. Herpetological explorations on the Rio 
Conchos, Chihuahua, Mexico. Herpetologica 19: 
205-215. 
SMITH, P. W. and M. M. HENSLEY. 1957. The 
mud turtle, Kinoslernon flavescens slejnegeri 
Hartweg, in the United States. Proc. Biol. Soc. 
Washington 70:201-204. 
SNEATH, P. H. A. and R. R. SOKAL. 1973. Nu- 
merical taxonomy. W. H. Freeman and Co., San 
Francisco. 
SOKOLOFF, V. P. and J. L. LORENZO. 1953. 
Modern and ancient soils at some archaeological 
sites in the Valley of Mexico. Amer. Antiquity 19: 
50-55. 
SOLORZANO PRECIADO, A. 1961. Contribucion 
al conocimiento de la biologfa del charal prieto del 
lago de Pa'tzcuaro, Michoac/n. Secretaria de In- 
dustria y Comercio, Direccio'n General de Pesca e 
industrias conexas, Mexico City. 71 pp. 
SPIETH, H. T. 1950. The David Rockefeller Mu- 
seum Expedition of the American Museum of Na- 
tural History. Amer. Mus. Novitates 1454:1-67. 
STEBBINS, R. C. 1966. A field guide to western 
reptiles and amphibians. Houghton Mifflin Co., 
Boston. 
STEJNEGER, L. H. 1899. Reptiles of the Tres Mar- 
ias and Isabel islands. N. Amer. Fauna 14:63-71. 

. 1902. The reptiles of the Huachuca 

Mountains of Arizona. Proc. U.S. Natl. Mus. 25: 
149-158. 
. and T. BARBOUR. 1917. A check list of 



North American amphibians and reptiles. Har- 
vard Univ. Press, Cambridge, Mass. 

STORER, T. I. 1930. Notes on the range and life 
history of the Pacific freshwater turtle Cleininys 
marmorata. Univ. California Publ. Zool. 32:429- 
441. 

STRAUCH, A. 1862. Chelonische Studien mit be- 
sonderer beziehung auf die Schildkrotensamm- 
lung der Kaiserlichen Akademie der Wissenschaf- 
ten St. Petersburg. Mem. Acad. Imp. Sci. St. 
Petersbourg (7)5(7): 1-196. 

. 1865. Die Vertheilung der SchildkrSten 

uber den Erdball. Mem. Acad. Imp. Sci. St. Pe- 



tensammlung in zoologischen Museum der kaiser- 
lichen Akademie der Wissenschaften zu St. Pe- 
tersburg. Mem. Acad. Imp. Sci. St. Petersbourg 
(7)38(2):1-I27. 

STRECKER, J. K. 1915. Reptiles and amphibians 
of Texas. Baylor Univ. Bull. 18(4):l-82. 

and W. J. WILLIAMS. 1927. Herpetolog- 
ical records from the vicinity of San Marcos, 
Texas, with distributional data on the amphibians 
and reptiles of Edward's plateau and central Tex- 
as. Contrib. Baylor Univ. Mus. 12:1-16. 

TAMAYO, J. L. 1962. Geografi'a General de Mex- 
ico, Vols. 1-4. Inst. Mex. Invest. Econ., Mexico. 

. 1964. The hydrography of Middle Amer- 
ica, p. 84-121. In R. WAUCHOPE and R. C. 
WEST (eds.). Handbook of Middle American 
Indians, Univ. Texas Press, Austin. 

TANNER, W. W. and W. G. ROBISON, JR. 1960. 
Herpetological notes for northwestern Jalisco, 
Mexico. Herpetologica 16:59-62. 

TAYLOR, E. H. 1936. Notes on the herpetological 
fauna of the Mexican state of Sinaloa. Univ. Kan- 
sas Sci. Bull. 24:505-537. 

. 1939. Some Mexican serpents. Univ. Kan- 
sas Sci. Bull. 26:445-487. 

_. 1952. Third contribution to the herpeto- 



logy of the Mexican state of San Luis Potosi. 

Univ. Kansas Sci. Bull. 34:793-815. 
TROSCHEL, F. H. 1855. Bericht uber die Leistun- 

gen in der Herpetologie wahrend des Jahres 1854. 

Arch. Naturgesch 21:411-425. 
. 1860. Bericht uber die Leistungen in der 

Herpetologie wahrend des Jahres 1859. Arch. 

Naturgesch 26:265-278. 
UDDEN, J. A., C. L. BAKER and E. BOSE. 1919. 

Review of the geology of Texas. Third edition. 

Univ. Texas. Austin. 
VAN DENBURGH, J. 1922. The reptiles of western 

North America. Vol. II. Snakes and turtles. Occ. 

Pap. CaHfornia Acad. Sci. 10:623-1028. 
. 1924. Notes on the herpetology of New 

Mexico with a list of species known from that 

state. Proc. California Acad. Sci. (4)13:189-230. 
and J. R. SLEVIN. 1913. List of the am- 



phibians and reptiles of Arizona, with notes on 
the species in the collection of the Academy. Proc. 
California Acad. Sci. (4)3:391-454. 

VAN DEVENDER, T. R. and C. H. LOWE, JR. 
1977. Amphibians and reptiles of Yepomera, Chi- 
huahua, Mexico. J. Herpetol. 11:41-50. 

and W. VAN DEVENDER. 1975. Ecolog- 
ical notes on two Mexican skinks (genus 
Eumeces). Southwestern Nat. 20:271-287. 

VELASCO, A. L. 1890a. Geografia y estadi'stica del 



No. 1 



Kinosternon Biosystematics 



73 



estado de Nuevo Leon. Geografia y Estadi'stica de 
la Republica Mexicana. Vol. 4. Secr. Fomento, 
Mexico, D.F. 231 pp. 

1890b. Geograffa y estadi'stica del estado 



de Guanajuato. Geografia y Estadistica de la 
Republica Mexicana. Vol. 5. Secr. Fomento, 
Mexico, D.F. 300 pp. 

1891. Geograffa y estadi'stica del estado 



de Queretaro-Arteaga. Geografi'a y estadistica de 
la Republica Mexicana. Vol. 8. Secr. Fomento, 
Mexico, D.F. 140 pp. 

1892a. Geografia y estadi'stica del estado 



de Guerrero. Geografi'a y estadistica de la Repub- 
lica Mexicana. Vol. 10. Secr. Fomento, Mexico, 
D.F. 248 pp. 

.. 1892b. Geografia y estadistica del estado 



de Tlaxcala. Geografi'a y estadistica de la Repub- 
lica Mexicana. Vol. 11. Secr. Fomento, Mexico, 
D.F., 138 pp. 
. 1892c. Geografia y estadistica del estado 



de Tamaulipas. Geografi'a y estadistica de la Re- 
publica Mexicana. Vol. 12. Secr. Fomento, Mex- 
ico, D.F., 204 pp. 

1893a. Geografi'a y estadistica del estado 



de Durango. Geografi'a y estadistica de la Repub- 
lica Mexicana. Vol. 13. Secr. Fomento, Mdxico, 
D.F., 196 pp. 
. 1893b. Geografi'a y estadi'stica del estado 



de Sonora. Geografia y estadistica de la Republica 
Mexicana. Vol. 14. Secr. Fomento, Mexico, D.F., 
248 pp. 
1894. Geografia y estadistica del estado 



de Zacatecas. Geografi'a y estadi'stica de la Repub- 
lica Mexicana. Vol. 15. Secr. Fomento, Mexico, 
D.F., 324 pp. 
. 1895. Geografia y estadistica del estado 



de Campeche. Geografia y estadistica de la Re- 
publica Mexicana. Vol. 16. Secr. Fomento, 
Mexico, D.F., 140 pp. 
. 1896a. Geografia y estadistica del estado 



de Aguascalientes. Geografia y estadistica de la 
Republica Mexicana. Vol. 17. Secr. Fomento, 
Mexico, D.F., 136 pp. 
. 1896b. Geografia y estadi'stica del estado 



de Colima. Geograffa y estadistica de la Republica 
Mexicana. Vol. 18. Secr. Fomento, Mexico, D.F., 
142 pp. 

1897. Geograffa y estadi'stica del Estado 



de Coahuila de Zaragoza. Geografia y estadistica 
de la Republica Mexicana. Vol. 19. Secr. 
Fomento, Mexico, D.F., 202 pp. 
. 1898. Geografia y estadistica del estado 



de Chiapas. Geografia y estadistica de la Repub- 
lica Mexicana. Vol. 20. Secr. Fomento, Mexico, 
D.F., 164 pp. 



WAGLER, J. G. 1830. Naturliches System der Am- 
phibien, mit voranghender classification der Sau- 
gethiere und Vogel. Ein Beitrag zur vergleichender 
Zoologie. Munich. 

. 1828-1833. Descripciones et icones am- 

phibiorum. Parts 1-3. Munich, J. G. Cotta. 

WAUER, R. H. and D. H. RISKIND (eds.) 1978. 
Transactions of the Symposium on the biological 
resources of the Chihuahuan desert region, United 
States and Mexico. U. S. D. I. Natl. Park Serv. 
Trans, and Proc. 3:1-658. 

WEBB, R. G. 1973. Trionyx spiniferus. Cat. Amer. 
Amph. Rept. 140:1-4. 

WEISE, J. G. 1962. A new material for marking 
Kordite line and various animals. Turtox News 40: 
165. 

WESTPHAL-CASTELNAU, A. 1872. Catalogue 
de la collection de reptiles de feu M. Alexandre 
Westphal-Castelnau. C. R. Congr. Scient. Fr., 
Montpellier 35:273-327. 

WERMUTH, H. and R. MERTENS. 1961. Schild- 
kroten, Krokodile, Bruckenechsen. G. FISCHER, 
Jena. 

. 1977. Liste der rezenten Amphibien and 

Reptilien. Testudines, Crocodylia, Rhynchoce- 
phalia. Das Tierreich 100:1-174. 

WHITE, S. E. 1962. Late Pleistocene glacial se- 
quence for the west side of Iztacci'huatl, Mexico. 
BuU. Geol. Soc. Amer. 73:935-958. 

WIEWANDT, T. A. 1971. Breeding biology of the 
Mexican leaf-frog. Fauna 2:29-34. 

, C. H. LOWE and M. W. LARSON. 

1972. Occurence of Hypopachus variolosus 
(Cope) in the short-tree forest of southern Sonora, 
Mexico. Herpetologica 28:162-164. 

WILLIAMS, K. L., H. M. SMITH and P. S. 
CHRAPLIWY. 1963. Turtles and lizards from 
northern Mexico. Trans. Illinois Acad. Sci. 53:36- 
45. 

and L. D. WILSON. 1966. Noteworthy 

Mexican reptiles in the Louisiana State University 
Museum of Zoology. Proc. Louisiana Acad. Sci. 
28:127-130. 

YARROW, H. C. 1875. Report upon the collections 
of batrachians and reptiles made in portions of 
Nevada, Utah, California, Colorado, New Mex- 
ico, and Arizona, during the years 1871, 1872, 
1873, and 1874. Rept. Geog. Geol. Expl. Surv. W. 
100 Mer. Wheeler. 5(4):509-584. 

. 1883. Check-list of North American Rep- 

tilia and Batrachia with catalogue of specimens in 
the U.S. National Museum. U.S. Natl. Mus. Bull. 
24:1-249. 

ZANGERL, R. 1939. Homology of the shell ele- 
ments of turtles. J. Morph. 65:383-410. 



74 Tulane Studies in Zoology and Botany Vol. 23 

ZEEVAERT, L. 1953. Outline of the stratigraphical 
and mechanical characteristics of the unconsoli- 
dated sedimentary deposits in the basin of the Val- 
ley of Mexico. Proc. 4th Cong. Int. Assoc. Quart. 
Res. (INQUA):1-14, Rome-Pisa, Italy. 

ZWEIFEL, R. G. 1960. Results of the Puritan- 
American Museum of Natural History expedition 
to western Mexico. 9. Herpetology of the Tres 
Marias Islands. Bull. Amer. Mus. Nat. Hist. 119: 
77-128. 

and K. S. NORRIS. 1955. Contribution to 

the herpetology of Sonora, Mexico: descriptions 
of new subspecies of snakes (Micruroides eury- 
xanthus and Lampropeltis getulus) and mis- 
cellaneous collecting notes. Amer. Midi. Nat. 54: 
203-249. 



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, 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 


o 


o 
o 






o 
o 




*-? 


o 




O 


CA 


c* 


<> 


o 


o 


o 


o 






o 


A 


<) 


(TA 


() 


o 




o 


o 








o 


o 


O 


O 


OA 


(> 


•o 




o 




o 


A-? 


A 


G* 


O 


O 


O 


o 


•o 


» 


o 


o 


o 




o 


O 


O 




o 


• 


•o 


K) 


o 


o 






o 


O* 


O 


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 


• 
• 

• 












• 




• 


? 


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 





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. 



Literature Cited 

BOSCHUNG, H. and P. O'NEIL. 1980. The effects 
of forest clearcutting on fishes and macroin- 
vertebrates. U.S.D.A. Forest Service, Atlanta, 
Georgia. 32 pp. 

BRAASCH, M. and P. SMITH. 1967. The life 
history of the slough darter, Elheostoma gracile 
(Pisces: Percidae). 111. Nat. Hist. Surv. Biol. 
Notes No. 58 12 pp. 

BRYANT, R. 1979. The life history and compar- 
ative ecology of the sharphead darter, Etheo- 
sioma acuticeps. Tennessee Wildlife Resources 
Agency Technical Report No. 79-50. 60 pp. 

BURR, B. and L. PAGE. 1978. The life history of 
the cypress darter, Elheostoma proeliare. in 
Max Creek. 111. Nat. Hist. Surv. Biol. Notes 
No. 106. 15 pp. 



FAHY, W. 1954. The life history of the northern 
greenside darter, Etheostoma blennioides. Elisha 
Mitchell Scientific Society 70:139-205. 

FLYNN, R. and R. HOYT. 1979. The life history of 
the teardrop darter, Etheostoma barbouri 
Kuehne and Small. Amer. Midi. Nat. 101:127- 
141. 

MATHUR, D. 1973. Some aspects of life history of 
the blackbanded darter, Percina nigrofasciata, 
in Halawakee Creek, Alabama. Amer. Midi. 
Natur. 89:381-393. 

. and J. RAMSEY. 1974. Reproductive bio- 
logy of the rough shiner, Notropis baileyi, 
in Halawakee Creek, Alabama. Trans. Amer. 
Fish. Soc. 103:88-93. 

MURDOCH, W., S. AVERY, and M. SMYTH. 
1975. Switching in predatory fish. Ecology 
56:1095-1105. 

PAGE, L. 1974. The life history of the spottail 
darter, Etheostoma squamiceps, in Big Creek, 
Illinois and Ferguson Creek, Kentucky. 111. Nat. 
Hist. Surv. Biol. Notes No. 89. 20 pp. 

. 1975. The life history of the stripetail darter, 

Etheostoma kennicoiti, in Big Creek, Illinois. 
111. Nat. Hist. Surv. Biol. Notes No. 93. 16 pp. 

. and B. BURR. 1976. The life history of 



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 
production in freshwater. Blackwell Scientific 
Publications, Oxford, England. 348 pp. 

ROBERTS, N. and H. WINN. 1962. Utilization of 
the senses in feeding behavior of the johnny 
darter, Etheostoma nigrum. Copeia 1962:567- 
570. 

SCALET, C.G. 1972. Food habits of the orange- 
belly darter, Etheostoma radiosum cyanorum 
(Osteichthyes: Percidae). Amer. Midi. Natur. 
87:515-524. 

. 1973. Reproduction of the orangebelly 

darter, Etheostoma radiosum cyanorutn. Amer. 
Midi. Natur. 89:156-165. 

SCHENCK, J. and B.C. WHITESIDE. 1977. Food 
habits and feeding behavior of the fountain 
darter, Etheostoma fonticola (Osteich- 
thyes: Percidae). Southwest Nat. 21(4):487-492. 

STARNES, W.C. 1977. The ecology and life history 
of the endangered snail darter, Percina (Imo- 
stoma) tansi Etnier. Tenn. Wildlife Resources 
Agency Tech. Report No. 77-52. 143 pp. 

STILES, R.A. 1975. The reproductive behavior of 
Etheostoma simoterum (Cope)(Perci formes: 
Percidae). Program Abstracts, American Soci- 
ety of Ichthyologists and Herpetologists 55th 
Annual Meeting, June 8-14, Williamsburg, Vir- 
gia. p. 121. (Abstr.) 



No. 1 Etheostoma Life History 83 

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

1978. Metabolism, critical oxygen tension, and 

habitat selection in darters (Etheostoma). 

Ecology 59:99-107. 
WINN, H. 1958a. Comparative reproductive be- 
havior and ecology of fourteen species of 

darters (Pisces: Percidae). Ecol. Mongr. 

28:155-191. 
1958b. Observations on the reproductive 

habits of darters (Pisces: Percidae). Amer. 

Midi. Natur. 59:190-212. 



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 






§i1 

S -a E 



> 



c . 
— Q 









tS; E I 

>< ?i "^ 

cxo. .S 
E - 
o 
o ^ 

E -o 

"£^ 
£li 

O '^ r\ 
to ^ 

E |g ^ 

5 o ? 



S Cu ^^ 



5^S 





;3 


■53 O 




•& 


3 s: 




% 






1 


most 
!ome 
emys 




•^ 

3 


13 C ^ 




1 


doide 
and i 




u 


^»«K 


g 


b '^* c 


X 


O 


^ b S 


^*^ 


M 


Uj 5 "^ 


i 


c 


.E|-S 


1 


i 


^"21 





O 




g o 2> 
C JJ ^ 




c 






lU 






b 'u •- 


E 


_aj 






o 


X3 




^ c« HJ 


p 


3 
O 


CU 


E ££ 



^^ D, 



S o 



18 



>J c« 



i^ -b lr> 



S-Si 



^ (0 



o ^ 

U J.. 

c Q 



><• 


^-^ 




c^- 


E 


CL 


^— 




o 


00 




C 




.^ 
o 


c 


w 


o 


■ 


u 


(fl 


eO 


00 

3 


3 


O 


3 
C/5 


C8 






co tn^ 

w 
XJ ~ 

u O 

E-§ 

3 2 

C lU ^ 

S 5 "■= 
15.2iS 

_ 1> D. 

S E >. 

1 ^t 

2 o S 



o ^^ 



- S 



o c 
X o 

ti -a — 

" w c 

^ — — 

o ° s 






2 = * 

ci -c "5 2 

"5 nJ 00 C 

^ « >^ o 



— .2 



UJ 



D..E 



7^ (Ll — 



^^ C 



k. L. U 



E I 



IJ 

c 




^ 




T3 


s: ^ 


iE 


U 


Sb 


T3 


3 


^ • 


C 


o 


? 00 


w 




•^ c 




_>. 


0:e 


"w 


15 


_ o 




E 


C w 


1) 


— o 



O Q. 
^ => 

1> 13 

00 C 

3 
T3 <-» 
C » 

X c 



E 3 
B3 O 
1/5 T3 



0^ 00 u .-.^ 



Ji n. "" "O 

E — ?; « 

II ll 

«> U O JO 



9 c 



E - c 



— o 



o -c 



_, — (U 

I) u- W > 

" c3 _[- 
E£ 

X 

o 



— ; 1> 



.2 t;; x c " 

■a o 

D Q. 

w O 



u. IJ 



rt 



3 ul 2 -= 

I" >, ii .^ 

^ — C "r 

2 E - -c 



J£ 3 J2 ,::; 



~ •- "^ 



^ i! iJ g w 



1> o 

Ei: 



2-S _ 



(U ^ 



CO -- -- o 

« E •- _ 

i_ t w. T3 

nJ O w. C 

U C O IJ 



O 



No. 1 



Malaclemys- Graptemys Relationship 



89 



5 g S E .E 5 

? -. '^ 1^ « -Q 

o 



5 Q, 



s ^' ^ ej - 



■Q « 



W 



(U 



-u _ T- • E O 

i^.E^^ 



•- ^ ^ ^ 



5 ^ £ ■- E ^ 

°"c.E2 o 

T3 pj — m ^S o 

S . 2 -S « ^ 
O <i, <u .O. o - 

c t °-S^ o I 

o ~ a> -^ g Ci. 

-Si "t« ~ -s; s 

- I .5 5 § ,. ^ 

o^^ g o E-a 

Z Q; E 8 a S rt 

•i S 2 o 

O — "^ TS 

ci c« a 2 



o cs 

— T3 



=*- O "O D 



1> ™ — 

^ E 



c^Q 



<!; J3 c 



o t: 



o :z: 



u, c 3 

~; P O 1^ '" 

> ui X) X) 3 



c 
o 


c 


1) 


£ 


^ 






X) 




-o 




15 

x: 


XI 
IS 


E 


c 


_>v 




o 

XI 


3 


"rt 




_ 


C 


E 


o 


(0 


O 




CO 


K 


o 

c 


C 

o 


x: 

D. 






o 




u 






HJ 


;a 


O 


o 


a 


« 


_>. 


(/^ 


"5 


D. 


■« 


k- 










U 


E 


aj 


1) 


ca 


W 


x: 




3. 






o 


■a 

c 


"o 








c 




u 


u 


> 


O 
o 




;o 



■§:5 
1^ 

2 S3 

^.E 

^ C 

2, «j 

5 2.- 

gQ S 

E ^ S 
o -Si 

E E ^ 
•a c s: 

c — := 

g| i 
2 c g 

u- -^ r* 
H Z -r> 



Cu u 
^^ o 
s? Z 



■a ^^ 



ZO § 



o 
Z 

Si 3 

3 "5 



1> 



03 



ii <r! 0- 

TO *- 

O 'S5 >- 

5 ^:! 

uo C O 

Du X) ;^ 



r: Q 



o 3 

i_ o 

o -o 

XI c 

aj 

nJ c • 

.1 15 :i 
5 ts = 



•- Z 



D. Cu 



O > C 

r ° I 

S i_ CIS 

a ^G 

«^ ^ 2! 

o Z ? • 



£ S?5 E 



E iS 2 



a — 

CO c« 



o t. 
Z c 



.4/ •- 

> 3 

O JJ 

3 2 

° 3 

XI J 

15 <^ 



X3 



cjo 3 
"O O 

15 -o 
a> D. 

31§ 



•a y 



~'CU C 



r~ "^ X) 
age 

^ JC CO 

-eS^ 



^Q 



x: ^ Ai 

3 .E 3 . 
T3 CO — ?s 



3 !>, ^ [^ 

C C X> _ 

o o _- .E 

o CO 

; - x: i> 

° P 3 O 

Z 5 c X) 



c 

■2 § 

O '-> 
X) 

TO ^J 

«lo Q 

U (U u 

^ r c 

< > Xi 



1> CO 

3 "Z 



<■ I 
=^ s 

C -C 

a| . 

CO ? ^ 

~ — ? 

1» CO ^ 

c .^ ? 

3 Q fc 



^ iT '^ 



id- 3 =^ 



__ C CO 
CO Cb c/5 

o OS S 



O 



I' I 

o 

II 



w- 'o 



aj CO 00 
T3 J= c 
c cj aj 

30^. 

't* j= — Ci 



Si S 
U I 



■a 
2 § 



^S. 



o 



•- -a cu 

o 5; 1^ 
p — 1^ 

5 0^ L^ 

I* js K 

a> O 3 
00 ^ •- 

c5 S O. 



o -s: 



y o 



d 



3 ^-CIO 3 ^ 

° ^ .5 -c o ^ 

ECO -a o c TO 

^ E c 3 J E 

< 55 3 c < b; 



3 a.) 



Ci, in 



f:: s 



c 

e8 

O 00 

D. 3 ^1 

a> CO ts 

'-> _ o^ 

S 2 fc 

15 ^'^ 

— x: E 

13 :2 Q 



j: x: -a 
■53 ° u 



.t; > 



15 Q 






:Ei 




ap_ 


c 

3 




■53 CO 




E 

CO 


g 


= 


„^ 


2 -= 






CO 


c .£ 


E 


60 


c/^ 


U 'C 



s 
a 

•g 






x:=ii-scj='-i 
^S6 O JJ E o § 

•^15 E ^B >:^ 

3 I- CO XI ^ C 

a> w <— x: ^ 3 ^ 

-5 g E 5:5 c c 
^ g -g 5 § T3 « .2 



O 1) 



1) 



-15 " 



tJ x: g .b .1-' 2 

- - t: ^. b o o 



CO o 



^ 2 .i 



3 O 1- 



aj 



a> 



^^ 



— •£ - 
j= ■£ £ 00 c cu 
_ 3 ^ 00— •- ^ 

■Sl5_^ y .SP'Clo 3 ^ 

u aj 3 ■ - 

x: j= 



— OJ w- 



tU c 






90 



Tulane Studies in Zoology and Botany 



Vol. 23 



5 S, 

c a 

c o 
<Z 



^ E 
S 9 

II 

o5. 



2 

3 



I 



(fl a «* 



s 5 

c/5 c 



6 I 

t I 



yi 



I 



XI 

.2 

> 



X> 

.2 
> 









^ S o 






I'll 



'3 '^ 

^ o 



O t3 
J| = 
'- C ^ <b 

— R ^ w 

S S 5, E 
^ ^ i= E 

^ §.Q & 
O E a 






c 
o 

o 
a 

T3 

c 

CO 



> 

s 

-Si 

o 

-S 
ce; 
<u 

E 

o 

H. 

u 

CJ 

X 

1> 

in 

o S 

^•5 



1 



s' 

o 



X X) 

^ >> 

c 5 
^-^ 
E'Z 

CO i> 



c? (U 
OD X 



^ o « — 
■5'ZS d 

*^ C CJ 



3 O 



id S o 



s 5 "? - 



1o 5 A 

— CO w-i 

2 w A 
1/1 m CO 

■^^ /^ = 

En 



u 



CO 



1^ ifl ^ 

U 1) > 



— <L) lU 
> 3 X 



T3 A 



' A 

(U — 



•5 A 






CO CO 



=: CO 



c 2 



o Q 



^ o 



o i: 

1/, c 






5 


^ 






(> 


o 


CO 




Q. 
CO 


C 

CTJ 


CO 


,, 


U 


o 



ti t3 



S - 



CO E 



I I 
s x> -a 



CO 



E 2i 



b a; ii.E 



.5 a^ 
11 

-s: 
.E S- 



5)Q 



<-s 



E 5 

u 
CO "T 
3 /^ 

J2E§ 

3 O '-' 

^■^ O 

£ CO c- 

- V b 
S lo ^ 



CO 






— o .y 



CJ Q.10 

5. V 
>^ — 
H CO 



V 



PD.5? cog !=;-^ 



-O (U o 



CO CO 

c 

'5b E 
« 2 
E- 



C ^ CO 



on CJ U 



c S 

1> CO 

> D 






c .ti 



< T3 



V A 

eg "w 

•^-^ 

iS Xi c^- 
3 CO CU 

E V 
o o 

— ■ w 00 



CO CJ 



CO o il^ 

.e| E 

O in i> 
CO r> u- 



•5 J> 6- 

(U Tj 



_ 3 O 



-2 o «^ 
•E -JQ 

E c lu ;^• 



■Op 

-^ .5 — 



3 3 CO 

CJ t; c ^^ 



•-so 
>> ^ F. 

H •= <ii 



CO 



CO 



o & 



CO C , 
u. L. lo 

c - c 
■| HO Q 

O w- C ^ 

— <U CO T3 

^ §.oS 
•2>75 S.2? 

° J o b 

S^^ § 
J E E 



.5 CJ c^. 

5 o r^ CO 
< Ji 00 CJ 



.5 o 



>. c 



u 



CO w 2 
y ?> "cO „, 



J D CO T3 

'^ CO « -S a 

"listed 



II; 



-. o - 

C OT) .S 



CO 



o« 



D. "^ 
<u 



CU CO 



2 d 



o 

o tS -o -^ 

C S « CO 
CO C -Q T3 



-a 

lU 
Q. 



O 1- 



o E 



T3 ^ 



O CU t^ 

Isl 



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|- 

25 

24 

23 

22 

21 
20 

19 

6 '8 

i ,7 

a 

I 15 

I "* 
_ 13 
€ 12 

f " 
O 

£ 10 
?• 9 
" 8 
7 
6 
5 
4 
3 
2 



■ A 









% " 



o o 



• + ♦ ♦ 



e 



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- 



No. 1 



Malaclemys- Graptemys Relationship 



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. 



98 



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 



99 



o 



CQ 



o 



> -^ 



o 




.^ 


^ 


1> 


T3 






nj 


•o 




3 



U5 



I + + + I 



+ + + + 



+ +-H + + ++ M + + + + 



a 

T3 



o 


% 


c 




o 










9> 






es 


c« 






1) 


u 



■o 

C 




^F 






tS o 


*_, 




p.-o 


o 






a 


j: 


=S-S 


J2 

u 
T3 


■a 


2 S 
o 2 



3 _^ 



_ >" T3 



"5 £? 

.£ =5 

3 "" 
00 >> 

c 13 



o 


j= 








* 




a. 










X! 


■o 


tj 


F 




o 


ni 


o 


c 


QU 




o 


>. 






n 


Lh 


O 


c 



>< o 
W X3 






O 3 





c 
t5 


C 

"3 


'3) 


iS 


CO 


3 


<L) 




c« 


X 


•i 


z 


U 




e 


< 


C3 



li O. c 00 o 

„ _ ■•" C J3 

§ ^ S ■.£ - 

« c o 2 

2 o Ji g o 



■5 J= 



W o 






.S" oo 



3 c -2 .5 



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- 
matics, fossil history, and evolution of the genus 
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 



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 
in CBE Style Manual, 4th ed., 1978. Manuscripts should be typewritten and double 
spaced. Two additional copies should accompany the original to expedite editing and 
publication. Legends for figures should appear on a separate page and in sequence. 
Illustrations should be proportioned for one or two column width corresponding to our 
printed page size, and should allow for insertion of the legend if occupying a whole 
page. Guidelines for letter and other extraneous markings should be done with a 
non-photo blue pencil such as Eagle Prismacolor. Photographs should be on glossy 
paper. 

Many tables, if carefully prepared with a carbon ribbon and electric typewriter, can be 
photographically reproduced, thus helping to reduce publication costs. Lettering in any 
illustrative or tabular material should be of such a size that no letter will be less than 1 V2 
mm high when reduced for publication. 

An abstract not exceeding three percent of the length of the article must accompany the 
manuscript. 

Separates of published articles are available to authors at a nominal cost. 

Page charges, calculated at $45/page, are solicited from authors who have funds for this 
purpose through their institutions or grants. Acceptance of papers is not dependent on 
ability to underwrite costs but excessive illustrations and tabular matter may be charged 
to the author. 

EXCHANGES, SUBSCRIPTIONS, ORDERS FOR INDIVIDUAL COPIES: Ex- 
changes are invited from institutions publishing comparable series. Subscriptions are 
billed in advance. A price list of back issues is available on request. Individuals should 
send their remittance, preferably money order, along with their orders. Remittances 
should be made payable to "Tulane University." Subscription rates: 

Volume 23. $8.50 domestic, $9.50 foreign. 

Copies of Tulane Studies in Zoology and Botany sent to regular recipients, if lost in the 
mails, will be replaced if the editorial offices are notified before the second subsequent 
issue is released. 

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 



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. 

Literature Cited 

Brown, F.A., JR., and M.N. HiNfES. 1952. 
Modifications in diurnal pigmentary rhythm of Uca 
affected by continuous illumination. Physiol. Zool. 
25: 56-70. 

Carlson, S.P. 1935. The color changes in Uca 
pugilator. Proc. Nat. Acad. Sci. 28: 549-551. 

Douglas, W.W198O. Histamine and 5-hydroxy- 
tryptamine (serotonin) and their antagonists, pp. 
609-646 In: A.G. Oilman et al., Eds. Goodman and 
Oilman's The pharmacological basis of thera- 
peutics, 6th ed., Macmillan Publishing Co., New 
York. 

Fantozzi, R., F. Franconi, P.E. Man- 
NAioNi, E. Masini, F. Moroni. 1975. 

Interaction of H,- and Hj- receptor antagonists 
with histamine uptake and metabolism by guinea- 
pig isolated atrium and mouse neoplastic mast cells 



No. 2 



Melanin Migration in Crabs 



107 



in vitro. Br. J. Pharmacol. 53: 569-574. 
FiNGERMAN, M. 1956. Black pigment concentrat- 
ing factor in the tiddler crab. Science 123: 585-586. 

M.M. Hanumante, S.W. 

FiNGERMAN, and D.C. REINSCHMIDT. 
1981. Effects of norepinephrine and norepine- 
phrine agonists and antagonists on the melano- 
phores of the fiddler crab, Uca pugilator. J. Crust. 
Biol. 1: 16-26. 
Fuller, R.W., and D.T. Wong. 1977. inhibi- 
tion of serotonin reuptake. Fed. Proc. 36: 
2154-2158. 

Hanumante, M.M., and M. Fingerman. 

1981a. Responses of the melanophores of the 
fiddler crab, Uca pugilator, to drugs affecting 
noradrenergic neurotransmission: Further evidence 
for norepinephrine as a neurotransmitter triggering 
release of melanin-dispersing hormone. Comp. 
Biochem. Physiol. 70C: 27-34. 

. 1981b. Inhibitory effect of histamine on 

the release of melanin-dispersing hormone in the 
fiddler crab, Uca pugilator. Amer. Zool. 21: 1011. 

. 1982a. Additional evidence for nore- 



pinephrine as a neurotransmitter triggering release 
of melanin-dispersing hormone in the fiddler crab, 
Uca pugilator: The effects of alpha, and beta 
adrenoceptor blocking drugs on melanin migration. 
Comp. Biochem. Physiol. 71C: 15-19. 
. 1982b. Further evidence for norepine- 



phrine as a neurotransmitter stimulating release of 
melanin-dispersing hormone in the fiddler crab, 
Uca pugilator: The changes in the melanophores of 
the crabs following reserpine, 6-hydroxydopamine 
and bretylium administration. Gen. Pharmacol. 13: 
99-103. 
1982c. Pharmacological involvement of 



presynaptic alpha: adrenoceptors in norepinephri- 
nergic neurotransmission triggering the release of 
melanin-dispersing hormone in the fiddler crab, 
Uca pugilator. J. Crust. Biol. 2: 22-30. 
, S.W. FiNGERMAN, and M. 



FiNGERMAN. 1981. Antagonism of the 
inhibitory effect of the polychlorinated biphenyl 
preparation, Aroclor 1242, on color changes of the 
fiddler crab, Uca pugilator, by norepinephrine and 
drugs affecting noradrenergic neurotransmission. 
Bull. Environ. Contamin. Toxicol. 26: 479-484. 



HERMAN, W.S., and S.H. DALLMANN. 1975. 
Linnilus chromatophorotropin: action on isolated 
Uca legs and in various crustaceans. Experientia 31: 
918-919. 

HOGBEN, L., and D. SLOME. 1931. The pig- 
mentary effector system - VI. The dual character of 
endocrine co-ordination in amphibian colour 
change. Proc. R. Soc. Lond. B. !08: 10-53. 

Isaac, L., and A. Goth. 1965. interaction of 
antihistaminics with norepinephrine uptake: 
cocaine-like effect. Life Sci. 4: 1899-1904. 

KOE, K. 1976. Molecular geometry of the inhibitors 
of the uptake of catecholamines and serotonin in 
synaptosome preparations of rat brain. J. 
Pharmac. Exp. Ther. 199: 649-661. 

LOFSTROM, A., and T. BACKSTROM. 1978. 
Relationship between plasma estradiol and brain 
catecholamine content in the diestrus female cat. 
Psychoneuroendocrinology 3: 103-107. 

Marco, E.J., G. Balfagon, J. Marin, 

B. Gomez, and S. LLUCH. 1980. indirect 
adrenergic effect of histamine in cat cerebral 
arteries. Naunyn-Schmiedeberg's Arch. 
Pharmacol. 312: 239-243. 

Owen, D.A.A., C.A. Harvey, and R.W. 

GRESTWOOD. 1979. Cardiovascular studies with 
impromidine (S. K. and F. 92676), a new very 
potent and specific histamine Hj-receptor agonist. 
J. Pharm. Pharmacol. 31: 577-582. 

PANTIN, C.P.A. 1934. The excitation of crusta- 
cean muscle. J. Exp. Biol. 11: 11-27. 

POLANIN, A., and J.H. McNEILL. 1981. Char- 
acterization of the histamine receptors in rabbit left 
atria. Can. J. Physiol. Pharmacol. 59: 19-24. 

T.E. Tenner, jr., and J.H. 

McNEILL. 1981. The characterization of cardiac 
histaminergic chronotropic receptors in the rabbit. 
Can. J. Physiol. Pharmacol. 59: 14-18. 
SANDEEN, M.I. 1950. Chromatophorotropins in 
the central nervous system of Uca pugilator, with 
special reference to their origins and actions. 
Physiol. Zool. 23: 337-352. 

Terrasawa, E., W.E. Bridson, J.W. 
Davenport, and R.W. Gay. 1975. Roie of 

brain monoamines in release of gonadotropin 
before proestrus in the cyclic rat. Neuroendocrin- 
ology 18: 345-359. 



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 


» 



112 



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, 



1 = E c E c = 3 c 



iSSoi^^uBS -iSmSS £ 2 



g I II 









S i- 8 S o S S 5 : 



11 |J 



lli £ 



;j|l|S|sil|u|S|i|(;|a 






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. 

Literature Cited 

AMEEL, D.J. 1934. Paragonimus, its life history 
and distribution in North America and its taxo- 
nomy (Trematoda: Troglotrematidae). Am. J. 
Hyg. 19: 279-317. 

Babbot, F.L., W.W. Frye, and J.E. 
Gordon. 1961. intestinal parasites of man in 
Arctic Greenland. Am. J. Trop. Med. Hyg. 10: 
185-190. 

BABERO, B.B. 1960. A survey of parasitism in 
skunks {Mephitis mephitis) in Louisiana, with ob- 
servations on pathology due to helminthiasis. J. 
Parasitol. 46: 26-27. 

1972. A record of progenesis in Trema- 
toda. Proc. Helm. Soc. Wash. 39: 128-131. 

, and J. Lee. 1961. Studies on the hel- 



minths of nutria, Myocastor coy pus (Molina), in 
Louisiana with checklist of other worm parasites 
from this host. J. Parasitol. 47: 378-390. 

BAFUNDO, K.W., W.E. WiLHELM, and 
M.L. Kennedy. 1980. Geographic variation 
in helminth parasites from the digestive tract of 
Tennessee raccoons, Procyon lotor. J. Parasitol. 
66: 134-139. 

Beaver, P.C. 1941. studies on the life history of 
Euparyphium melis 'Trematoda: Echinostomi- 
dae). J. Parasitol. 27: 35-44. 

Bennett, H.J. 1938. a partial check list of the 
trematodes of Louisiana vertebrates. Proc. La. 
Acad. Sci. 4: 178-181. 

, and A.G. Humes. 1939. studies 

on the pre-cercarial development of Stichorchis 
subtriquetrus (JTemaioda.: Paramphistomidae). J. 
Parasitol. 25: 223-231. 

, and L.L. Jenkins. 1936. The longe- 



No. 2 



Trematodes of Mammals 



121 



vity of the miracidium of Cotylophoron cotylo- 
phorum. Proc. La. Acad. Sci. 8: 5-13. 
BRIDGMAN, J.F. 1969. Life cycles of Carneo- 
phallus choanophallus n. sp. and C. basodactylo- 
phallus n. sp. (Trematoda: Microphailidae). 
Tulane Stud. Zooi. & Bot. 15: 81-105. 

Burrows, R.B., and W.G. Lillis. i965. 

Trematodes of New Jersey dogs and cats. J. Para- 
sitol. 51: 570-574. 

BYRD, E.E., and R.W. MACY. 1942. Mam- 
malian trematodes. III. Certain species from bats. 
J. Tenn. Acad. Sci. 17: 149-156. 

, and R.J. REIBER. 1942. Mammalian 

trematodes. II. Three flukes from small mam- 
mals. J. Tenn. Acad. Sci. 17: 143-156. 

, and M.V. Parker. 



1942. The anatomy of a lung fluke from the opos- 
sum (Didelphis virginiana Kerr). J. Tenn. Acad. 
Sci. 17:116-129. 

Chandler, A.C, and R. Rausch. i946. a 

study of strigeids from Michigan mammals, with 
comments on the classification of mammalian 
strigeids. Trans. Am. Microsc. Soc. 65: 328-337. 

Custer, J.W., and D.B. Pence. 1981. Ecolo- 
gical analysis of helminth populations of wild 
canids from the Gulf Coastal prairies of Texas and 
Louisiana. J. Parasitol. 67: 289-307. 

DAWES, B. 1946. The Trematoda with special refer- 
ence to British and other European forms. Cam- 
bridge Univ. Press, 664 p. 

DiKMANS, G. 1931. A new nematode worm, Vian- 
naia bursobscura, from the opossum with a note on 
the other parasites of the opossum. Proc. U.S. 
Natl. Mus. 79: 1-4. 

1945. Check-list of the internal and ex- 
ternal animal parasites of domestic animals in 
North America. Am. J. Vet. Res. 6: 211-241. 

Dubois, G. 1937. Sur quelques Strigeide. Rev. 
Suisse Zool. 44: 391-396. 

1944. A propos de la specificite' parasi- 

taire des Strigeida. Bull. Soc. Neuch. Sci. Nat. 69: 
5-103. 

1970. Statut des Alariinae Hall et 



Wigdor, 1918 (Trematoda: Diplostomatidae) et 
revision de quelques alariens. Bull. Soc. Neuch. 
Nat. 10: 259-727. 

Fleming, W.J., C.F. Dixon, and J.W. 

LOVETT. 1977. Helminth parasites of river otters 
{Lutra canadensis) from southeastern Alabama. 
Proc. Helminthol. Soc. Wash. 44: 131-135. 

HARKEMA, R. and G.C. MiLLER. 1964. Hel- 
minth parasites of the raccoon, Procyon lotor in the 
southeastern United States. J. Parasitol. 50: 60-66. 

HERBER, E.C. 1942. The life history studies on 
two trematodes of the subfamily Notocotylinae. J. 
Parasitol. 28: 179-196. 



Hoffman, G.L. 1957. studies on the life cycle of 

Cryptocotyle concavum from the common sucker 

and experimentally in the chick. Proc. North 

Dakota Acad. Sci. 11: 55-56. 
Kaplan, E.H. 1964. HeterobUharzia americana 

Price, 1929, in the opossum from Louisiana. J. 

Parasitol. 50: 797. 
KINSELLA, J.M. 1971. Growth, development, and 

intraspecific variation of Quinqueserialis quinque- 

serialis (Trematoda: Notocotylidae) in rodent 

hosts. J. Parasitol. 57: 62-70. 
Larson, O.R., and W.C. SCHARF. 1975. New 

helminth records from Minnesota mammals. Proc. 

Helminthol. Soc. Wash. 42: 174-175. 
La Rue, G.R., and D.J. AMEEL. 1937. The 

distribution of Paragonimus. J. Parasitol. 23: 

382-388. 
LUMSDEN, R.D., and C.A. WiNKLER. 1962. 

The opossum, Didelphis virginiana (Kerr), a host 

for the cyathocotylid trematode Linstowiella szidati 

(Anderson, 1944) in Louisiana. J. Parasitol. 48: 

503. 
, and J. A. ZlSCHKE. 1961. Seven 

trematodes from small mammals in Louisiana. 

Tulane Stud. Zool. & Bot. 9: 87-98. 
, and 1963. Studies on the 



trematodes of Louisiana birds. Z. Parasitenk. 22: 
316-366. 

Malek, E.A., L.R. Ash, H.F. Lee, and 

M.D. Little. 1961. HeterobUharzia infection 
in the dog and other mammals in Louisiana. J. 
Parasitol. 47: 619-623. 
Martin, D.R. 1976. New host and distribution 
records of helminth parasites of the Mexican free- 
tail bat, Tadarida brasiliensis, from Texas and 
Louisiana. Proc. Helminthol. Soc. Wash. 43: 
85-86. 

Miller, G.C, and R. Harkema. i964. 

Studies on helminths of North Carolina 
vertebrates. V. Parasites of the mink, Mustela 
vison Schreber. J. Parasitol. 50: 717-720. 

, and 1968. Helminths of 

some wild mammals in the southeastern United 
States. Proc. Helminthol. Soc. Wash. 35: 118-125. 

NASIR, p., and M.T. DiAZ. 1972. Avian flukes 
from Venezuela. Rivista di Parassitologia 33: 
245-276. 

PENN, G.H. 1942. Parasitological survey of Loui- 
siana muskrats. J. Parasitol. 28: 348-349. 

Rausch, R. 1946. The raccoon, a new host for 
Microphallus sp., with additional notes on M. 
ovatus from turtles. J. Parasitol. 32: 208-209. 

1947. Some observations on the host re- 
lationships of Microphallus opacus (Ward, 1894) 
(Trematoda: Microphailidae). Trans. Am. 
Microsc. Soc. 66: 59-63. 



122 



Tulane Studies in Zoology and Botany 



Vol.23 



ROTHENBACHER, H., and W.D. LlND- 

QUIST. 1963. Liver cirrhosis and pancreatitis in a 

cat infected with Amphimerus pseudofelineus. J. 

Am. Vet. Med. Assoc. 143: 1099-1102. 
SAYWER, T.K. 1961. The american otter, Lutra 

canadensis vaga, as a host for two species of trema- 

todes previously unreported from North America. 

Proc. Helminthol. Soc. Wash. 28: 175-176. 
SHOOP W.L., and K.C. CORKUM. 1981a. 

Some trematodes of mammals in Louisiana. 

Tulane Stud. Zool. & Bot. 22: 109-121. 
, and . 1981b. Epidemiology 

of A laria marcianae mesocercariae in Louisiana. J. 

Parasitol. 67: 928-931. 

_, and . 1982. Progenesis re- 



considered in Ribeiroia ondatrae (Price, 1931) nee 
Babero, 1972, a junior synonym of Gyrosoma 
singulare. J. Parasitol. 68:424. 

SKRJABIN, K.L., and E.I. BASHKIROVA. 
1956. Echinostomatidae Dietz, 1909. In 
SKRJABIN, K.I., Trematodes of Animals and 
Man. 12: 53-932. 

SOGANDARES-BERNAL, F. 1961. Sellacotyle vi- 



lellosa, a new troglotrematid trematode from the 
mink in Louisiana. J. Parasitol. 47: 911-912. 
. 1965a. Parasites from Louisiana cray- 



fishes. Tulane Stud. Zool. & Bot. 12: 79-85. 
. 1965b. Studies on American paragoni- 



miasis. 1. Age immunity of the snail host. J. Para- 
sitol. 51: 958-960. 
, and J.F. BRIDGMAN. 1960. Three 



Ascocoiyle complex trematodes (Heterophyidae) 
encysted in fishes from Louisiana, including the 
description of a new genus. Tulane Stud. Zool. & 
Bot. 8: 31-39. 

Thatcher, V.E. 1970. The genus Amphimerus 
Barker, 1911 (Trematoda: Opisthorchiidae) in Co- 
lombia with the description of a new species. Proc. 
Helminthol. Soc. Wash. 37: 207-211. 

WOOTTON, D.M. 1957. The life history of Cryp- 
tocotyle concavum (Creplin, 1825) Fischoeder, 
1903 (Trematoda: Heterophyidae). J. Parasitol. 
43: 271-279. 

YAMAGUTI, S. 1971. Synopsis of digenetic trema- 
todes of vertebrates. Vol. I. Keigaku Publ. Co., 
Ltd., Tokyo, 1,100 pp. 



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 



dE'xaaAO 
Xsupiii 




^OOC/iOO C/50C/) 

■b "b 'b^ ■b -D ^ tj ^ 


z 
■b 


/I CO 

z z 

bo 


yjtotototototoocococototncotocotoo "3t2i3 
zzzzzzz -zzzzzzzzz^zzz 

bo'b'bO'b'b'b'b'D'b'b'D'b'b'bo*" ^ii°* 


"b 


z z 

■b -b 


aOEdSJ33UT 

J8ATT 

-IJEBl) 


b 

■b 


O 1 

jn c/1 c/1 t/) o 1 

Z Z Z Z • 1 

^■bf>--b ^ 


Z 1 1 

■b 


z 

■b 


II 


1 IZZZZZZIZZZI IZZZI 'g^ 

'b^i'boo'b ooo 'o'b'b '^ "o 


1 


to o 




o 

■b 


~^l o o -1 in -I 
o o o o o o 

■b 'b -b -b ■b -b 


*-l 
n o 
o o to 

■bbl 


z 

•b 


o 

to O 


oo^^ .-H^o o inocNo^ o ^ 
ooootoooocootooooootoo ^o^ 

■b-b'o'bl'b'b^^'b^'b'b'b'b'b^'b ■b'b'b 


o 

o 

b 


o 

CO o 


H3du3x 

A3Upi51 -J 


o 

b 


-M -1 O -H m 
O O O O CO o 

•b "b •£ 'b^ "b 


to o to 


CO 

z 

■b 


o 

CO o 


.-1 O O .H .H O 

totototoototootoototootoococoo MCOCO 
zzzz-zz.*z.*zz.*z.-zz.* zzz 

■o'b'b'b^'b'b^'bT^'b'b^'b^'b^^ iioo 


o 
o 

•b 


o o 
■b-b 


II53U3X 

•Xq TIES 


1 


z z z z ri 1 

0.0.r,.,> -n 


1 1 1 


1 


Z 1 

■b 


1 1 ! itototootoitocoi icocol 1 'H;' 

1 1 1 IZZZ -ZIZZI IZZl 1 IZl 

■bo'b'^'fa ^'> c/*o* o 


] 


1 1 


J3AIX 


1 


c/1 c« to O C/1 1 
Z Z Z • Z 1 

o. a. "D o. c» 


1 1 1 


1 


to CO 

z z 


1 1 1 itotototooitotoi ItOOl 1 ii"] 
1 1 1 IZZZZ -IZZl IZ-I 1 IZl 

o ■□ ooO "bo o*o o* 


i 


- 


ASUpi^l "X 

•3S0d 


■b 


z z z z z z 

•b -0 o ■bOo 


z z z 


z 

o 


z z 

■b -b 


inorgino-— 1 in CN m t-HOm 
tococooooooococoocoocoococo ooo 

^ooooooooooo'-b'^o^o'b <><>o* 


z 

o 


z z 

o o* 


Aaupi^ -X 
•3UB 


1 


O ^ O CNi ,-( 

CW o o o o o 


^ o 

O O C« 


o 

o 


to O 

o o 


0'-l*H^OOOOOO^'-l O^OtM^*-^OC^ 
OOOOOOOOOOOOCOOOOOOOOO 

o*oo*oooooooooo*o*oooo ooo 


o 
o 

o 


o o 
(> o* 


CH Cw o o ^ ^ 


O* O* o* 


Aaupt^i -J 
■ 3Sod 


1 


en c/l en to en c« 
z z z z z z 

•d o -D -o 'b o 


to to CO 

z z z 

0*0 0* 


z 

o 


to o 
z • 

oo 


,_4,_i(/^,_)0'Hinooo CM i-i cNjin --lo 
ooooooooootoocootoootoooto 
2 -2 'Z • 'Z • 'Z 

OOOOOOOOOOo.O*oOo*OOi^3 OOo* 


z 

o 


to O 

o °* 


Xaupt^i -a 
•3UB 


1 


C -^ C 

tn o o tn to o 
z • • z z • 

0+0*0*0 0*0* 


in o 
o o to 

O O o* 


o 
o 


o 
to o 


^r-< .H^^ ^r-I.H ^ -H.-I •-* ^ 

OO^OOO-HOOO'-lO.-I^OOrHO ino 

oooooooooooooooooo oo^ 


o 
o 

o 


tN O 

o o 
o o 


o o 


000*OOOC>000*0*IOOOOOOO OOo 


•xq -xTeS 
•qsod 


1 
1 


1 
to to to to to I 

Z Z Z Z Z 1 

o-b-oo*^ 


1 1 1 


1 


in 1 

O 1 

-6 ' 


llll cNinl 11^ III 1 
1 1 1 itntocoooitotoi lotoi 1 igl 

1 1 1 IZZZ ■ -IZZl 1-ZI 1 IZl 

■b ■doO*'^ -b -o ^ -b -b 


1 


1 1 


jaAXX 
• 5Sod 


1 


to to to CO to 1 

z z z z z 1 

0+ "b "d o* "o 


1 1 1 
1 1 1 


1 


c 
o to 

b? 


1 1 1 Itotototooitotoi lOOl 1 1^' 
1 1 1 IZZZZ -IZZl l.-^-l 1 IZl 

o*.boo^ •bo bo II 


1 


1 
1 z 


J3ATX 
■3UE 


O 

■d 


O eg 1 

to o to to O 1 
z • z z • 1 

■b b o* -o •□ 


O 1 1 

■o 


o 


o 

o to 
• z 

■d 'b 


111 r-H 1 1 in 1 --^ 

1 itoitototoototototoi itootoiotoco 

1 IZIZZZ-ZZZZI IZ.-ZI'ZZ 

•d fe b II ^ "d o -d o -d d o ^ ■o\) 


i 


^ o 
o o 


aJEaq 
•jsod 


z 


u-1 .-1 O 

o o to to o to 
• ■ z z • z 

■b ■b 'b o •b o* 


to to to 

z z z 

•b o "b 


o 

■b 


o 

o to 
• z 

■b -b 


1 .H o in .-1 1 in 
tococoto 1 totootototocoototootoo oto 
ZZZZIZZ -ZZZZ -ZZ -Z •! -z 

•b'b'bb ^■b>Doii'bo'b'b'b'bo'b "bii 


o 
o 


O '-I 

o o 

■b -b 


CO / 

CO / 
•C / 
CJ / 

/ ^ 

/ c3 


'a 
c 
c: 

1. 
a: 

."■ 

C 

c 

K 


O .Sf -til ^ tJ 't! 

<<S -W O g C» -rJ 

o :5^ CO o ix^-^i 
^ <» ^ i^ to s 

.a 

-a 

g 


to .s; -vJ 
i^ « o> 

<3 

s; 

<32 


to 

« 

Cm 

1 

1^ 


o 
o 

•tJ 'pi 

2i SX 

13 O 

.a 

!, 
CO 

o 
to 


g^<r.n^cr- +^t3to O <« 
O g.<;mra<tm Sati-aeo caeocococo 
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 

l3-"!,33t*c»ts3'9SH<~> en's P ^ V a 5^ .^ -^ 

^.QU5jy^<J)<a'a(35EEs;0!Xf*s.»«>coco 

CO 


3 

« 

c 

o 

£X 
o 


C3 S* 

•vi IS 

(3 
•t-1 

i 



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 ■-< 


^ 


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 
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: 



Xaupi:^ 






q^Suai; 
Xaup"p^ '2 



jaATi 



•3Sod 



Xaupx^ 'X 



-* vO ro CT\ lO 
r~ cs >— I I— t u-i 



0-) iH vo rH r^ I 
r-| t-t CN ~3- r^ I 



r^ r^ -J- ~* iH ro 

U-| iTl lO lO lA -J" 



ON I I r^ 




n in vo I 



1 I I l~- 



O rS; -v^ rCl "tS ^ 
t-^ +^ 1^ E <i) -f^ 
^ S5 CO O Cuf<s 

35 ?^ ,a r« -^ « 



•v^ E q fc 

Q) rC -r^ +^ 
« (35 ?^ CO 



o 

E 
o 
-p 

S5 tX, 

(3 -^ 
(U t) 

"Xi o 

(3 






■-C CO 
O 4^ 



I v£> o in 



CO "5^ 




s 


35 ^ 




« , 


rs; Q) 


CO 


•r^ w 


O '--i 


^"ll 


t) -t^ 


a +^ 


?; 


?; 'tl 


S^ 3 


(■V 


(3 « 


^ rCl 


*i) 


fc- !^ 



(3 U 



No. 2 



Visceral Topography of Snakes 



133 





^ 










vO^Om^o^oo^ 1 O^O 


vO V.O ^D 0^ 0^ 


■H 1 


\0 CM 


3DBdsa9:}ut 


(NCS)CN<N<NI(N |r-ICN 


>H (N CM iH iH 


r-s 1 


CN CO 


a3Ai][-5ap3q 


m a> r^ in rH 1 i ~* i 


in sj- o o CM 


1 1 


CM m 


rH rH fO rH ,H 1 1 O" 1 


rH sj- rn rO CN 


1 1 


CN iH 




<rrH,-i>^,HcNjrgvooo 


>H rH c^ m o 


CO -3- 


00 CJs 


dB^aSAO 


1— l-a'iHirivJ-vOCMi— IvO 


in m -a- -* r-i 


r^ 00 


r^ in 


AaupT^ 


•— lrO<3-CTN<rrHvOU~im 


in oo CO .H -vf 


<r 1 


so in 


m<rm<tm<f-*rovo 


m vD -3- -^ CN 


in 1 


r^ vD 




omc^^o^>^r^lrlCT^<^J 


.H ro iH CJN 00 


CM VO 


m CN 


qaSua^ 


ro<rro<fin<T(^iHm 


in <}• m <f ro 


<7s 00 


OS C3S 


it9UpT5l •\ 


ro-^r^vo-— icNr^cNCN 


CN r-s CM ro 00 


■H 1 


O SO 


<fcNmr^~3-rocNroro 


ro in vo <r ro 


00 1 


O 00 




iHromvOC»0000-*iH 


iH -d- CN ON OS 


CN rH 


r^ CTs 


qaSu9X 


<f<l--cr~3-mvX3rOCN<t 


in 00 ro <r -J- 


<7S 00 


OS 00 


Xaupxit -a 


rO00r^rsl^nO^r-~\OvO 


in o> r-s rH tH 


<r 1 


r-s CN 


<fo-|<f^DrO.HCN^-<f 


ro in in <j St 


00 1 


OS CTs 


q:)3u3i 


1 n Lo m Lo ro 1 1 1 


1 a> 1 in in 


1 in 


00 vO 


1 m CM m cvi ro 1 1 1 


1 r^ 1 CN CM 


1 rs 


00 OS 


aaATj 


CM rH ,H M ^ 1 1 1 1 


1 ^O 1 .H 1 


1 1 


1 1 


-* CNI CN CM CN 1 1 1 1 


1 CN 1 CN 1 


1 1 


1 1 


XaupT^i '\ 


ro<r)rHfon2rovoin 


00 CO <■ in 00 


CO CM 


CO -sf 


CO rs 00 CO CO 


sT CO 


<r in 


• 5 sod 


O^a^r000u-lOv£)'-l^-- 


iH CN r^ in cN 


CO 1 


o o 




u-imr^r^ror^<i-oou-i 


~* vO in vD vO 


<r 1 


CO CO 


XaupT^i '1 


rHr~<3-rHvOCNCT\0000 


~d- 00 CO tH si- 


St CO 


s3- -J- 


mu~imoo<j-CT\-3- ^n 


in r-s r-^ in in 


00 rs 


00 00 


• 5UB 


rOrOONcNinooOvooO 


M .H 00 vO 00 


<r 1 


00 so 




r^r^0OQ\vDr^^ r^\£> 


m 00 r^ r^ \o 


00 1 


r-s r^ 


iC9up-i2i -a 


vCvO^fvOONr^LOvOOO 


in CNi ^o in CT> 


r-s 00 


so in 


oor-~oooo<roNror^m 


^o vc r^ CO ^ 


Csl CO 


St CO 


• 3 sod 


vomoOOOONtHCM iH iH 


OS 00 i-H CO tH 


r^ 1 


.H C3S 




r^r^r^r^^oovooo m 


in vD 00 t-s in 


in 1 


in <r 


Xgupiij • X 


a^00cOO^CT^CN^o^M•-3■ 


tH 00 O OS CJN 


<)■ 00 


so <f 


L^lvo^^oo<ra^<t■^om 


in r^ r^ in in 


00 ts~ 


00 CO 


• 3UB 


rHOOONVOrOrH^a-OOr^ 


sj- tH vo vO S3- 


vO 1 


CM so 




c»r^oooor-.oOLor^Ln 


in 00 00 r^ in 


00 1 


OS 00 


SBgjDUBd 


1 0> 1 lO ro n 1 1 1 
1 m 1 ro ^ vo 1 1 1 


1 m 1 in cN 

1 vO 1 rO CM 


1 .H 
1 OS 


00 00 

St sl- 


• 3 sod 


m u~i ro vo r-^ 1 1 1 1 
vo rn in r^ ^ 1 1 1 1 


1 CN 1 cys 1 
1 00 1 in 1 


1 1 
1 1 


1 1 
1 1 




1 O 00 ^o o vr> 1 1 1 


1 00 1 o 00 


1 vo 


00 so 


a9AT^ 


1 vD ^D r-^ vC c^ i 1 i 


1 00 1 00 sl- 


1 r-s 


so CJs 


•3 sod 


CTN <r ON a^ S3- 1 1 1 1 


1 ON 1 St 1 


1 , 


1 m 




r^ 00 rs. rs 00 1 i i i 


1 rs. 1 00 1 


1 1 


1 OS 




1 00 00 o <!• in 1 00 iH 


in iH -sT m in 


1 CN 


CN St 


agATi 


1 vo in cTv r-s in 1 v£) vo 


in rs r~. vo in 


1 St 


in r^ 


•auB 


-a- 00 CN CM VO 1 1 iH r^ 


ro rH -sT OS .H 


1 1 


00 o 




in \o 00 00 rn 1 1 ^£3 in 


sf r-s. vO 1^ vo 


1 1 


so in 




CNOOOOv^OOCN y£) ~* 


CO s^- C30 in cNi 


vD r^ 


r^ o 


:}aB9i{ 


»or^vooooOvc^D r^ in 


sj- in vo vo vo 


00 OS 


in rs 


• 3 sod 


mrOCTsrsir^^rroO 1 


sO OS O CO OS 


CO 1 


t^ vD 




vomoocTvvoinvc^ i 


vD 00 00 00 00 


00 1 


r-s <j- 


/ 


CO 


^co 
3 


<: PQ 




/ 


!^ CO 3 


r-:» 


^ p 




u / 


^ •ri +i 


a 


s s 




01 / 


+:> to CO (3 


rs; < m 


-w +i 




4J / 


<cQ<;mcocjQ) +^ 


Cl, CO 


(3 (3 




y / 


a rSr "^ O CO 


^ -^ CO CO 


Q) ^ 




<8 / 


•r^ -c^ CO CO Cn ^ -ri S 't^ 


t) CO •<>s -t-i -t-i 


s s 


(3 


i-i / 


•r^-r^csoao S !^ 


o ex, s f-i t-i 


•<J -ri 


I<S Q) 


* / 


^rs;o(3£;c)s; fxa 


CO o a a a 


rss T-~i 


3 a 


6 / 


OOCTiCDyS^-rJ-r^t^i 






■<^ -fj 


3 S (U <i) r-i Cn'tJ H-, a 


S 


a s^ 


/ 


0<ycaQ)EK;o S CO 


rS; 35 o -t^ •!>S 


O 


•rJ cu 


/ 


y y (33 CO CO 


•fJ 


SL, ^-i 


/ 






SS 


■t-^ <3 


/ 


CO 


CO 


O 


CO ;i 


/ 


•ri 


•r^ 


t-^i 




/ 


r«; 


rs; CO 


ti 


a 


/ 


fX CO 


EX -^-i 


rS 


•t^ 


/ 


o p 


O t-^ 


^ 


s 


/ S 


s 5 CL 


S « 5- 


•V^ 


•t-i 


E (33 3 


fc 4-i 3 


£X 


<35 


/ X 


<3 <i) o 


Q s^ o 


O 


S^ 


/ ^ 




6-1 Co 00 


^ 


•:^ 



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 
. 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 
. 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 















■^^ 




/-^ /-^ ^-y /'-v ^^ ^-^ 


^■^ /~^ ^-^ 


^*^ 




/.-S 


/-N r^ 




o 00 o m <r -» 
cN rn fn rH en en 


<T. 00 -J- 
in CM s£) 


in 


sO<rini£iO-3--3-r-r-~rsienr^rH 





• 

. rH 

-3 1 

1 in 
• 

. rH 
r-\ 1 


enso<J-in<}-vDr-tcMso<roocn-<r 


o 


1 1 1 1 1 1 

%c o r-, o vo vc 

CD ^' ^' ^' O -H 


1 1 1 1 
CM in -H 1 

... 1 

f-H r-l CM 


1 1 

1 
• 1 


1 1 1 1 1 1 1 1 1 1 1 1 1 1 
1 O<rOCMr^Q0P^CM<r-3"enr^o 




en 


rHCMcncMCMi-IOOmOCMOCM 


















CM m 00 fn 00 CM 
1-1 CNl I—I rH rH Csl 


in <r vo 
en CM en 


'^ 


en<7sQ0asLnt7s(7\(7sCMincMp^f-l 


in 
en 


CM 

c--i 


j: 


w 


cMcnenenencnOr-iineMincMcn 


u 


as 














C 














B 


<u 














0) 




r^ O i-~ >H o 


<r 


CTs rH 


cnr^oOiXJr^'— ir^r^ <j-in rH 






Vj 


(Ni <r cN <r <r 


<r 


en en 


incnsD^tco^ToocM r^in m 






(U 


<v 


1 1 1 1 1 1 


1 1 1 1 


1 1 


1 1 1 1 1 1 1 1 1 1 1 1 1 1 


1 


1 1 


o 


•o . 


u-i o o o en 1 


1 1 1 00 


m 


enOcnininr~.r-^cn lOO 100 1 


1 


1 1 


nj 


TD JD 


1 


1 1 1 • 




1 • • 1 • 1 


1 


1 1 


a 


CO Id 


O <N ^ 1-H O 


rH 


is 


rHCMCMCMCMr-I.HCM cnc^l CM 






CO 
















(V 


x> 


en cJ^ r^ O en 


-3- 


en \D 


osvocnincMp^cMvo ooo 






^ 


.— t CM ^ CNl tH 


en 


CM rH 


cMCM>3-cn<tcs4encM inen -3- 






B 


I— 1 














•rf 


CO 














M 


















^^ ^-- /-. 


^^ 




^-v 


^^ /-^ 




<r tN iH Ov O i-H 

en <T m CNi m m 


en m ^ 
en CM en 


OS 


LninP^in(3sr^r^\D0OCMint-H^ 


en 


rH m 
en en 


CMCMCMCMrHCMCnCMT-lmcMcncM 
1 1 1 1 1 1 1 1 1 1 1 1 1 ' 




1 1 1 1 1 1 

rH <r CNl ,-1 CN CN 


1 1 1 1 

vD en m 1 


in 1 


1 cjsenr-iooen^r'— tencsjoocMcnos 


CM 


r- vD 








1 






c 


CN rH CM CNI CN CN 


rH CM iH 


.-( 


I— IrHCMi— 1<— If— ICM(— IrHrHi— IrHrH 


CM 


•-\ ^ 


01 














00 vo r- <3- in -a- 

CN CM CM CM CM CM 


-3- <r ^ 


CM 


f— lO^fcMr^rHini— ir^iHincMf-i 


Csl 
CM 


CM CM 


CMCMCMCMrHCMCMCMiHCMi-IC-JCM 


T3 - 














13 














« 


O ^ ^ C3> VO 





-a- <r 


0Oi-ienir>in<rvD'-i <rin <r 






J3 


CM CM cn CM cn 


en 


CM en 


CMcncMCMtMCMCMCM CMCM CM 






03 


1 1 1 1 1 1 

<t o in <i" in 1 


1 1 1 1 

1 1 1 


in vc 


oooocMincMcMcno icMcn 100 1 


1 


1 1 


1 


III* 


rH >-l 


rHrHiHCMrHrHiHCM r-\ ^ r-A 






00 














CN CM -3- O 00 


in 


CM <f 


cn<-Q0inr^rHrHO lOO 








^^ ^ ^^ 


t^ 


^ ^' 


oJ csl ^ ^ .^ cs] r.' cm" Ac:, o] 






1) 


CTv 











CM 

en 


u 


<f ^-s r-^ en cr^ ^ 


en <r in 


p-~ 


-3-os^^-vcn^^r^r-<— Iino0v£) 


<r 


-a- /-v 


(0 


^N en ^^ ^^ ,~. ^~. 










^^ in 


& 


<f . p^ m <r <t 


in r^ i£> 


CM 


CJsos . .vXJOsOOSinsOOsOsOO 


00 


00 

• in 


fi 


. o • • • • 










u 


^ T— 1 ^D ^ r-^ r^ 


<y\ \0 (y\ 




vDCTSrHrHr-OOrOinOsCOOOsI-^ 


00 


00 1 


5 o 


1 1 1 1 1 1 


1 1 1 1 


1 1 


1 1 1 1 1 1 1 1 i 1 1 1 1 1 


1 


1 -3- 


Z t> 


o o .-H -H <r ^c 


^ in 1 


CM 1 


1 osenvD^f-ioo-3-sOrHenooooeM 


i~^ 


. CM 


c 






. 1 


1 




T-t 


CM <r en CM iH en 


<J en \0 


CM 


rH<rr^min<tT-iOi^<— lenfHin 


vO 


rH 1 
















0) 
13 


CM en O ■-• CM <f 
m in in en <■ in 


CM r^ <f 
p^ m 00 


CM 


^m^r^rOSf— tCMOOsOiHCOCMCM 


C7S 


<r -H 


^^^o^oOln'^cMm^^ln^vOsc 


to ■ 








vD 






XI 


^ iH 






t^ ■-< in t-i 








00 r- 00 vD iH 


<f 


00 \D 


^ in rH y-^ ^N \o ^*^ en Oi /^ c]0 






.H 








y^/— s/— vin^fy'vcM/^ ^sin ^ 






r-H 


r~ 00 r-H 00 en 


<r 


r-l C3S 


in r^ • • vD • <■ OS . CM 






(0 








■ . . rH . CM . • 






60 


m C30 vo m vD 


c^ 


sO <r 


CTSvOCJSt— li— IvOtHin OOi— 1 CJS 
1 1 1 1 1 1 1 1 1 1 i 1 1 1 




1 1 


<U D 


1 1 1 1 1 1 

r^ .H en oj 00 i 


1 1 1 OS 


<r I-- 


OOenenmOincTs ir^oo lio 1 


1 


1 1 


O <r CM CN o 


<f 


CM r4 


cninin^<rcn-3-<r <r-3- sd 




















.H 


CO en CT^ VD ^ 





r^ en 


cnoinosencMoocM mm ^o 








CM vX) en en en 


00' 


<r r^ 


^^f^oor-^ini — ^in r^r^ p^ 








S^ 


a 




to 








Ci) 


+i 


s 


^•ri 








+^ 








Q> »--i 


s 






CO <3 


-w 




-u C3 <: oa 


pa 






S « Sh 


+^ 


^ 


<:pqco<:ffl toco.* 

•vi S«Oeo cococo 


■tj 


<3 




O C35 <3 ^ 


■^ -t-i 


c^ 


!S g 


t-i ca 




•ri O 4^ 't-. C 


•<J s 


+i 


to -ri "tJ to CO to S <3j 3 S S -ri -iJ 


S 


s _a 




O .JC "t^ -Cl ^ ^ 




•tJ -^ 


•ri-«->-^a,Ks; «osE +jtoTO 


r~-i +i 


« « S 




a ») ix 


,«.c.s;oooto-^s;ov^"^'3a 

OS3?HC»<B2!S'<>(3>0^3t.SH 
poo Sj'--' t-^ <3- (3 <a •vi Sh <3 a -vi -ri 


« 




C3 T-^ -^i O E ci> -T-i 


(i .s; -^ -u 


trS'?^ 


.S ^ 






•t^ o Si CO o a.'-i 




t3 S 


c 


cs Q> y 


•t^ -tJ 


■tJ -M « 


O 


t) C» H-, Ss CO a 


•<J « Gl !^ to 


s. 13 


g<D«OcBCflcaEEs;CL,S^coco«o 


Ch'-i 


5jj CO a 


^ 


Sh 


tn 










s^ 




H 


=^ 


^ 


03 


g 


s 


S 



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 



o o o o 



^r~- OO OO OO OO OO OO OO OO 0-10 oo 



.HO OO OO OO OO tr^o- OOCsl OvO OO OO 



cnon cNro OO c^jn OO oOvT) cn(>j on OO OO 



OO H04 OO Or^ OO <r.H OO OO HO 



CN ^D O O O 



OO OO OO OO 



r^O <rro OO OO 



OO OO OO OO OO 



CO OO OO OO rooo >x>m n-)<r oo oo 



OO oo oo oo oo oo oo oo 






gs g 



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 2 i 6 



% Males 

10 12 -4 2 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 



< m < 






m -, 



o 



fs CM ,^ "ro 



CN 



CO tx 

t^ IV K f^ 
c^ ^ "^ ts. 



N (X 



» o- 



e* CNK Nootor^'^hv 
fv 00 rs o> 

00 



rv. 00 



0» « 



GO o^ ^^ 03 00 ^ 

00 OO 00 



gg Oe r^. CD 



a 


^» 


5J) 








oc 


t/5 


II 


II 


m 


0\ 


ti 


.5" 


t3 
P 




(11 


!>: 


^ 


S 


II 


a 


* 


CM 


h. 


•a 





-sr 




Q 


a 


O 


3 


sr 





o 








O 


lo 



■5 s 



c 


a 




B 


3 














tuj 


■T3 


C 







a 


C 





^, 


w 


•a 


II 




w 


n, 


(Tl 




3 


x> 




hi 


«5 


■a 


ou 




-c 


II 


8 


^ 


* 


c 






c 


•«: 


cw 


S 




i^ 








II 


■3 


a 

3 


< 












on 





fi 


!0 


s- 


PS 




ri 


u 






CHI 




\^ 


n) 


>t 







tM 


■x:i 




^ 


Ci. 


C 


II 








VO 


ft; 




^ 




II 


^ 


v, 


s: 




? 


V 


<3 








K, 


a 




sr 


tM 


?= 





51 


iX 


CO 


3 


II 





■* 


t3U 



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. 

Literature Cited 

ATWOOD, W.H. 1916. The visceral anatomy of 
the black snake {Zamenis constrictor). Washington 
Univ. Stud. Ser. Sci. 4(13): 3-38. 

1918. Visceral anatomy of the garter 

snake. Trans. Wisconsin Acad. Sci. Arts and Let- 
ters. 19: 531-552. 

BEDDARD, F.E. 1908. A comparison of the neo- 
tropical species of Corallus, C. cookii, with C. 
madagascariensis; and on some points in the ana- 
tomy of Corallus caninus. Proc. Zool. Soc. 
London. 1908: 135-158. 

1909. Some notes upon Boa occiden talis 

and Boa {Pelophilus) madagascariensis. Proc. 
Zool. Soc. London. 1909: 918-927. 

Bergman, R.A.M. 1941. Acrochordus Javanl- 
cus Hornst. Treubia, Buitenzorg. 18: 207-211. 

1950. The anatomy of Natrix vittata 

(L.). Zool. Meded. 31 : 13-24. 

1951. The anatomy of Homalopsis buc- 



cata. Proc. Kon. Nederlandse Akad. van Weten., 
Ser. C. 54: 511-524. 
1952. L'anatomie du genre Ptyas a Java. 



Rivista di Biologia Coloniale. 12: 5-42. 
1953. The anatomy of Cylindrophis 



rufus (Laur.). II. Proc. Kon. Nederlandse Akad. 
van Weten., Ser. C. 56: 657-666. 
. 1954. Thalassophis anomalus Schmidt. 



Universidad Nacional de Cordoba, Rep. Argen- 
tina. 1954: 1-16. 
1955a. The anatomy of Xenopeltis uni- 



color. Zool. Meded. 33: 209-225. 
. 1955b. Dendrophis pictus. Proc. Kon. 



Nederlandse Akad. van Weten., Ser. C. 58: 
206-218. 
. 1955c. L'anatomie de Cerberus rhyn- 



chops. Archives neerlandaises de Zool. 11: 

113-126. 
1955d. L'anatomie de Enhydrino schis- 



tosa D. Archives neerlandaises de Zool. 11: 
127-142. 



'Standard museum acronyms (Duellman, Fritts, 
and Leviton, 1978) are used throughout this paper. 



162 



Tulane Studies in Zoology and Botany 



Vol. 23 



. 1955e. L'anatomie de Enhydris enhy- 

dris. Revista di Biologia Coloniale. 15: 5-28. 
1956a. The anatomy of Dryophis pra- 



sinus. II. Proc. Kon. Nederlandse Akad. van 
Weten., Ser. C. 59: 272-279. 
1956b. The anatomy of Matrix submi- 



niata. Biol. Jaarb. 23: 306-326. 
. 1956-58. L'anatomie de £'/ff/7o/rfe5 /w5- 



cus B. Rivista di Biologia Coloniale. 16: 9-32. 
1958a. The anatomy of the Acrochor- 



dinae. I, II, IV. Proc. Kon. Nederlandse Akad. 
van Weten., Ser. C. 61: 145-166, 173-184. 
. 1958b. The anatomy of Matrix piscator. 



Biol. Jaarb. 26: 77-99. 
. 1959a. Matrix chrysarga. Biol. Jaarb. 27: 



73-86. 



27: 87-97. 



.. 1959b. Matrix trianguligera. Biol. Jaarb. 



. 1960. The anatomy of some Homalop- 

sinae. Biol. Jaarb. 28: 119-139. 
. 1961a. The anatomy of Coluber radiatus 



and Coluber melanurus. Pacific Sci. 15: 144-154. 
. 1961b. The anatomy of some Viperidae. 



(I, II, and III). Acta Morph. Neer. Scan. 4: 
195-230. 
. 1962a. The anatomy of Hydrophis fas- 



ciatus atriceps. Biol. Jaarb. 30: 389-416. 
1962b. Die anatomic der Ela- 



pinae. Zeitschrift fiir Wissenschaftliche Zoologie. 
167(3/4): 291-337. 
1963. The anatomy of Ablabes balio- 



deira, a colubrid snake from Java. J. Ohio Herp. 
Soc, 4: 1-14. 

.. 1965. The anatomy of Calamaria multi- 



punclata (Boie). Bull. Nat. Mus. Singapore. No. 

33, Pt. 7: 35-56. 
BOGERT, CM. 1968. A new genus and species of 

dwarf boa from southern Mexico. Amer. Mus. 

Novit. (2354): 1-38. 
BRAGDON, D.E. 1953. a contribution to the sur- 
gical anatomy of the water snake. Matrix sipedon 

sipedon; the location of the visceral endocrine 

organs with reference to ventral scutellation. Anat. 

Rec. 117: 145-161. 
BRONGERSMA. L.D. 1951. Some notes upon the 

anatomy of Tropidophis and Trachyboa (Ser- 

pentes). Zool. Meded. 31: 107-124. 
1957a. Notes on the trachea, the lungs 

and the pulmonary artery in snakes. I-II. Proc. 

Kon. Nederlandse Akad. van Weten., Ser. C. 60: 

299-313. 
1957b. Notes upon the trachea, the lungs 



and the pulmonary artery in snakes. III. Proc. 
Kon. Nederlandse Akad van Weten., Ser. C. 60: 
451-457. 



CAMAZINE, B., W. GARSTKA, and D. 
Crews. 1981. Techniques for gonadectomizing 
snakes (Thamnophis). Copeia 1981: 884-886. 

Collins, R.F., and C. Carpenter. 1970. 

Organ position-ventral scute relationship in the 
water moccasin (Agkistrodon piscivorus leucos- 
toma), with notes on food habits and distribution. 
Proc. Oklahoma Acad. Sci. 49: 15-18. 

Con ANT, R. 1961. A new water snake from Mex- 
ico, with notes on anal plates and apical pits in 
Matrix and Thamnophis. Amer. Mus. Novit. 
(2060): 1-22. 

DUELLMAN, W.E., T. FRITTS, and A.E. 
LEVITON. 1978. Museum acronyms. Herpetolo- 
gical Review. 9: 5-9. 

FRENKEL, G., and E. KOCHVA. 1970. Visceral 
Ematomy of Vipera palaestinae: an illustrated pre- 
sentation. Israel J. Zool. 19: 145-163. 

GARRIGUES, N.W. 1962. Placement of internal 
organs in snakes in relation to ventral scalation. 
Trans. Kansas Acad. Sci. 65: 297-300. 

Hull, C.H., and N.H. NIE, eds. 1979. SPSS 
update: new procedures and facilities for releases 7 
and 8. McGraw-Hill Book Company, New York, 
238 p. 

KOFRON, C.P. 1978. Foods and habitats of aqua- 
tic snakes (Reptilia, Serpentes) in a Louisiana 
swamp. J. Herpetol. 12: 543-554. 

Manna, D.C, and A.K. Sircar. 1978. Mor- 
phology of the hemipenis and cloacal gland and 
seasonal changes in the testes of the snake 
Ahaetulla nasuta. British J. Herpetol. 5: 711-717. 

Matthews, L.H., and F.H.A. Mar- 
shall. 1956. Cyclical changes in the reproduc- 
tive organs of the lower vertebrates. In A.S. 
Parkes, ed., Marshall's physiology of reproduc- 
tion, Ch. 3, 1: 156-225 Longmans, Green and 
Co., London. 

McDowell, S.B. 1979. a catalogue of the 
snakes of New Guinea and the Solomons with 
special reference to those in the Bernice P. Bishop 
Museum. Part III. Boinae and Acrochordoidea 
(ReptiHa, Serpentes). J. Herpetol. 13: 1-92. 

MUSHINSKY, H.R., and J.J. HEBRARD. 
1977. Food partitioning by five species of water 
snakes in Louisiana. Herpetologica. 33: 162-166. 

NiE, N.H., C.H. Hull, J.G. Jenkins, K. 

STEINBRENNER, and D.H. BENT. 1975. 
SPSS: Statistical package for the social sciences, 
2nd ed. McGraw-Hill Book Company, New York, 
675 p. 

Prasad, M.R.N. , and P.R.K. Reddy. 1972. 

Physiology of the sexual segment of the kidney in 
reptiles. General and Comparative Endocrinology, 
Supplement, (3): 649-662. 



No. 2 



Visceral Topography of Snakes 



163 



RASMUSSEN, J.B. 1979. An intergeneric analysis 
of some Boigine snakes — Bogert's (1940) Groups 
XIII and XIV (Boiginae, Serpentes). Vidensk. 
Meddr dansk naturth. Foren. 141: 97-155. 

ROSSMAN, D. A. 1963a. The colubrid snake genus 
Thamnophis: a revision of the Sauritus group. 
Bull. Florida State Mus. 7: 99-178. 

. 1963b. Relationships and taxonomic sta- 
tus of the North American natricine snake genera 
Liodytes, Regina, and Clonophis. Occ. Papers 
Mus. Zool., Louisiana St. Univ., (29): 1-29. 

RUNYON, R.P., and A. HABER. 1968. Funda- 
mentals of behavioral statistics. Addison-Wesley 
Publ. Co., Reading, Massachusetts, xi + 304 p. 

RUTHVEN, A.G. 1908. Variations and genetic re- 
lationships of the garter-snakes. Bull. U.S. Natl. 
Mus. 61: 1-201. 

Simpson, G.G., A. Roe, and R.C. Lewon- 

TIN. 1960. Quantitative zoology. Harcourt, Brace 
and Company, New York, BurUngame, 440 p. 

Smith, H., and J.E. HUHEEY. i960. The 
watersnake genus Regina. Trans. Kansas Acad. 
Sci. 63: 156-164. 

Thompson, J.C. 1913a. contributions to the 
anatomy of the Ophidia. Proc. Zool. Soc. 
London. 1913: 414-425. 

. 1913b. The variation exhibited by main- 
land and island specimens of the Hibakari snake. 
Matrix vibakari (Boie). Proc. U.S. Natl. Mus. 46: 
157-160. 

. 1914. Further contributions to the ana- 



tomy of the Ophidia. Proc. Zool. Soc. London. 
1914: 379-402. 

Thorpe, R.S. 1975. Quantitative handling of 
characters useful in snake systematics with particu- 
lar reference to intra-specific variation in the 
Ringed Snake, Natrix natrix (L.). Biol. J. Linn. 
Soc. 7: 27-43. 

Underwood, G. 1976. a systematic analysis of 
bold snakes. In A. d'A. Bellairs and C. Barry Cox, 
eds.. Morphology and biology of reptiles. Linnean 
Soc. Symposium Ser. No. 3: 151-175. 

VALLE, J.R. 1944-45. Sobrevida da parelheira 
{Philodryas sp.) depois da adrenalectomia. Mem. 
Inst. Butantan, Sao Paulo. 18:237-240. 

VARKEY, a. 1979. Comparative cranial myology 
of North American natricine snakes. Milwaukee 
Public Mus. Publ. Biol, and Geol. (4): 1-76. 

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