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
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48
Tulune Sludles in Zoology und Botany
Vol. 23
Range. Primarily distributed on the
Mexican Plateau, Kinosternon hirdpes
ranges from Alamito Creek in Texas in
the United States and the Rfos Santa
Maria, Carmen, and Conchos in northern
Mexico south and eastward to the Chap-
ala, Zapotlan, San Juanico, Pa'tzcuaro,
and Valle de Me'xico basins of the Sierra
Volcanica Transversal of southern Mex-
ico. It is known from between at least 800
and 2600 m in elevation.
Specimens examined and Additional
Records. See locality list.
Etymology. The specific name hirtlpes
is from the Latin, hirtus, meaning rough,
and pes meaning foot, and refers to the
rough scales on the feet of the species.
Kinosternon hirtlpes hirtlpes Wagler
Valley of Mexico Mud Turtle
CInosternon hirtlpes Wagler, 1830:187
(see species synonymy).
Ozotheca odorata Duges, 1869:143 (in
part; State of Mexico).
CInosternon pennsylvanlcum Duges,
1888:107 (in part; Valley of Mexico).
CInosternum /?//7//7e5 Boulenger, 1889:38;
Siebenrock, 1906:94, 1907:551 (State of
Mexico); Gadow, 1908:5 (Chalco lakes,
Valley of Mexico).
CInosternon pensvlvanlcum Herrera,
1890:330, 1891:46 (in part; Valley of
Mexico).
Cynostenum pennsylvanlcum Herrera,
1893:342 (in part; Valley of Mexico).
Cynosternon pensylvanlciim Herrera
and Lope, 1899:131 (in part; Valley of
Mexico).
CInosternom pennsllvanlcum Cope, 1900:
1229 (in part; Valley of Mexico).
CInosternon sp. Herrera, 1904:6 (Mexi-
calzingo. Valley of Mexico).
CInosternum pennsylvanlcum Gadow,
1905:209 (in part; Valley of Mexico).
Kinosternon hirtlpes Martin del Campo,
1938:391 (Valley of Mexico); Caballero
y C, 1939:279 (Xochimilco, Mexico,
Distrito Federal); H.M. Smith and
Taylor, 1950a:25 (Distrito Federal);
Glass and Hartweg, 1951:50 (Valley of
Mexico); Schmidt, 1953:89; Beltz,
1954:124 (Mexico City, Mexico);
Martin del Campo, 1955:66 (Valley of
Mexico); Deevey, 1957:240 (Valley of
Mexico); Casas Andreu, 1965:385 (Dis-
trito Federal); Kranz, Smith, and
Smith, 1971:23 (near City of Mexico);
Greene, 1972:2 (in part; Mexico City,
Puebla locality based on A'. Integrum);
Perez Villegas and Reyna Trujillo,
1978:215 (southern region of Valley of
Mexico).
Kinosternon hirtlpes hirtlpes Mertens and
Wermuth, 1955:336 (first use of this
combination; in part; State of Mexico);
Wermuth and Mertens, 1961:20 (in
part; State of Mexico); Duellman,
1961:57, 1965:653 (in part; Michoacan
localities not this subspecies); H.M.
Smith, Williams and Moll, 1963:209;
Liner, 1964:221 (in part; Guanajuato
records not this subspecies); Pritchard,
1967:37 (in part; State of Mexico);
Casas Andreu, 1967:44 (in part; State
of Mexico); Parsons, 1968:1238; Legler
and Webb, 1970:163 (in part; Chihua-
hua records based on A", h. murrayl);
Mittermeier, 1971:16 (Mexico City);
Moll and Legler, 1971:92 (in part; Chi-
huahua records based on K. h.
murrayl); Wermuth and Mertens, 1977:
7; Pritchard, 1979:537 (in part; Mexico
City).
Kynosternon hirtlpes Lopez 1975:2 (Val-
ley of Mexico.
Kinosternon sp. Niederberger, 1979:134
(Valley of Mexico archeological re-
mains: 5500 BC).
Types. Only the holotype (Fig. 25) is
available (see Remarks below).
Diagnosis. A subspecies oi Kinosternon
hirtlpes with 1) the adult nasal scale tri-
angular, rhomboidal, or bell-shaped (fur-
cate behind in subadults, but not in large
adults); 2) a mottled head pattern, typical-
ly organized into a light streak extending
posteriorly from the angle of the jaw,
with a similar light postorbital streak vari-
ably evident; 3) one or (typically) two
pairs of mental chin barbels, the anterior
pair largest; 4) medium body size (maxi-
No. 1
Kinosternon Biosystematics
49
mum male size 140 mm CL; female, 140
mm); 5) relatively short bridge length
(male BL/CL, 17.6<7o; female
21.7'^o); 6) relatively short interfemoral
seam length (male IF/CL, 6.9%;
female , 7.1%); 7) relatively long inter-
anal seam length (male lAN/CL,
20.6%; female ,25.8%) and 8) popula-
tions confined to the Valley of Mexico.
Remarks. As mentioned under SYN-
THESIS (above), the allocation of the
holotype of Kinosternon hirtipes to the
Valley of Mexico must remain uncertain
until additional material is available from
the southern and southwestern margins of
the Mexican Plateau.
Specific natural history data are un-
available for Kinosternon hirtipes hir-
tipes. A photograph of the plastron of
UMMZ 99449, an adult female, appears
in H.M. Smith and R. B. Smith (1980;
plate 19, bottom).
Range. Kinosternon hirtipes hirtipes is
known only from the drainages of the
Valley of Mexico.
Specimens Examined and Additional
Records. See locality lists.
Etymology. See species account.
Moll and Legler, 1971:92 (in part; Chi-
huahua records based on k. h.
murrayi); Wermuth and Mertens, 1977:
7; Pritchard, 1979:537 (in part; Mexico
City).
Kynosternon hirtipes Lopez 1975:2 (Val
ley of Mexico.
Kinosternon sp. Niederberger, 1979:134
(Valley of Mexico archeological re-
mains: 5500 BC).
Types. Only the holotypes (Fig. 27) is
available (see Remarks below).
Diagnosis. A subspecies of Kinosternon
hirtipes with 1) the adult nasal scale tri-
angular, rhomboidal, or bell-shaped (fur-
cate behind in subadults, but not in large
adults); 2) a mottled head pattern, typical-
ly organized into a light streak extending
posteriorly from the angle of the jaw,
with a similar light postorbital streak vari-
ably evident; 3) one or (typically) two
pairs of mental chin barbels, the anterior
pair largest; 4) medium body size (maxi-
mum male size 140 mm CL; female, 140
mm); 5) relatively short bridge length
(male x BL/CL, 17.6%; female x ,
21.7%); 7) relatively short interfemoral
seam length (male x IF/CL, 6.9%;
female x , 7.1%); 8) relatively long inter-
anal seam length (male x lAN/CL,
20.6%; female x, 25.8%) and 9) popula-
tions confined to the Valley of Mexico.
Remarks. As mentioned under SYN-
THESIS (above), the allocation of the
holotype of Kinosternon hirtipes to the
Valley of Mexico must remain uncertain
until additional material is available from
the southern and southwestern margins of
the Mexican Plateau.
Specific natural history data are un-
available for Kinosternon hirtipes hir-
tipes. A photograph of the plastron of
UMMZ 99449, an adult female, appears
in H.M. Smith and R. B. Smith (1980;
plate 19, bottom).
Range. Kinosternon hirtipes hirtipes is
known only from the drainages of the
Valley of Mexico.
Specimens examined and Additional
Records. See locality lists.
Etymology. See species account.
Kinosternon hirtipes murrayi
Glass and Hartweg
Murray's Mud Turtle
Ozotheca odorata Duges, 1869:143 (in
part; Guanajuato); Velasco 1890b: 54 (in
part; Guanajuato).
Cinosternon hirtipes Westphal-Castelnau,
1872:278 (Guanajuato).
Cinosternum hirtipes Cope, 1887:23 (in
part; city of Chihuahua, Guanajuato).
Cinosternum pennsylvanicum Duges,
1896c:479 (Guanajuato).
Cinosternon pennsilvanicum Cope, 1900:
1229 (Toluca Valley northward through
Guanajuato.
Kinosternon sonoriense Dunn, 1936:472
(in part; Rio Conchos, Julimes, Chi-
huahua); H.M. Smith and Taylor
1950a:26 (in part; Chihuahua and Dur-
ango); Mertens and Wermuth, 1955:
338 (in part; Chihuahua and Durango);
50
Tulane Studies in Zoology and Botany
Vol. 23
Casas Andreu, 1965:386 (in part; Chi-
huahua and Durango); Legler and
Webb, 1970:163 (in part; Rfos Papi-
gochic and Tomuchic in western Chi-
huahua); Iverson, 1976:1 (in part;
Upper Rfo Yaqui, Chihuahua; see Iver-
son, 1978).
Kinosternon hirtipes Caballero y C,
1940b: 562 (Rio Lerdo, Guanajuato);
Caballero y C. y Cerecero, 1943:534
(Rio Lerdo del Valle de Santiago,
Guanajuato); H.M. Smith and Taylor,
1950b:25 (in part; Chihuahua, Guana-
juato); Williams, Smith, and Chrapliwy,
1960:36 (Chihuahua, 1 mi. E La Cruz);
Casas Andreu, 1965:385 (in part; Chi-
huahua, Guanajuato); Conant, 1978:
465 (Texas, Chihuahua, Durango and
Zacatecas).
Sternothenis odoratus Brown, 1950:230
(in part; Presidio Co., Texas; based on
holotype oiK. murrayi; see Conant and
Berry, 1978:15).
Kinosternon murrayi Glass and Hartweg,
1951:50 (Type-locality, "Harper
Ranch, 37 miles south of Marfa, Pres-
idio County, Texas." Holotype,
TCWC 650, a young male, collected 12
August 1941 by S.H. Wheeler.); Peters,
1952:54 (Texas); Legler, 1960:139 (Jet.
Rio San Pedro and Conchos, and Ojin-
aga. Chihuahua).
Kinosternon hirtipes murrayi Schmidt,
1953:89 (first use of combination;
Texas); Mertens and Wermuth, 1955:
336 (Texas); H.M. Smith, Williams and
Moll, 1963:207 (Chihuahua); Casas
Andreu, 1967:45 (Texas, Chihuahua,
and Durango); Parsons, 1968:1238;
Cochran and Goin, 1970:135 (Texas);
Moll and Legler, 1971:92 (Durango and
Chihuahua); Ernst and Barbour, 1972:
66 (Texas); Hambrick, 1976:292
(Texas); Wermuth and Mertens, 1977:7
(Texas); Conant and Berry, 1978:1
(Texas and Chihuahua); Iverson, 1978:
476 (Chihuahua).
Kinosternon hirtipes hirtipes Mertens and
Wermuth, 1955:336 (in part; Chihua-
hua); Duellman, 1961:57 (in part ?;
Michoacan, 8 km W Ciudad Hidalgo
and Lago'de Cuitzeo); Casas Andreu,
1967:44 (in part; Chihuahua, Michoa-
can, and Guanajuato); Legler and
Webb, 1970:163 (in part; Rios Papi-
gochic and Tomuchic, western Chihua-
hua); Moll and Legler, 1971:92 (in part;
Chihuahua); Wermuth and Mertens,
1977:7 (middle and western Mexico);
Pritchard 1979:537 (in part).
Kinosternon flavescens Stebbins, 1966:82
(in part; Durango; see Iverson, 1978).
Kinosternon hirtipes murryi Ashton et al.,
1976:51 (lapsus pro murrayi).
Kinosternon hertipes Semmler, et al.,
1977:18 (near Galeana, Chihuahua).
Types. Holotype: see subspecies synon-
ymy. Paratypes: USNM 15860, adult
male, preserved whole, from "Marfa,
Presidio County, Texas", collected by
Vernon Bailey: UMMZ 101294, adult
male, preserved whole, and UMMZ S-
1083, shell of adult male, both topotypic
and collected 12 June 1950 by Herndon
G. Dowling.
Diagnosis. A subspecies oi Kinosternon
hirtipes with: 1) a large posteriorly furcate
nasal scale (typically exending posterior to
the orbits); 2) an extremely variable mot-
tled to reticulated head pattern; 3) typical-
ly two pair of mental chin barbels, the
anterior pair largest; 4) medium to large
body size (maximum known male size,
182 mm CL; female, 157 mm);^) relative-
ly long bridge length (male x BL/CL,
20.0<^o; female x, 23. 7%); 6) relatively
long gular length (male x GL/CL,
14.7<^o; female x, 15.8%); and 7) popu-
lations confined to the Big Bend region of
Texas and adjacent Chihuahua southward
across the Mexican Plateau to northern
Jalisco, northern Michoacan, and eastern
Mexico (state).
Remarks. As discussed in the results,
there appears to be a slight morphometric
distinction between populations of K. h.
murrayi in the Ri'o Nazas northward, and
populations in the Rio Aguanaval south-
ward. This difference is not considered
significant enough to warrant subspecific
designation, but has some interesting zoo-
geographic impUcations.
No. I
Kinosternon Biosystematics
51
Range. Kinosternon hirtipes murrayi is
known from the following basins in
Aguascalientes , Chihuahua, Coahuila,
Durango, Guanajuato, Jalisco, Mexico,
Michoacan, San Luis Potosi, Texas, and
Zacatecas: Santa Maria (Chihuahua),
Carmen, El Sauz, Conchos, Bustillos,
Papigochic, Nazas, Viesca, Aguanaval,
Santiaguillo, Mezquital, El Salto, Santa
Maria (San Luis Potosi; presumably in-
troduced), Aguascalientes, Verde, Lerma
(except Chapala), Cuitzeo, Balsas, and
Villa Victoria (with reservation).
Specimens examined and Additional
Records. See locality lists.
Etymology. The subspecific name mur-
rayi is a patronym, honoring Dr. Leo T.
Murray of Texas A & M College.
Kinosternon hirtipes chapalaense ssp. nov.
Lake Chapala Mud turtle
Cinosternum integrum Gadow, 1908:518
(in part; Laguna de Zapotlan, Jalisco).
Kinosternon hirtipes Altini, 1942:153 (in
part; Lake Chapala, Jalisco).
Kinosternon hirtipes hirtipes Duellman,
1961:57, 1965:653 (in part; Jiquilpan,
La Palma, Lago de Came'cuaro = 14
km E Zamora, Michoacan).
Kinosternon hirtipes chapalaense Pritch-
ard, 1979:557 (nomen nudum; Lake
Chapala).
Holotype. UMMZ 97128, adult male,
preserved whole, from Lake Chapala,
0.25 mile off Chapala, Jalisco, Mexico
[20°18'N, 103°12'W]; collected 15 July
1947, by Norman Hartweg.
Paratypes. All preserved whole: UMMZ
97122-23, topotypic adult females; UMMZ
97124, topotypic subadult male; 97125-27
and 97129-30, topotypic adult males; and
UU 12126-12128, adult male, subadult fe-
male, and juvenile, Lago de Chapala, 3.2
km W Chapala; and UU 12125, adult fe-
male, Lago de Chapala, 6.1 km W Ajijic,
all collected on 21-22 June 1969 by Clyde
Barbour.
Diagnosis. A subspecies of Kinosternon
hirtipes with 1) a reduced crescent-shaped
nasal shield, which nearly always lies an-
terior to the orbits (Figure 23); 2) a reduc-
tion of dark pigment on the head and
neck, dark markings confined to isolated
spots or reticulations dorsally (Figures 23
and 24), but laterally sometimes organ-
ized as two dark, nearly parallel post-
orbital stripes; 3) the neck and chin virtu-
ally unmarked and the mandibular and
maxillary sheaths bearing only a few dark
streaks, if any; 4) one, two, or three pairs
of mental barbels present, the anterior-
most pair (near the mandibular symphysis)
usually the largest; 5) medium body size
(maximum known size for males is 152
mm CL; females, 149 mm); 6) relatively
long bridge length (male x BL/CL,
20.3<^o; female x, 25.3%); 7) relatively
long interanal _seam (male x lAN/CL,
19.1%; female x, 25.2%); and 8) popula-
tions confined to the Chapala and Zapot-
lan (and possibly Duero) basins of Jalisco
and Michoacan.
Remarks. Field notes accompanying
the topotypes provide no additional geo-
graphical and ecological information.
However, Clyde Barbour (pers. comm.)
obtained the non-topotypic paratypes
(during the night 21-22 June 1969) along
the shore of Lake Chapala, on trot-lines
baited with liver. These lines were neces-
sarily buoyed off the lake bottom with
floats to avoid bait removal by crabs.
Peter Meylan found a single rotten car-
cass of this species on the south shore of
Lake Chapala just east of Tuxcueca dur-
ing my field trip to the area on 15 June
1978; trapping at that locality produced
no turtles. Trapping in isolated spring-fed
pools just northeast of the town of Chap-
ala on 9 May 1981 produced only K. inte-
grum.
Range. Kinosternon hirtipes chapalense
is known only from the Lago de Chapala
and Laguna de Zapotlan drainage basins
in Jalisco and Michoacan, Mexico. Speci-
mens from the Rio Duero basin are tenta-
tively considered intergrades with K. h.
murrayi.
Specimens Examined and Additional
Records. See locality list.
Etymology. The subspecific name chap-
52
Tulane Studies in Zoology and Botany
Vol. 23
alaense refers to Lake Chapala wherein
the type series was collected.
tKinosternon hirtipes megacephalum
ssp. nov.
Viesca Mud Turtle
Holotype. SM(BCB) 1 1466, adult male,
preserved whole, from 3.2 km SE Viesca
[25 °2rN, 102 °48'W], Coahuila; collected
4 June 1961 by Bryce C. Brown and John
Wottring by seining a drying pond.
Paratypes. SM(BCB) 11460-65, adult
females, preserved whole, all topotypic
(11461 photographed in H.M. Smith and
R.B. Smith, 1980); and SM(BCB) 9823,
adult male, preserved whole, from 9.7 km
SW Viesca, also collected on 4 June 1961
by Brown and Wottring (see Figs. 22 and
24).
Diagnosis. A subspecies of Kinosternon
hirtipes with: 1) enlarged head, hyper-
trophied head musculature, and broad al-
veolar jaw surfaces (Fig. 24); 2) the nasal
scale furcate posteriorly; 3) the head pat-
tern mottled or reticulated as in K. h.
murrayi; 4) three to four pairs of chin
barbels are present, two to three mental
pairs (anterior usually the largest) and one
small pair at level of anterior edge of
tympanum; 5) small body size (maximum
known size for males 99 mm CL; females,
1 17 mm); 6) plastron extremely reduced in
size (Fig. 22); 7) relatively short bridge
length (male x BL/CL, 17.3^70; female x ,
23.90/0); 8) relatively short gular length
(male x GL/CL, ll.Oo/o; female x,
12.8°7o); 9) relatively short interanal seam
length (male x lAN/CL, 15.9"7o; female
X , 20.90/0); and 10) populations confined
to southwestern Coahuila.
Remarks. This subspecies is known
only from the type series. Field work in
the area of the type locality (see MATER-
IALS AND METHODS) suggests that K.
h. megacephalum is now extinct; natural
permanent water habitats apparently no
longer exist near Viesca. Future field
work should be concentrated in the
mountains south of the city of Viesca in
hope of discovering permanent water situ-
ations where turtles (and fishes?) might
still exist.
The distinctive trophic apparatus of
this subspecies is likely an adaptation to
stenophagous molluscivory.
Range. Known only from the two local-
ities in Coahuila at which the type series
was collected.
Etymology. The subspecific name
megacephalum is from the Greek mega,
meaning large, and kephale, meaning
head, and refers to the enlarged head, di-
agnostic of the subspecies.
Kinosternon hirtipes tarascense ssp. nov.
Patzcuaro Mud Turtle
Kinosternon hirtipes Altini, 1942:153 (in
part; Lake Patzcuaro, Michoacan).
Kinosternon hirtipes hirtipes Duellman,
1961:57, 1965:653 (in part; Lago de
Patzcuaro, Michoacan); Casas Andreu,
1967:45 (in part; Patzcuaro, Canal de la
Tzipecua, Michoacan).
Holotype. UF 43506, adult male, pre-
served whole, from Lago de Patzcuaro,
adjacent to city of Patzcuaro [19°32'N,
101°36'W]; purchased in Patzcuaro
market 13 June 1978 by John B. Iverson.
Paratypes. All topotypic and preserved
whole: UF 43505 and 43596, adult
females; and UF 43507 and 43595, adult
males.
Diagnosis. A subspecies of K. hirtipes
with: 1) a typically finely mottled to
spotted head (Fig. 24); 2) variable red-
brown to brown staining on the otherwise
light yellow plastral scutes [The dark plas-
tral scutes are apparently a result of
natural staining; the character is exhibited
to variable degrees by individuals and the
dark color is lost when plastral scutes are
shed (Fig. 22)]; 3) the large nasal scale
posteriorly furcate; 4) two pairs of mental
chin barbels typically present; 5) small to
medium body size (maximum known size
for males 136 mm CL; females, 132 mm)^:
6) relatively short bridge length (male x
BL/CL, 18.0^^0; female x, 21. 40/0); 7) rel-
No. 1
Kinosternon Biosystematics
53
atively short gular length (male x
GL/CL, 10.6%; female x , 12.6%); 8) rel-
atively long interpectoral seam length
(malex IP/CL, 10. lo/o; female x, 8.5%);
and, 9) populations confined to the Lago
de Patzcuaro drainage basin.
Remarks. Despite considerable study of
other components of the biota of the
Lago de Patzcuaro (see review in Cole,
1963 and Barbour, 1973), the mud turtles
have been ignored. Reproductive infor-
mation resulting from my studies appears
in Table 2.
Range. Known only from the basin of
the Lago de Patzcuaro, Michoacan.
Specimens examined and Additional
Records. See locality list.
Etymology. The subspecies name tara-
scense honors the native tribe of Indians,
the Tarascas, inhabiting the Patzcuaro
area.
Kinosternon hirtipes magdalense ssp. nov.
San Juanico Mud Turtle
Holotype. UF 45035, an adult male,
preserved whole, from along the face of
the dam at Presa San Juanico, Michoacan
[ca. 19°50'N, 102°40'W] (Fig. 28). Holo-
type collected 15 June 1978 by John B.
Iverson, Peter A. Meylan, and Ron
Magill.
Paratypes. UF 45036, a subadult fe-
male, UF 45038, female shell; UF 45039-
40, male shells, all topotypic; and TUL
18677, aduU male, collected atop Presa
San Juanico 9 August 1963 by Clyde D.
Barbour and Salvador Contreras-Balderas.
Diagnosis. A subspecies o{ Kinosternon
hirtipes with: 1) a finely mottled to spot-
ted head pattern with jaw streaking mini-
mal or absent; 2) a large nasal scale, fur-
cate behind; 3) two pairs of mental chin
barbels present; 4) small body size (max-
imum known male size 94 mm CL; fe-
male, 91 mm); 5) a relatively small plas-
tron (male x PWB/CL, 41.9%; female
X, 43.5%); 6) relatively short bridge
length (male x BL/CL, 18.5%; female x ,
19.7%); 7) relatively short gular length
(male x GL/CL, 9.9%; female x,
11.0%); 8) relatively long interpectoral
seam length (male x IP/CL, 8.7%; fe-
male X, 11.0%); and, 9) populations re-
stricted to the Magdalena Valley, Micho-
acan.
Remarks. As mentioned earlier (MAT-
ERIALS AND METHODS), the turtles
inhabiting the Presa San Juanico are
poorly known. Future field work in the
area should help delimit the subspecific
range within the Magdalena Valley and
also provide basic natural history infor-
mation.
Range. Kinosternon hirtipes magdal-
ense is known only from the type series,
all from the reservoir above Presa San
Juanico in the Magdalena Valley of
Michoacan, Mexico.
Etymology. The subspecific name mag-
dalense refers to the Magdalena Valley of
Michoacan to which the subspecies is
apparently endemic.
Figure 28. Holotype (UF 45035) of Kinosternon
hirtipes magdalense.
54
Tulane Studies in Zoology and Botany
Vol. 23
Evolution
Based on the derived characters of the
turtles of the Kinosternon hirtipes species
group (Table 3), I have constructed a phy-
logeny of the included taxa (Fig. 29). For
reasons discussed by Farris (1966) and
Kluge and Farris (1969) (e.g., high intra-
familial variation and indiscrete character
shifts), I have not always assumed that
taxa sharing derived morphometric char-
acters are closely related. In fact, the dis-
tribution of some character states among
taxa clearly indicates that those characters
are not a result of single origin, but rather
of convergence. For example, the length-
ening of the interanal seam in K. sonor-
iense (longifemorale) and K. hirtipes (hir-
tipes and chapalaense) certainly illustrates
multiple origin of a derived character
state. In addition, Viesca (megacephal-
um), Patzcuaro (tarascense), and San
Juanico turtles (magdalense) all share a
relatively short bridge (with Valley of
Mexico turtles), a short gular, and small
body size (the latter two also share a long
interpectoral seam), yet geographically
and zoogeographically (Iverson, in prep-
aration) the three populations likely do
not represent a monophyletic divergence
from a pre-murrayi stock. Rather, the
evolution of these character states is more
likely a response to selection in similar,
very narrow adaptive zones (i.e., isolated,
very small basins). Unfortunately, the
functional significance of those characters
is unknown, as is that of most of the other
characters herein examined (but see Iver-
son, MS 2)
Table 3. Tally of subspecific taxa exhibiting derived character states in the Kinosternon
hirtipes species group. Primitive states are discussed in the text.
Derived Character
1
small plastron
2
short bridge
3
short gular
4
long gular
5
long interpectoral
6
short interpectoral
7
short interfemoral
8
long interanal
9
short interanal
10
V-shaped nasal
11
reduced nasal
12
multiple, long chin
barbels
13
reduced head
pigment
14
well-developed
head stripes
15
large head
16
female > male
17
small body size
18
tendency toward
unicarination
Taxa exhibiting derived character
magdalense, megacephalum
magdalense, tarascense, hirtipes, megacephalum
magdalense, tarascense, megacephalum
sonoriense and longifemorale
magdalense, tarascense
sonoriense, longifemorale
hirtipes
chapalaense, hirtipes, longifemorale
megacephalum
murrayi, magdalense, tarascense, megacephalum,
chapalaense
chapalaense
sonoriense, longifemorale
chapalaense
hirtipes
megacephalum
sonoriense, longifemorale (possibly tarascense,
megacephalum)
magdalense, tarascense, megacephalum
murrayi, hirtipes, magdalense, tarascense,
megacephalum, chapalaense
No. I
Kinosternon Biosystematics
55
The Kinosternon hirtipes species group
apparently evolved on the Mexican Pla-
teau from an ancestor as yet unknown.
Despite the fact that several coastal
streams have come to drain the Plateau
due to headwater stream erosion (e.g.,
Rios Yaqui, Me^quital, Santiago, Balsas;
see Fig. 4), K. hirtipes has nowhere left
the Plateau. This is surprising since K.
integrum has apparently moved both up
and down several of these basins (Balsas,
Santiago-Lerma, and Mezquital; Iverson,
unpublished). K. sonoriense apparently
evolved from a K. hirtipes-\\ke ancestor
isolated in the Sonoran Desert, possibly
following migration across the well-
documented Sonora Desert-Chihuahua
Desert filter barrier in southeastern Ari-
zona, southwestern New Mexico and ad-
jacent Mexico (see review in Morafka,
1977). Because so much geological infor-
mation concerning the Mexican Plateau is
now available (see reviews in Barbour,
1973 and Wauer and Riskind, 1978), a
discussion of the historical zoogeography
of the K. hirtipes species group will
appear elsewhere (Iverson, in prepara-
tion).
The relationship between the Kino-
sternon hirtipes species group and other
Kinosternon is unclear. Siebenrock
p* „.''^/^ ..^' y ./- j^"- y
Figure 29. A theory of relationships among the
subspecific taxa of the Kinosternon hirtipes
species group. Numbers refer to derived character
states listed in Table 3. Solid lines cutting line-
ages mark identical shifts (convergence) in
character states.
(1907:551) included K. hirtipes and K.
sonoriense, K. baurii, K. subrubrum, K.
flavescens, and K. steindachneri (= K.
subrubrum) in the K. subrubrum species
group. However, I believe that K. baurii
and K. subrubrum (including steindach-
neri) represent a species group distinct
from the K. hirtipes group, and that K.
flavescens is similarly distinct. Perhaps
the closest relative of the hirtipes group is
K. herrerai (found in the Tampico Em-
bayment of eastern Mexico; i.e., non-
Plateau), which shares with most K. hir-
tipes the elevated scale patches on the
hindlegs of males, the tendency toward
unicarination in adults, the furcate nasal
scale, the reduced plastron, the broad
inguinal-axillary contact, and several
morphometric plastral characters. Un-
fortunately, the determination of the
phylogenetic relationship of the K. hir-
tipes group to the other species groups in
the genus must await further analysis.
Key To Adult Turtles Of The
Kinosternon hirtipes SPECIES GROUP
lA. Nasal shield triangular, rhomboidal,
or bell shaped; largest 2 pairs of chin
barbels relatively long (at least one
pair > half orbit diameter, with one
pairmental and other at mid-tympan-
um level); interpectoral length
averages 5.0% of plastron length in
males (less than 8% in 95% of cases)
and 4.0% in females (less than 7% in
96% of cases); posterior width of
plastral forelobe (PWB) averages
47.2% of carapace length in males
(more than 44% in 95% of cases) and
49.0% in females (more than 45% in
96% of cases); maximum gular width
averages 19.7% of carapace length in
males (more than 18% in 94% of
cases) and 19.1% in females (more
than 17% in 94% of cases); first
neural bone often (38.1%) in contact
with nuchal bone; northwestern
Chihuahua and Sonora, Mexico and
adjacent New Mexico, Arizona and
California Kino-
sternon sonoriense 2
56
Tulane Studies in Zoology and Botany
Vol. 23
IB. Nasal shield large and deeply notched
posteriorly (V-shaped), or reduced to
crescent-shaped scale lying fully
anterior to level of orbits, or
triangular, rhomboidal, or bell
shaped if from Valley of Mexico;
largest 2 pairs of chin barbels
relatively short (< half orbit
diameter), mentally located, with
anterior pair larger; interpectoral
length averages 8.2*^0 of plastron
length in males (more than 4.5% in
97<^o of cases); and 6.6% in females
(more than 3.5% in 94% of cases);
posterior width of plastral forelobe
(PWB) averages 42.8% of carapace
length in males (less than 48% in 98%
of cases) and 47.6% in females (less
than 51% in 95% of cases); maximum
gular width averages 17.3% of
carapace length in males (less than
20% in 98% of cases) and 17.0% in
females (less than 20% in 98% of
cases); first neural rarely (10.2%) in
contact with nuchal; Chihuahua,
Mexico and adjacent Texas south-
ward to Jalisco, Michoacan, and
Me'xico, Mexico (state)
Kinosternon hirtipes 3
2A. Interanal seam length averages
19.5% of carapace length in males
(more than 16.5% in 97% of cases)
and 23.0% in females (more than
21% in 90% of cases); interfemoral
seam length averages 10.1% of
carapace length in males (less than
13% in 93% of cases) and 10.1% in
females (less than 12.5% in 95% of
cases); maximum first vertebral width
averages 24.4% of carapace length in
males Oess than 28% in 97% of cases)
and 25.5% in females (less than 28%
in 90% of cases); and gular width
averages 20.0% in males (more than
18.5% in 93% of cases) and 19.4% in
females (more than 17.5% in 90% of
cases); Bill Williams, lower Colorado,
Gila, Sonora, Magdalena, Yaqui,
southwest New Mexico, and Casas
Grandes basins K.s. sonoriense
2B. Interanal seam length averages
14.4% of carapace length in males
(less than 16% in 90% of cases), and
18.5% in females (less than 22% in
100% of cases); interfemoral seam
length averages 12.8% of carapace
length in males (more than 10% in
100% of cases) and 13.5% in females
(more than 11.5% in 91% of cases);
maximum first vertebral width
averages 28.9% of carapace length in
males (more than 28% in 90% of
cases) and 28.8% in females (more
than 26% in 100% of cases); and
gular width averages 17.7% of
carapace length in males (less than
19% in 100% of cases) and 17.8% in
females (less than 20% in 100% of
cases); Rio Sonoyta basin, Arizona,
and Sonora, Mexico
K. s. longifemorale
3A. Nasal shield reduced to crescent-
shaped scale lying anterior to level of
orbits; dark reticulate head markings
reduced or nearly absent; plastral
width at humero-pectoral seam
(PWA) averages 33.3% of carapace
length in males (less than 35.5% in
100% of cases) and 37.0% in females
(less than 40% in 93% of cases);
bridge length averages 20.3% of
carapace length in males (over 18% in
100% of cases) and 25.3% in females
(more than 22% in 100% of cases);
gular length averages 11.8% of
carapace length in males (less than
13% in 93% of cases) and 14.5% in
females (less than 18.5% in 100% of
cases); forelobe length averages
30.5% of carapace length in males
(less than 33.5% in 100% of cases)
and 31.8% in females (less than 34%
in 100% of cases); interhumeral seam
length averages 14.0% of maximum
plastron length in males (more than
12% in 93% of cases) and 12.7% in
females (more than 10% in 88% of
cases); interabdominal seam length
averages 28.6% of maximum plastron
length in males (more than 26% in
No. I
Kinosternon Biosystematics
57
93*^0 of cases) and ZQ.S'Vo in females
(more than 25. 5 "^o in 100<Vo of cases);
Lake Chapala and Lake Zapotlan
basins, Michoacan and Jalisco,
Mexico
.... Kinosternon hirtipes chapalaense
3B. Nasal shield large and deeply notched
posteriorly, triangular, rhomboidal or
bell shaped; head with abundant dark
head markings; plastral width at
humero-pectoral seam (PWA) av-
erages 36.1% of carapace length in
males (more than 32.5% in 98% of
cases) and 38.1% in females (more
than 35% in 97% of cases); bridge
length averages 19.9% of carapace
length in males (less than 22% in 95%
of cases) and 23.4% in females (less
than 26% in 93% of cases); gular
length averages 14.6% of carapace
length in males (more than 12% in
95% of cases; excluding turtles from
Patzcuaro, San Juanico, Cuitzeo, and
Viesca basins) and 15.6% in females
(more than 12.5% in 97% of cases;
excluding turtles from Patzcuaro, San
Juanico, Cuitzeo, and Viesca basins);
forelobe length average 31.2% of
carapace length in males (more than
28.5% in 100% of cases) and 34.4%
in females (more than 30% in 98% of
cases); interhumeral seam length aver-
ages 11.8% of maximum plastron
length in males (less than 15% in 90%
of cases) and 12.6% in females (less
than 17% in 96% of cases); interab-
dominal seam length averages 28.1%
of maximum plastron length in males
(less than 31% in 96% of cases) and
28.0% in females (less than 31% in
95% of cases); Chihuahua, Mexico
and adjacent Texas southward to Ja-
lisco, Michoacan, and Mexico, Me'x-
ico, except Chapala and Zapotlan
basins 4
4A. Gular length averages 10.5% of cara-
pace length in males (less than 13% in
100% of cases) and 12.4% in females
(less than 14% in 93.0% of cases);
plastron width at humero-pectoral
seam (PWA) averages 34.6% of cara-
pace length in males (less than 38% in
100% of cases) and 35.9% in females
(less than 38% in 93% of cases); pos-
terior width of plastral forelobe
(PWB) averages 42.5% of carapace
length in males (less than 45.6% in
100% of cases) and 45.8% in females
(less than 47.5% in 88% of cases); an-
terior width of plastral hindlobe
(PWC) averages 39.3% of carapace
length in males (less than 43% in
100% of cases) and 42.8% in females
(less than 46% in 93% of cases); max-
imum carapace length 140 mm in
males, 135 mm in females; Patzcuaro,
San Juanico, and Viesca basins
populations with small plastron .... 5
4B. Gular length averages 14.8% of cara-
pace length in males (more than 12%
in 94% of cases) and 15.9% in fe-
males (more than 13% in 94% of
cases); plastron width at humero-pec-
toral seam (PWA) averages 36.2% of
carapace length in males (more than
33% in 97% of cases) and 38.3% in
females (more than 35.5% in 96% of
cases); posterior width of plastral
forelobe (PWB) averages 42.9% of
carapace length in males (more than
39% in 94% of cases) and 47.6% in
females (more than 45% in 94% of
cases); anterior width of plastral hind-
lobe (PWC) averages 38.0% of cara-
pace length in males (more than 34%
in 96% of cases) and 43.2% in fe-
males (more than 40% in 97% of
cases); maximum carapace length 185
mm in males, 160 mm in females;
Chihuahua and Texas to Jalisco,
Michoacan, and Mexico, except Cha-
pala, Zapotlan, Patzcuaro, San Juan-
ico, and Viesca basins popula-
tions with large plastron 7
5A. Head enlarged, jaws with extremely
broad alveolar surfaces; carapace
width averages 61.9% of carapace
length in males (less than 65% in
100% of cases) and 68.1% in females
(less than 71.5% in 100% of cases);
58
Tulanc Studies in Zoology and Botany
Vol. 23
plastral torelobe length averages
28.7% of carapace length in males
(less than 30% in lOOo/o of cases) and
29.0% in females (less than 30.5% in
100% of cases); plastral width at fe-
moro-anal seam (PWD) averages
28.2% in males (less than 29% in
100% of cases) and 31.4% in females
(less than 32.5% in 100% of cases);
interpectoral seam length averages
6.8% of carapace length in males (less
than 8% in 100% of cases) and 4.8%
in females (less than 6.5% in 100% of
cases); bridge length averages 17.3%
of carapace length in males (less than
17.5% in 100% of cases) and 23.9%
in females (more than 23% in 100%
of cases); Viesca area, Coahuila
K. h. tnegacephalum
5B. Head not enlarged, jaws with narrow
alveolar surfaces; carapace width av-
erages 72.0% of carapace length in
males (more than 65% in 100% of
cases) and 72.7% in females (more
than 66.5% in 100% of cases); plas-
tral forelobe length averages 31.2%
of carapace length in males (more
than 29.5% in 100% of cases) and
33.3% in females (more than 30.5%
in 100% of cases); plastral width at
femoro-anal seam (PWD) averages
29.1% in males (more than 28% in
100% of cases) and 34.0% in females
(more than 31% in 100% of cases);
interpectoral seam length averages
9.6% of carapace length in males
(more than 8% in 100% of cases) and
8.9% in females (more than 6% in
100% of cases); bridge length aver-
ages 18.2% of carapace length in
males (more than 16% in 100% of
cases) and 20.9% in females (less than
23.5% in 100% of cases); Pa'tzcuaro
and/or San Juanico basins, Michoa-
can 6
6A. Plastral scutes usually immaculate,
not darkly stained; maximum plastral
hindlobe length averages 30.1% of
carapace length in males (less than
32% in 100% of cases) and 31.5% in
females (less than 33% in 100% of
cases); plastral width at humero-pec-
toral seam (PWA) averages 33.7% of
carapace length in males (less than
38% in 100% of cases) and 34.8% in
females (less than 36% in 100% of
cases); posterior width of plastral
forelobe averages 41.9% of carapace
length in males (less than 43% in
100% of cases) and 43.5% in females
(less than 45.5% in 100% of cases);
interpectoral seam length averages
10.3% of maximum plastron length in
males (less than 12% in 100% of
cases) and 12.3% in females (more
than 11.5% in 100% of cases); first
vertebral scute width averages 22.3%
of carapace length in males (less than
23.5% in 100% of cases) and 21.4%
in females (less than 22.5% in 100%
of cases); San Juanico basin,
Michoacan K. h. magdalense
6B. Plastral scutes often stained red-
brown to dark brown; maximum plas-
tral hindlobe length averages 31.3%
of carapace length in males (more
than 29% in 100% of cases) and 34%
in females (more than 32% in 100%
of cases); plastral width at humero-
pectoral seam (PWA) averages 35.4%
of carapace length in males (more
than 33.5% in 100% of cases) and
36.8% in females (more than 35% in
100% of cases); posterior width of
plastral forelobe averages 43.6% of
carapace length in males (more than
41% in 100% of cases) and 46.8% in
females (more than 43.5% in 100% of
cases); interpectoral seam length aver-
ages 11.7% of maximum plastron
length in males (more than 9% in
100% of cases) and 9.1% in females
(less than 12% in 100% of cases); first
vertebral scute width averages 22.9%
of carapace length in males (more
than 20.5% in 100% of cases) and
24.5% in females (more than 22% in
100% of cases); Lake Patzcuaro
basin, Michoacan
K. h. tarascense
No. 1
Kinosternon Biosystematics
59
7A. Nasal scale triangular, rhomboidal,
or bell shaped; maximum plastron
length averages 86.4% of carapace
length in males (less than 90.5% in
100% of cases) and 91.6% in females
(less than 94% in 100% of cases);
bridge length averages 17.6% of cara-
pace length in males (less than 19.5%
in 100% of cases) and 21.7% in fe-
males (less than 23% in 94% of
cases); interabdominal seam length
averages 22.7% of carapace length in
males (less than 23.5% in 100% of
cases) and 24.3% in females (less than
26% in 94% of cases); interfemoral
seam length averages 6.9% of cara-
pace length in males (less than 8.5%
in 1(X)% of cases) and 7.1% in fe-
males (less than 8.5% in 100% of
cases); inter anal seam length averages
20.6% of carapace length in males
(more than 19% in 100% of cases)
and 25.8% in females (more than
23.5% in 100% of cases); Valley of
Mexico K. h. hirtipes
7B. Nasal scale deeply notched posteri-
orly (V-shaped); maximum plastron
length averages 86.0% of carapace
length in males (more than 81% in
95% of cases) and 92.6% in females
(more than 88% in 96% of cases);
bridge length averages 20.0% of cara-
pace length in males (more than 18%
in 97% of cases) and 23.7% in fe-
males (more than 21% in 95% of
cases); interabdominal seam length
averages 24.2% of carapace length in
males (more than 21.5% in 98% of
cases) and 26.0% in females (more
than 23% in 96% of cases); interfe-
moral seam length averages 9.0% of
carapace length in males (more than
6% in 98% of cases) and 9.0% in fe-
males (more than 6% in 99% of
cases); interanal seam length averages
18.6% of carapace length in males
(less than 22% in 95% of cases) and
23.4% in females (less than 26% in
93% of cases); Chihuahua and Texas
south to Michoacan, Jalisco, and
Mexico K. h. murrayi
Aknowledgments
I am deeply indebted to many persons
for the loan or gift of specimens or infor-
mation, including T. Alvarez, Walter
Auffenberg, R. H. Baker, CD. Barbour,
J. F. Berry, R. L. Bezy, J. Black, Bryce
Brown, C. C. Carpenter, A. F. Carr, G.
Casas Andreu, A. H. Chaney, J. Christ-
iansen, J. T. Collins, R. Conant, R.
Crombie, J, Cross, B. J. Davis, W. G.
Degenhardt, J. R. Dixon, N. H. Douglas,
H. A. Dundee, W. E. Duellman, M. J.
Fouquette, T. Fritts, D. Frost, A. L. Gen-
naro, J. W. Gibbons, U. Gruber, D.
Hahn, L. M. Hardy, M. M. Hensley, H.
Hidalgo, D. Hoffmeister, J. F. Jackson,
E. D. Keiser, A. G. Kluge, J. M. Legler,
A. E. Leviton, E. A. Liner, D. W. Linzey,
C. H. Lowe, J. D. Lynch, E. V. Malnate,
R. F. Malnate, R. F. Martin, H. Marx, T.
P. MasHn, C. J. McCoy, R. R. Miller, E.
O. Moll, O. Mooser, D. J. Morafka, R.
W. Murphy, C. W. Myers, M. A. Nick-
erson, R. Nussbaum, A. H. Price, G. G.
Raun, R. Reynolds, M. D. Robinson, D.
A. Rossman, J. F. Scudday, M. E. Seidel,
D. Smith, H. M. Smith, P. W. Smith, R.
C. Stebbins, W. Tanner, E. H. Taylor, D.
W. Tinkle, F. Truxal, T. M. Uzzell, R.
W. Van Devender, T. R. Van Devender,
R. Vogt, R. G. Webb, E. E. WiUiams, V.
Wilson, R. D. Worthington, J. W.
Wright, G. R. Zug, and R. G. Zweifel.
Tom Van Devender generously provided
numerous hve specimens from Sonora.
Permission to collect in Mexico was
granted through Miguel Angel Hernandez
Garcia and Ignacio Ibarrola Bejar of the
Direccion General de la Fauna Silvestre.
For unequaled field assistance, Diderot
Gicca, Sheila Iverson, Ron Magill, Peter
Meylan, and C. R. Smith deserve special
mention. Joan and Jill Iverson assisted in
computation of preliminary data. The
University of Florida, Florida State Mu-
seum and Earlham College provided sup-
port and study space. Portions of the field
work were supported by grants from
Sigma Xi, the Theodore Roosevelt Memo-
rial Fund, the American Philosophical
Society, the Earlham College Faculty
60
Tulane Studies in Zoology and Botany
Vol. 23
Development Fund, and the National
Science Foundation (DEB-8005586). Jim
Berry, Roger Conant, Mike Seidel, and
Hobart Smith each offered helpful com-
ments on the manuscript. Sheila Iverson
typed the manuscript.
RESUMEN
Se analizaron las variaciones geograficas del
escudo y las medidas de las conchas (mediante ana-
lisis estadi'stico multivariado), tamaTio del cuerpo,
morfologia de las escamas de la cabeza y del
menton, tamano del primer hueso neural, escama-
cio'n irregular, asi como tamano de la cabeza y los
patrones de poblacidnes de la tspecit ^ Kinosternon
hirtipes. Los resultados sustentan la retencidn de las
especies alopatricas A^. sonoriensey K. hirtipes cofho
especies completas dentro del grupo, y el recono-
cimiento de dos subespecies alopatricas (una de ellas
nueva) de K. sonoriense y de seis subespecies (cuatro
de ellas nuevas y todas aparentemente alopatricas)
de K. hirtipes. La descripcion de cada taxon incluye
datos completes de sinonimias, ecologia y repro-
duccidn. Tambien estan incluidas claves para
adultos y una discusion de todos los taxa.
Specimen List
All specimens examined as well as local-
ities plotted in Figure 1 are listed below by
drainage basin sample used in the analy-
sis. Basins are Usted under the appropriate
taxon in approximate geographic order
from northwest to southeast. Localities
(including literature records) within each
basin are listed alphabetically by state,
county, and specific locality. Specimens
marked with an asterisk were not exam-
ined. All distances are in km. The fol-
lowing abbreviations are used throughout
the list: C = city or ciudad; Cn = can-
yon; Cr = creek; Hwy = highway;
Mtn(s) = mountain(s); nr = near; R
= river or ri'o; Rd = road; Spg(s) =
spring(s); trib = tributary; and vie =
vicinity.
K. sonoriense sonoriense.
BILL WILLIAMS (BIG SANDY) RIVER (WILL).
ARIZONA. Mojave Co.: Big Sandy Basin, NW
Wickenburg, UAZ 30826*; Burro Cr Camp-
ground, ASU 13785; 14.5 km E Burro Cr
Campground, ASU 13786; Trout Cr (Hulse,
1974).
GILA and LOWER COLORADO RIVERS
(GILA).
ARIZONA. Cochise Co.: Babacomari R, ca. 4.8
km W Huachuca C, UAZ 38861*; Bear Cn,
16.1 km W Coronado International Memorial,
ASU 13783*; Bear Cn, Huachuca Mtns, Monte-
zema Pass Rd, UAZ 27982*; Fort Huachuca,
first cienega above post, USNM 17780-81*,
19680*, 21718-19*, 45305* (Stejneger, 1902); nr
Hereford, San Pedro R, KU 15927*, CAS-SU
48886-87*; Huachuca Mtns, AMNH 19450,
USNM 20975-77*, 20979-80* (Van Denburgh
and Slevin, 1913; Van Denburgh, 1922); Lewis
Spgs, AMNH 15165-69, 18103, 18656-57,
UMMZ 118269; 3.2 km S Miller's Peak, Hua-
chuca Mtns, Cochise Cn, CAS-SU 13888*;
Pyeatt Ranch nr West Gate Fort Huachuca, JBI
410-14; San Pedro R, USNM 20547-55*; San
Rafael Valley, UMMZ 88476*; Hwy 80 at St.
David (Kauffeld, 1943); Vasquez Ranch, St.
David, UAZ 32960*. Gila Co.: Cibecue Cr nr
Salt R, ASU 10530* (Hulse, 1974); Coyote Cn,
ASU 10903-04* (Hulse, 1974); 66 km NNE
Globe, Salt R, UMMZ 105791 (Duellman,
1955); Mezquite Flat at SaU R, ASU 10527-29*
(Hulse, 1974); Natural Spgs, just N Payson
(J. F. Berry, pers. comm.); Payson, ASU
4142*; 4.8 km N Punkin Center on Tonto
Creek, ASU 12061-68*; Rice, San Carlos Indian
Reservation, USNM 59738*; Roosevelt Reser-
voir (Little, 1940); San Carlos River, N San
Carlos, UMMZ 105821 (Duellman, 1955);
Spring Cr, 16.1 km W Young, UMMZ 105756
(Duellman, 1955); Tonto Cr nr Gisela, ASU
2372* (Hulse, 1974, 1976). Graham Co.: Bonita
Cr, NE of Safford, UMMZ 105792 (Duellman,
1955); Marijilda Cr (Nickerson and Mays,
1971); 8.0 km S Safford (Nickerson and Mays,
1971); 9.7 km S Safford, UMMZ 105765,
105293 (Duellman, 1955); no further data,
USNM 55627-28 (Van Denburgh, 1922 as K.
flavescens; Iverson, 1978). Greenlee Co.:
Virden, 1 .6 km W New Mexico State line, UNM
15561. Maricopa Co.: Agua Caliente, CAS-SU
39102*; Box Cn, 8.0 km N Wickenburg (Gates,
1957); Cave Cr, CAS-SU 17282*, KU 15926*.
UAZ 35948*; Cave Cr, Fairbank, CAS-SU
20643*, 35157* (Van Denburgh and Slevin,
1913; Van Denburgh, 1922); Granite Reef Dam,
ASU 4549*; Guadalupe, ASU 1972*; Hassay-
ampa R, 8.0 km S Wickenburg, CHAS 16177
(Gates, 1957); Hassayampa R, 8 km SE Wick-
enburg, UIMNH 85839, 85842; Mesa, ASU
336*; Phoenix, AMNH 73821-22*. ASU 4268*.
UMMZ 69417-20, 72497. USNM 55625-26*
(Van Denburgh. 1922); Phoenix, Salt R. KU
No.
Kinosternon Biosystematics
61
2908, UMMZ 15755*. USNM 15755* (Iverson,
1978); 48.3 km SW Phoenix, Gila R, KU
15928*. Sycamore Cr at Sunflower. ASU
13801-03*. CM 57121. 57113-14 (Hulse. 1974,
1976); Sycamore Cr at Hwy 87, ASU 12105*;
Sycamore Cr, 1.6 km S. Sunflower. UU 11537-
39*; Tempe. ASU 1004*. Navajo Co.: Fort
Apache (Hulse, 1974); Rock Cr Cn, S. Camp
Apache. USNM 1103* (Yarrow 1875 as K. hen-
rici; Van Denburgh. 1922). Pima Co.: Annilo
Tank, R17E, T14S. Sec 3, NE %, UAZ 36510*;
Arivaca, 0.8 km SW of Post Office, UAZ
30821, 30823; 0.8 km E Arivaca, UAZ 30824;
Madrona Cn. Rincon Mtns. UAZ 27985*,
36512*, FB 1551; Molina Basin, Santa Catalina
Mtns, UAZ 27998*; Posta Quemada Cn, SE
side Rincon Mtns, UAZ 24753*; Rincon Mtns,
end of Kennedy Rd via Speedway, UAZ
30825*; Rincon Stock Farm, nr Tucson,
UMMZ 89871-73; Sabino Cn, Santa Catalina
Mtns, CAS-SU 8637-38*, FMNH 74777,
SDNHM 14225, UAZ 27997* (Van Denburgh
and Slevin, 1913; Van Denburgh, 1922); Santa
Catalina Mtns, AMNH 4520; Tanque Verde
Ranch, SDNHM 16232-37; Tucson, Santa Cruz
R, AMNH 2565, 20538, CAS-SU 33850-66*,
MCZ 1920, USNM 67*, 17018-21*, 16835-36*
(LeConte, 1854; Agassiz. 1857; Baird. 1859;
Yarrow. 1883; Gunther. 1885; Van Denburgh
and Slevin. 1913; Van Denburgh, 1922); Tucson
Sewage Disposal Area, UAZ 28002*; nr Xavier,
16.1 km S Tucson, CM 19287. Pinal Co.: Boyce
Thompson SW Arboretum, 6.4 km W Superior,
AMNH 66336, CHAS 9494-97. 9644. 9648.
10324. UMMZ 85076 (14 specs); Queen Cr,
Arboretum, CHAS 9879-80, 13634-44; Superi-
or, CHAS 10325. UAZ 27994-95*. Santa Cruz
Co.: Alamo Cn. 4.0 km SW Pena Blanca
Camp. Pajarito Mtns, MVZ 50903-06. UAZ
15104*; Babacomari R at Babacomari Ranch.
ASU 12107-113*; G. A. Jones Ranch at Parker
Cn. UAZ 27986*; Lochiel, ASU 13804*; Mon-
key Spg. ASU 12077*; Nogales. USNM 17127-
36*. ASU 13787* (Van Denburgh. 1922); 19.3
km W Nogales. CM 25209; 6.8 km S Patagonia
on Hwy 82. LACM 64223; Pena Blanca Spg.
TUL 15040-41, UMMZ 75814, 75855 (Camp-
bell, 1934); Santa Rita Mtns. CAS-SU 48885*
(Van Denburgh. 1922); SW of Tucson. AMNH
2559-62. UMMZ 118268; Tumacacori Mtns.
SDNHM 5720, CAS-SU 81457-58*; Turkey Cr
at Canelo, UAZ 27988*. Yavapai Co.: Bard,
SDNHM 33866; 12.9 km S Camp Verde,
SDNHM 17889; 4.8 km N Clarksdale, Verde R,
UU 15078-84*; Ft. Verde, USNM 14807-09,
15708 (Van Denburgh, 1922 as K. flavescens
and K. sonoriense; Iverson, 1978); Fossil Cr,
9.7 km N Verde R, ASU 12151-56* (Hulse,
1974); Hassayampa R at Wagoner, CHAS
16631; Hassayampa R. 3.2 km S Wagoner.
CHAS 15834; Montezuma's Well. ASU 4573*.
UU 13031*; Peck's Lake, NE Clarksdale, JBl
386-88; Rock Spgs, CM 47751. MSU 3578;
Stehr Lake, ASU 13790*; Sycamore Cr. E of
Dugas, UMMZ 105822 (Duellman. 1955);
Sycamore Cr at Verde R, ASU 12074-76*; Tule
Stream, ASU 10962-67*, CM 57115. 57122
(Hulse, 1974, 1976); Entrance to Tuzigoot
National Monument, ASU 13789*; Verde R.
above Camp Verde. UMMZ 105823*
(Duellman. 1955); Verde R in Cottonwood. JBl
524. Yuma Co.: Gila C. Gila R, USNM
21716-17*, 21817*; Gila R. Adonde Siding,
USNM 21715* (Van Denburgh. 1922); North
Gila East Main Canal, 1.6 km SW Laguna
Dam, RSF 468* (Funk, 1974); Warshaw, Mex.
Boundary line, USNM 21712-
14*; Yuma (Van Denburgh and Slevin, 1913;
Van Denburgh, 1922).
CALIFORNIA. Imperial Co.: Palo Verde, MVZ
6282 (Van Denburgh, 1922); No further data,
CAS-SU 33408 (Van Denburgh and Slevin,
1913; Van Denburgh, 1922).
NEVADA. Clark Co.: Pyramid Cn (LaRivers,
1942, as K. flavescens, but see Iverson, 1978).
NEW MEXICO. Catron Co.: Glenwood, San
Francisco R, CM 18310; Taylor Cr, 2.4 km NE
Wall Lake, UMMZ 134282-84, UNM 2568
(Niles, 1962); Wall Lake, 13.7 rd km SSE
Beaverhead, UMMZ 134281, UNM 20552, 20609-10
(Niles, 1962). Grant Co.: Bennett Ranch. W
Cliff. UNM 8157-69; 3.2 km ENE Cliff (Niles,
1962); 1.6 km E Bedrock Post Office, S side
Gila R, UNM 20611. Undetermined Co.: Gila
R, ANSP 83 (holotype of Kinosternum henrici).
SONORA. R Nutrias, above Nutrias Dam, UM-
MZ 105817; R San Pedro, above Elias Dam,
UMMZ 105816, 105818-20; R Santa Cruz. 6.4
km S Arizona border. UMMZ 105814-15; San
Pedro R, USNM 20968 (Van Denburgh. 1922);
Sierra Magallones. UAZ 36497*.
SW NEW MEXICO INTERIOR DRAINAGES
(SWNM).
ARIZONA. Cochise Co.: N of Rodeo, nr New
Mexico border, UMMZ 86081-86 (Niles, 1962).
NEW MEXICO. Hidalgo Co.: Clanton Cn, 16.1
km N Cloverdale. LACM 7967-70, 7994; 8 - 9.7
km W Cloverdale Store, UNM 20558; Guada-
lupe Cn, 3.1 km E, 2.3 km N Arizona-New
Mexico border, UNM 14061; W slope Pelon-
cillo Mtns, T32S, R21W, Sec 16, NE '/a, UNM
15618; 24.1 km N Rodeo, San Simon Marsh.
62
Tulane Studies in Zoology and Botany
Vol. 23
NMSU 3050*; San Simon Cienega, UMMZ
105800 (Niles, 1962); Skeleton Cn, Peloncillo
Mtns, AMNH 109056, MVZ 70350.
RIO MAGDALEN A (MAGD).
ARIZONA. Santa Cruz Co.: California Gulch,
ASU 13633-37*. CM 57116-20; Ruby, UIMNH
4129, UMMZ 107480 (Dueliman, 1955); Syca-
more Cn, UAZ 28000*, 30822*, 33582*.
SONORA. Imuris. UIMNH 85832; 14.5 km N.
Imuris, KU 44503-25; 14.5 km NNE Imuris,
KU 48562-63, 50734*, 51429; 1.1 km S Magda-
lena, UAZ 28010; nr Magdalena, MCZ 46649*;
25.1 km NNE Magdalena, UMMZ 126442; 42
km S Nogales, Rancho de Tascara, AMNH
73004; 69.5 km S Nogales on Hwy, 2, LACM
61107; R Arizona, vie. Rancho de la Arizona,
UAZ 28010-11; R Magdalena, 1.6 km SE Cab-
orca, MVZ 51355 (Zweifel and Norris, 1955).
RIO SONORA (SNRA}.
SONORA. Arispe, UAZ 27976, 28003-07, 28012-
14, 28016-18, 28020-21; 24.1 km W Cananea,
AMNH 67503-05, 67507; 4.8 km downstream
from Cucurpe, UAZ 36509; Hermosillo, AM-
NH 74945; 24.1 km N Hermosillo (Taylor,
1936); Cjenega nr Rancho Agua Fria, E
Cucurpe, 'jBI 799-803, 866-870; 16.1 km E
Ures, R Son ora, NMSU 4101*.
RIO YAQUI. (YAQ).
ARIZONA. Cochise Co.: Ashton Spg, nr San
Bernardino Ranch, UAZ 28001*; Black Dam,
San Bernardino Ranch, UAZ 27999*; Chirica-
hua Mtns, USNM 33929-30* (Van Denburgh,
1922); 8.0 km S McNeal on Hwy 666, LSU
9861; San Bernardino Ranch, 27.4 km E Doug-
las, CM 40407, ASU 13784*; San Bernardino
Ranch, Mex. boundary, USNM 21104*; nr
Turkey Cr Ranger Station, UMMZ 105675
(Dueliman, 1955).
CHIHUAHUA. Bavispe R, below 3 Rivers,
Chihuahua-Sonora border, BYU 14629; R
Gaviian, 11.3 km SW Pacheco, MVZ 46646.
SONORA. Guadalupe Cn, nr Monument 72,
Mex. boundary line, USNM 20970 (Agassiz,
1857; Baird, 1859; Yarrow, 1883; Van Den-
burgh and Slevin, 1913; Van Denburgh, 1922);
14.2 km W Maicova, UAZ 39968; Ranchito
Finos Altos, Sierra Nacori, UAZ 31613-14;
San Bernardino Ranch, USNM 20981-88 (Van
Denburgh, 1922). Yecora, UAZ 28211, 35209-
11*; 18.0 km E Yecora, UAZ 40105.
Rio CASA GRANDES INTERIOR BASIN (CSGR).
CHIHUAHUA. 3.2 km N Old Casas Grandes,
BYU 14132-33; Colonia Juarez, R Piedras
Verdes, FMNH 1873 (2), UNM 30393-99, UU
11522-36; 2.6 km NW Colonia Juarez, UF
47642-43, JBI 946-47; 10.5 km NW Cohania
Juarez, ASU 5207-08*; Ramos, MVZ 46647-50.
RIO FUERTE (FRTE).
CHIHUAHUA. Cerocahui, BYU 14625, 14627,
14628 (see text).
QUESTIONABLE DATA.
JALISCO. 12.1 km N Magdalena, BYU 14630
(Tanner and Robison, 1960).
Kinosternon sonoriense longifemorale
RIO SONOYTA (SNTA).
ARIZONA. Pima Co.: Quitobaquito Pond, JBI
391, 696-699, 701-706, UF 47719-20 (para-
types); Organ Pipe National Monument Col-
lection (4 uncatalogued specimens), LACM
105399, SDNHM 47316, UAZ 27987 (para-
types,) 27993, 27996 (paratypes) (Stebbins,
1966).
SONORA. Sonoyta, USNM 21709-11 (paratype,
holotype, and paratype, respectively); Sonoyta
R, USNM 21725; Sonoyta R, 4.8 km from Son-
oyta, USNM 21708 (Van Denburgh, 1922)
(paratype); 29.0 km W Sonoyta on Hwy 2,
LACM 105400.
Kinosternon hirtipes murrayi
RIO SANTA MARIA INTERIOR BASIN (STMR).
CHIHUAHUA. Galeana, R Santa Marfa, BYU
15266-76; nr Galeana, R Santa Maria, BYU
16846-47, UMMZ 117783-84 (Semmler et al,
1977); 4.8 km N and 3.2 km W Galeana, R
Santa Mari'a, UU 4457-80, 1251 1; ca. 4.8 km
SE Galeana, UAZ 36349*; 9.7 km NW Gale-
ana, R Santa Maria, MCZ 62516-22; Ojo de
Galeana, 7.2 km SE Galeana, ASU 5169-82*,
5185-95*, FB 1695*, 1844*, JBI 808-09, 815-20,
838-43, 850, 958-61, UAZ 27965-70*, 34766*,
UF 40536-49, UNM 32600-12; outHow of Ojo
de Galeana, 3.4 km S Galeana, ASU 5196-205*;
nr Progreso, R Santa Maria, UMMZ 118284-
89, USNM 105026-28, 105031-34; R Santa
Maria, USNM 30841-43; San Buenaventura,
below Presa El Tintero, R Santa Man'a (Casas
Andreu, 1967).
RIO CARMEN ( = SANTA CLARA) INTERIOR
BASIN (CRMN).
CHIHUAHUA. 3.2 km W Carmen, R Carmen,
UU 8539-43; 1.6 km S and 0.8 km E Santa
' Clara, R Santa Clara, MVZ 72819-43,
89676-77; 3.2 km S Santa Clara, MVZ
70688-95; R Carmen at Ricardo Flores Magon,
UMMZ 125362.
RIO SAUZ INTERIOR BASIN (SAUZ).
CHIHUAHUA. Arroyo El Sauz, El Sauz, UU
8549-53; 5 mi N Cerro Campana, MVZ 68915;
No. 1
Kinosternon Biosystematics
63
nr Encinillas, UMMZ 117781-82, 117785; Ojo
Laguna, MVZ 70696-98; Sauz, FMNH 1405 (5);
UMMZ 117426-29.
ALAMITO CREEK DRAINAGE. (TEX).
TEXAS. (See discussion in Conant and Berry,
1978). Presidio Co.: Casa Piedra, Willie Russell
Ranch, DMNH 985, 1095-96; Marfa, USNM
15860 (paratype) (Glass and Hartweg, 1951)
(data obviously in error; see Conant and Berry,
1978); 48.3 km S Marfa, Harper Ranch, USNM
198055; 59.5 km S Marfa, Harper Ranch,
TCWC 650 (holotype) (Glass and Hartweg,
1951); 60.3 km SSE Marfa, UMMZ S1083,
101294 (paratypes) (Glass and Hartweg, 1951;
Peters, 1952).
Rib CONCHOS (CNCH).
CHIHUAHUA. Boquilla Culebra, UIMNH
52198 (Smith et al, 1963); 1.6 km N Camargo,
UU 8548; 8 km N Camargo, UMMZ 118075;
20 km W Camargo, Arroyo del Vado o La Pal-
oma, Presa La Boquilla (Casas Andreu, 1967);
27.4 km SW Camargo, UU 8469-89, 8490-98;
27.5 km SW Camargo, UIMNH 43528; R Cata-
lina, 24.1 km N Villa Ocampo, Durango, UU
12758-59; 8 km N Chihuahua, MVZ 66121*;
8 km N Falomir, UIMNH 52199-201 (Smith et
al, 1963); 0.8 km N Guadalupe Victoria, KU
51237-38*, 51259-60; Guardiola, UIMNH
52194-97 (H.M. Smith et al, 1963); 4.8 km S
Hidalgo del Parral, UU 8468; 12.9 km SW
Hidalgo del Parral, TCWC 20812; 4.8 km SW
Jimenez, KU 53758-84; Julimes, ANSP 20106-
08, UIMNH 52190-93, UU 8546-47 (Smith et
al, 1963); 9.7 km NE La Boquilla, UNM 467;
0.8 km E La Cruz, KU 48259-62; 0.4 - 1 .6 km E
La Cruz, UIMNH 43511-27 (Williams et al,
1963); cited erroneously as Lago Toronto by
Casas Andreu, 1967); Meoqui, R San Pedro,
MVZ 52256; 8 km N and 8 km E Meoqui, KU
33903*; nr Ojinaga, AMNH 113858-59* (Con-
ant and Berry, 1978); 1.6 km NW Ojinaga, KU
52159, 69849 (Legler, 1960); R San Pedro, 78.8
km SE Chihuahua, MVZ 57467; Mouth of R
San Pedro, KU 51221-33, 51239-56, 51276,
51291-98, 51316-20, 52147-57, 56163-64,
9136572 (Legler, 1960); 1.6 km upstream from
mouth of R San Pedro, KU 51234-36, 51257-58;
12.9 km SE Santa Barbara at Rafael, AMNH
6792325; Santa Rosalia, FMNH 5930 (2).
DURANGO. 4.8 km E Las Nieves, R Florido,
MSU 3180-89.
LAGUNA BUSTILLOS INTERIOR BASIN (BUST).
CHIHUAHUA. 27.4 km N Cuauhtemoc, trib to
Laguna Bustillos, UMMZ 125358-61.
RIO PAPIGOCHIC DRAINAGE (PAP).
CHIHUAHUA. 8 km N, 1.6 km W Cd Guerrero,
R Papigochic, KU 45020-25, 51425-26, 87854;
El Riyito, 17.7 km WNW Cocomorachic, KU
51311, 51313-14; Minaca, FMNH 1102, MVZ
58967-70; 3.2 km W Minaca, KU 51261-309,
52142-43, 87853, 91364, 91373-78; 5.5 km
NE Minaca, BYU 16848; Ri'os Papigochic and
Tomochic (Legler and Webb, 1970; erroneous-
ly recorded as K. sonoriense and K. hirtipes hir-
tipes); Yepbmera, FB 1545-46, 1595-97, JBI
403-404, MSU 3579, UAZ 34168*; 1.6 km N
Yepbmera, JBI 821-23, 835-37, UF 40389-400,
UNM 32588-599; 3 km N Yepbmera, UAZ
34169-70*; 3 km W Yepdmera, MCZ 79029-38,
79039-46; 4 - 5 km N Yepomera, Arroyo de la
Huachin, UAZ 34171-72*.
RIO NAZAS INTERIOR DRAINAGE (NAZ).
DURANGO. Lerdo, USNM 61687-88; 24.1 km
SW Lerdo, AMNH 67496-500, UMMZ 118267;
between Lerdo and La Goma, USNM 105262-
64; R Nazas, at Cardenas Dam, nr El Palmito,
JBI 826-31, UU 8461-66; 22.5 km NE Pedriceffa
UIMNH 19339; La Concha, nr Penon Blanco,
AMNH 88883; Presa Francisco Zarco on R
Nazas nr Graseros, ENCB 10893-94, JBI 948-
50, UF 47602; Trib to R San Juan at Hwy 45,
5.6 km N turnoff to Primo Verdad, UU 12075-
77; Rodeo, AMNH 87654-57, 96589; 13.5 km
S San Jacinto, R Nazas, UF 40425-27; 16.1 km
W Torreon, R Nazas, USNM 105270-71.
RIO AGUANAVAL INTERIOR DRAINAGE
(AG UN).
ZACATECAS. 24.1 km NW Fresnillo, R Florido
AMNH 85285-91; 25.7 km N Fresnillo, UMMZ
118056-057, and 118060*; La Florida, R Flor-
ido, UU 12078-80; Rancho Grande, R Medina,
AMNH 85296; 1.6 km N Rancho Grande, R
Nieves, UU 8499-538, 8544-45; 17.7 km E Som-
brerete, UIMNH 28155; 46.7 km E Sombrerete,
UMMZ 126284.
LAGO SANTIAGUILLO INTERIOR DRAINAGE
(STGO).
DURANGO. 22.5 km SE Chinacates, AMNH
88882; trib to Lago Santiaguillo, at bridge in
Guatimape, UF 40428-30.
RIO MEZQUITAL DRAINAGE (MEZ).
DURANGO. ca. 5 km from Colonia Hidalgo, km
937, Torrebn-Durango Hwy (Casas Andreu,
1967); 4.8 km E Durango, AMNH 85294; 9.7
km E Durango, R Tunal, AMNH 85292-93;
10.5 km E Durango, R Tunal, UU 4481-520,
12512-15; 15.8 km N Durango, UIMNH 7051,
23844; 16.1 km N Durango, R Canatlan, MVZ
57333-35; 17.7 km E Durango on Hwy 45, TUL
18680; 17.7 km E Durango, R Santiago, MVZ
58222; 6.4 km E and 11.3 km S Durango, R
Santiago, MSU 4245-56; 25.4 km SW Durango,
64
Tulane Studies in Zoology and Botany
Vol. 23
R Chico on Hwy 40, LSU 34319, JBI 954-55,
UF 47603-04; ENCB 10904-08; 27.4 km N Dur-
ango, CU 46115-16; 37 km N Durango, MSU
7869; kilometer 48.5, N of Durango, Hwy 45,
UF 40424; nr Durango, 6.4 km E and 3.2 km
NE jet. hwys to Torreon and Fresnillo, UMMZ
122245-54; 0.8 km N Graceros, KU 68733-36,
68738-45 (KU 68737 is K. integruml); 6.4 km
SW La Pila, KU 51083-84, MSU 2680-82, 2684,
2686-89, 10197-98; 9.7 km NW La Pila, KU
51085-86; R Mezquital, at Mezquital, 86.7 km
SSE Durango, TUL 18670; 6.4 km S Morcillo,
MSU 4243-44 (basis of Stebbins' 1966 southern
Durango /lavescens record; see Iverson, 1978);
Ojo de Agua de San Juan, 1 .6 km N Los Berros,
UMMZ 129824-28; Otinapa, AMNH 68382; R
La Sauceda at Hwy 40, ENCB 10894-903, JBI
825, 832-34, 951-53, UF 40401-23, UNM 32588-
99; R Soledad, La Soledad, MSU 2683, 2685;
6.4 km S Villa Union jet. Hwy 45, CM 53987.
EL SALTO (ACAPONETA) BASIN (SALT).
DURANGO. 9.7 km ENE El Salto, Hwy 40.
ENCB 10909-14, JBI 956-57, LSU 34320, UF
47605-06.
RIO SANTA MARIA BASIN (SLP).
SAN LUIS POTOSI. Laguna de las Rusias, LSU
7873-75 (Williams and Wilson, 1966); Arroyo
la Hilada, ca. 1 km N Presa El Refugio ( =
Laguna de las Rusias), UF 42803-815.
Rfo AGUASCALIENTES DRAINAGE (AGUAS).
AGUASCALIENTES. Aguascalientes, MCZ
79047; Aguascalientes, R Morcinique, MU 793;
2.1 km E Aguascalientes, UIMNH 43582; R
Penueia nr Aquido, CAS-SU 19702-03; R Jo-
coque Dam, SE end Presa Jocoque, CAS-SU
19692-95; 1.2 km W Santiago, R Jocoque,
CAS-SU 19696-701.
RIO VERDE DRAINAGE (VERDJ.
JALISCO. El Olivo, 19.3 km W Lagos de Mo-
reno, AMNH 117953; Presa el Cuarenta nr
Paso de Cuarenta, JBI 896-900, UF 44064-65,
44078; 3.2 krn NE Valle de Guadalupe, trib to R
Verde, Hwy 80, JBI 893-95, TUL 18671, UF
44077.
MARAVATIO BASIN (MAR/.
GUANAJUATO. 1.6 km SE Inchamacuaro, KU
43637.
BAJIO BASIN (BAJ).
GUANAJUATO. No further data (Westphai-
Castelnau, 1872); R Turbio, 12.9 km E Pen-
jamo, UU 12081-82; R Urdo, Valle de Santiago
(Caballero y C. y Cerecero, 1943; Caballero y
C, 1940a); 16.1 km N San Miguel de Allende,
AMNH 93363; 22.5 km N San Miguel de Allen-
de, AMNH 85295; Arroyo el Sauz, ca. 10.5 km
N Yuriria-Salvatierra Hwy (Casas Andreu,
1967); Taboado, 9.7 km NW San Miguel Al-
lende, AMNH 71033, FMNH 71029; Hwy 51,
6.0 km S jet. Hwys 51 and 110, UF 43613-15;
11.9 km S jet. Hwys 51 and 110 at Sebastian,
UF 44074, JBI 908.
JALISCO. R Lerma, 0.8 km NW jet. Hwys 90
and 110, UU 12120.
LAKE CUITZEO INTERIOR BASIN (CUIT).
MICHOACAN. Lago Cuitzeo (Casas Andreu,
1967); Lake Cuitzeo, San Agustin, UMMZ
97136 (Duellman, 1961).
VILLA VICTORIA BASIN (VILLA).
MEXICO. 11.3 km W Villa Victoria, USNM
108719-26, UMMZ 118295-296; 3.7 km S La
Presa, JBI 928; 8.9 km S La Presa, JBI 927.
RIO BALSAS DRAINAGES (BALS).
MICHOACAN. 8 km W C Hidalgo, AMNH
62257 (UIMNH 24707 from the same locality
is K. integrum, not K. hirtipes, as listed in
Duellman, 1961).
PUEBLA. Trib to R Atoyae, 4.5 km S Molcaxae,
UU 2096 (Data questionable).
Kinosternon hirtipes megacephalum
VIESCA INTERIOR BASIN (VCSA).
COAHUILA. 3.2 km SE Viesea, SM 11460-66
(paratypes and holotype); 9.7 km SW Viesea,
SM 9823 (paratype).
Kinosternon hirtipes tarascense
LAGO PATZCUARO INTERIOR BASIN (PATZ).
MICHOACAN. Lago Patzeuaro, FMNH 1397,
2036, JBI 880-84, UF 43505-07 (paratype, holo-
type, and paratype), 43595-96 (paratypes),
UMMZ 96988-91, 97131, 99762, 1 17798 (Duell-
man, 1961); Lago Patzeuaro, nr E end, UF
7075; Isla Janitzio, Lago Patzeuaro, CU 16142;
Canal de la Tzipecua, SW margin Lago Patz-
euaro (Casas Andreu, 1967); Tzintzuntzan,
AMNH 82128.
Kinosternon hirtipes magdalense
SAN JUANICO ( = MAGDALENA or TOCUMBO)
VALLEY INTERIOR BASIN (SNJ)
MICHOACAN. Atop Presa San Juanico (road to
dam meets Hwy 15 ca. 56.3 km W of Zamora),
TUL 18677 (paratype); Presa San Juanico, at
dam, UF 45035-36 (holotype and paratype),
45038-40 (paratypes), and 45041.
No.
Kinosternon Biosystematics
65
Kinosternon hirtipes hirtipes
VALLEY OF MEXICO (VALLE)
DISTRITO FEDERAL. Mexico C. Senck 47875*
(Greene, 1972); vie Mexico C (Beltz, 1954);
San Juan Tezompa, 19.3 km E Xochimilco,
UMMZ 99446-60; Valley of Mexico, Xochimil-
co, USNM 61247; Xochimilco, UMMZ 69264
(Caballero y C, 1939); Lake Xochimilco, nr
Mexico C, MCZ 7866, UMMZ 80356-57.
MEXICO. Chalco, FMNH 1406 (Gadow, 1908);
Teotihuacan, San Juan, AMNH 17859-62;
Lake Texcoco, nr Mexico C, AMNH 68699;
Valle de Mexico, CAS-SU 5849-50 (Martin
del Campo, 1938; Hartweg and Glass, 1951;
Deevey, 1957; Kranz et al, 1970).
STATE UNCERTAIN. "Mexico", ZSM 1374/0
(Holotype of Cinosternon hirtipes; Wagler,
1830).
Kinosternon hirtipes chapalaense
LAGO DE CHAP ALA BASIN (CHAP).
JALISCO. Lago de Chapala, Beach at Chapala,
UMMZ 97190; Lago de Chapala, 0.4 km off
Chapala, UMMZ 97121-130 (includes holo-
type and paratypes); Lago de Chapala, 3.2 km
W Chapala, UU 12126-28; (paratypes) Lago de
Chapala, 0.8 km E Tuxcueca, JBI 890; Lago
de Chapala, 6.1 km W Ajijic, UU 12125; para-
type 3.2 km S Jamay, AMNH 17856; 3.2 km
SE Ocotlan (El Fuerte), UMBM 2403; Ocotlan,
UMMZ 76129, 1 17796-97 (UMMZ 1 17801 from
this locality is K. integrum.)
MICHOACXn. Jiquilpan (Duellman, 1961); La
Palma, USNM 108718 (Duellman, 1961).
LAGO DE ZAPOTLAn INTERIOR BASIN
(ZAPO).
JALISCO. 1.6 km NW C Guzman, Lago de
Zapotlin, UMMZ 1 17259-66; 3.2 km N C Guz-
man, UMMZ 102154; Laguna Zapotlan, BM-
NH 1906.6.1.253-5* (Gadow, 1908 as K. inte-
grum).
Kinosternon hirtipes chapalaense x murrayi
RIO DUERO DRAINAGE (DUER).
MICHOACAN. Lake Camecuaro, 14.5 km E
Zamora, JBI 885-889, UF 43603-610, 44062-63,
44075-76, UMMZ 97132-35, 102150-53 (Duell-
man, 1961).
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.
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LIFE HISTORY OF ETHEOSTOMA COOSAE (PISCES: PERCIDAE)
IN BARBAREE CREEK, ALABAMA
PATRICK E. O'NEIL
Geological Survey of Alabama,
P.O. Drawer O,
University, Alabama 35486
Abstract
Collections of Eiheostoma coosae (Coosa darter)
were made from April 1977 to April 1978 in Bar-
baree Creek (Coosa River system), Clay County,
Alabama. The principal habitat of this species was
cobble and/or gravel raceways, riffles and pools.
Spawning occurred on the surface of rocks and small
boulders from mid March to late May with peak
activity in mid April. The spawning position was
either inclined or semi-inverted horizontally. Indi-
viduals reached sexual maturity and spawned by the
first year. Maximum age was three years. By the end
of the second year, the average size of males and
females -was 41 .0 mm and 36.5 mm SL, respectively.
The sex ratio, 1:1.3, was significantly different from
1:1. The principal diet consisted of Copepoda,
Cladocera, Ephemeroptera (Isonychia) and Diptera
(Chironomidae, Simuliidae).
INTRODUCTION
Eiheostoma (Vlocentra) coosae is
endemic to the Coosa River system of
Alabama, Georgia, and Tennessee. The
biology of the subgenus as a whole is
largely unknown. Published papers in-
clude studies by Winn (1958a, 1958b) on
the reproduction of two undescribed
forms. Stiles (1975) on the reproductive
behavior of Etheostoma simoterum, and
Ultsch et al. (1978) on habitat selection by
Etheostoma duryi. This study reports on
the life history of E. coosae in Barbaree
Creek, an eastern Alabama stream.
STUDY AREA
A section along Barbaree Creek,
T.18S., R.7E, Sec. 22, Clay County,
Alabama (Coosa River system) was cho-
sen as the study site. Barbaree Creek is a
perennial stream flowing through north-
ern Piedmont physiography. Its head-
waters originate in the Talladega Moun-
tains.
The substrate consisted of gravel and
sand shoals interspersed with patches of
cobble and boulders that were regularly
broken by cobble or slab riffles. Bedrock
was usually exposed below riffles whereas
the pools contained unconsolidated mat-
erial. The ranges of measured water
quality values were: disssolved oxygen,
7.9-13.0 ppm; pH, 6.8-7.3; turbidity,
0.7-2.8 JTU; conductivity, 17-45 umbos;
and stream temperature 3.7-25 °C
(Boschung and O'Neil, 1980).
METHODS
Specimens of E. coosae were collected
monthly from April 1977 through April
1978. Small mesh minnow seines, 3/16
inch delta weave, and a backpack shocker
were each operated approximately 1.5
hours during each monthly collection,
sampling a variety of habitats. Upon
capture the fishes were preserved in a
20-percent formalin solution.
In the lab, fishes were blotted dry and
then weighed to the nearest .01 g on a
Mettler electronic balance. Standard
length (SL) and sex were determined. The
specimens are deposited in the University
of Alabama Ichthyological Collection.
Aging to year class was determined by
EDITORIAL COMMITTEE FOR THIS PAPER:
Dr. David C. HEINS, Assistant Professor of Biology, Millsaps College,
Jackson, Mississippi 39210
Dr. Royal D. SUTTKUS, Professor of Biology and Director of Museum of
Natural History, Tulane University, New Orleans, Louisiana 70118
75
76
Tulane Studies in Zoology and Botany
Vol. 23
scale annuli. From 5 to 10 scales per fish
were analyzed to reduce the chance of
aging error due to the presence of false
annuli and regenerated scales. Aging to
month was accomplished by the technique
outlined by Page (1974). The following
symbols were used to indicate year
classes: -1, 0 to 12 months; 1 + , 13 to 24
months; 2 + , >24 months.
Reproductive condition and length of
the spawning season were determined by
field observations and by determination
of female gonosomatic indices (GSI). The
GSI is the ratio of gonad weight to
corrected body weight. Corrected body
weight is total weight minus the viscera
and gonad weight (Mathur and Ramsey,
1974).
Fecundity is defined as the number of
ova equal to or exceeding 0.2 mm
diameter. A large number of ova less than
0.2 mm diameter were present in each
ovary, but past studies of darters with
retracted spawning seasons (Fahy, 1954;
Winn, 1958a; Scalet, 1973) have suggested
that these minute oocytes never differen-
tiate into fully yolked, enlarged ova and
were, therefore, not spawned that year.
The smallest differentiating ova of the
larger egg group was 0.2 mm diameter, so
this size was used as the lower Hmit.
For food and feeding studies, whole
stomachs were removed, and the contents
were identified to family and enumerated.
RESULTS AND DISCUSSION
Habitat
Etheostoma coosae adults and juveniles
were consistently collected over rubble in
raceways and around boulders, near sand
bars and occasionally in the foot of rif-
fles. This habitat preference was main-
tained seasonally with no indication of
age or size specific habitat utilization for
foraging or reproduction.
The basis of habitat selection by darters
is influenced, if not determined, by phys-
iological and/or ecological requirements
of the species. Ultsch et al. (1978) con-
ducted a series of critical O2 experiments
with six species of Etheostoma and ob-
served that four ecological groups exist
with respect to oxygen requirements ver-
sus habitat selection. They suggested that
one such group, typified by E. (Ulocentra)
duryi and E. (Catonotus) flabellare,
preferred relatively fast water but main-
tained its ability to tolerate periods of
hypoxia. This group was the most diverse
physiologically in terms of oxygen use
strategies. As a result of this, these darters
maintained a diverse array of habitat
types. The applicability of this expla-
nation to habit selection by E. coosae Ues
in the close phylogenetic and ecological
relationships between it and E. duryi.
Demography
Etheostoma coosae was the dominant
percid species in Barbaree Creek. It com-
prised 5.9 percent of the total number of
fish specimens collected (Table 1). The
overall age class distribution of E. coosae
for the year studied is seen in Table 2.
Approximately 64 percent of the popula-
tion occupied the - 1 age class, 29 percent
the 1 + age class, and 7 percent the 2 +
age class.
Seasonal changes in age class compo-
sition (Figure 1) indicate that maximum
contribution to population size occurred
during winter in the - 1 age class as it
approached 12 months of age. From this
point, percent contribution to population
size declined throughout the older age
classes.
Of the 750 specimens examined, 32.9
percent and 55.6 percent of the males and
females, respectively, survived from the
- 1 to the 1-1- age class, whereas 10.8
percent and 10.3 percent of the males and
females respectively, survived from the
1 + to the 2-1- age class (Table 2).
The overall sex ratio, 1:1.3, was signifi-
cantly different (X ' = 9.86; p < .01) from
the expected 1:1. This skewed sex ratio
was most evident in the 1 + age class,
1:1.8 (X' = 18.96; p <.01), whereas the
- 1 and 2 -^ age class sex ratios were not
statistically different from 1:1.
Age and Growth
The oldest individuals collected, two
females and one male (Figure 2), were 36
No. 1 Etheostoma Life History 77
Table 1 . Percent relative abundance and frequency of occurrence of fishes collected in
Barbaree Creek from April 1977 through April 1978.
Species Abundance Occurrence
Family Cyprinidae
Campostoma anomalum
Notropis asperifrons
Notropis callistius
Notropis lirus
Notropis stilbius
Notropis trichroistius
Notropis venustus
Notropis xaenocephalus
Phenacobius catostomus
Semotilus atromaculatus
Family Catostomidae
Hypentelium etowanum
Moxostoma duquesnei
Family Ictaluridae
Ictalurus natal is 0.04 8.3
Family Centrarchidae
Ambloplites rupestris
Lepomis cyanellus
Lepomis gulosus
Lepomis macrochirus
Lepomis megalotis
Micropterus coosae
Micropterus punctulatus
Family Cyprinodontidae
Fundulus stellifer 0.02 25.0
Family Percidae
Etheostoma coosae
Etheostoma jordani
Etheostoma stigmaeum
Percina caprodes
Percina nigrofasciata
Family Cottidae
Cottus carolinae
Total
1.69
91.7
5.87
100.0
8.81
100.0
0.19
25.0
2.97
75.0
45.62
100.0
0.34
41.7
16.90
100.0
0.01
8.3
0.09
25.0
2.00
100.0
0.39
66.7
0.07
41.7
0.01
8.3
0.07
33.3
0.04
8.3
0.76
66.7
1.11
91.7
0.06
25.0
5.93
100.0
1.18
100.0
0.97
100.0
0.31
66.7
0.96
91.7
2.94
100.0
99.35
78
Tulane Studies in Zoology and Botany
Vol. 23
Table 2. Age-class distributions and survival of Etheostoma coosae collected in
Barbaree Creek from April 1977 through April 1978. S, and S2 equal survival
calculated from the - 1 and 1 + age classes, respectively.
Sex
Year
Number of
specimens
Surviva
1
class
S,
S,
-1
231
1.000
_
1 +
76
0.329
1.000
2 +
25
0.108
0.329
-1
252
1.000
—
1 +
140
0.556
1.000
2 +
26
0.103
0.185
-1
483
1.000
—
1 +
216
0.447
1.000
2 +
51
0.106
0.236
Males
Females
Combined
Sexes
months of age assuming a May hatching.
Each specimen had two annuli and the
third was in the process of being estab-
lished. Scale studies of E. coosae in
Barbaree Creek indicated that annuli are
established in early to middle spring.
Males grew faster than females, and
were on the average significantly longer (p
<.05) by the first spawning season
(Figure 3). Females attained 70.3 percent
and 90.0 percent of their average maxi-
mum standard length (40.5 mm) in 12 and
24 months, respectively, whereas males
Eq
_^a_^B_
M
SUMMER
Figure 1 . Seasonal changes in age-class composition
of Eiheosloma coosae collected in Barbaree Creek.
Age-class designations are: - 1 , 1 to 1 2 months; 1 -i- ,
13 to 24 months; and 2 + , >24 months.
attained 69.5 percent and 94.5 percent of
their average maximum standard length
(43.1 mm) in 12 and 24 months. The
longest male and female were 47.0 mm
and 44.1 mm, respectively (Figure 2).
There were no significant differences (p
>.05) between male and female length-
weight equations as tested by analysis of
covariance (Fg = 0.867, df = 1,136). The
relationship for combined sexes was log
wt. (grams) = 3.1657 log SL -5.1200, N
= 140, r = .978.
Page and Schemske (1978) stated that
one possible factor affecting size in
darters of the subgenus Catonotus was
interspecific competition. They concluded
that maximum size in sympatric popula-
tions of Catonotus was determined by
the presence or absence of Etheostoma
squamiceps. In the presence of E. squaini-
ceps, other Catonotus species were re-
duced in size, whereas in the absence of £.
sqamiceps their maximum size was
greater. They suggested that the major
function of the size differences was for
size-specific utilization of potential repro-
ductive habitats.
The overall growth trend of E. coosae
was for rapid growth, both sexes reach-
No. 1
Etheostoma Life History
79
ing approximately 70 percent of the aver-
age maximum standard length, the first
year of life, with a subsequent reduction
of this rate in later years (Figure 3). This
pattern is quite common in darters (Page,
1974 and 1975; Page and Burr, 1976;
Starnes, 1977) and in fishes generally. As
Ricker (1971) has pointed out, this phe-
nomenon is usually attributed to physio-
logical size limitations primarily influ-
enced by the heavier reproductive effort
by older individuals.
Reproduction
Female gonosomatic indices (Figure 4)
and field observations indicate that E.
coosae spawned from mid March through
early to mid May with peak spawning in
April. The spawning periods for species
of the subgenus Vlocentra are similar.
Winn (1958a, 1958b) reported that males
of Etheostoma sp. (Barren River form) in
Tennessee established territories near the
beginning of April, and spawning began
in about one week. Stiles (1975) reported
that E. simoterum spawned from early
E
E.
I
»-
O
z
lU
-*
O
et
<
O
z
<
^
A-<^
A
^
d^
A
A
A
o
A
o
OV
A
A
A
A
A
O
OA
A
O
A
o
o
Oft
A
A
A
OA
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
0
299
Mean
256.4
88.0
38.4
382.8
Percent
67.0
23.0
10.0
100.0
No. 1
Etheostoma I ife History
81
the anterior dorsal and ventral interradial
membranes of the caudal fin. The inter-
radial membranes throughout the length
of the spinous and soft dorsal fins had
rusty-red quadrate spots. Females main-
tained a ground color of light tan overlaid
by brown to black lateral blotches and
mottling above the lateral line.
The total egg complement increased
proportionally with length, r = .682 (Table
3). Fecundity studies of other darters have
substantiated positive size-fecundity rela-
tionships: E. squamiceps, r = .692 (Page,
1974); E. barbouri, r = 530 (Flynn and
Hoyt, 1979); E. kennicotti, r = .631
(Page, 1975); and Percina nigrofasciata, r
= .721 (Mathur, 1973). An exception to
this general relationship was reported for
E. proeliare (r <.l) by Burr and Page
(1978). This vagary was attributed to the
short life span (one year) of the popula-
tion studied, which yielded females of a
similar size.
Feeding
The overall diet of E. coosae, consisted
of 78 percent Diptera (Chironomidae and
Simuliidae), 12 percent Crustacea (Cope-
poda and Cladocera), 3 percent Ephemer-
optera (Baetidae and Siphlonuridae) and
5 percent miscellaneous items (Acarina,
Mollusca, Nematoda, Trichoptera, and
sand). The diet of various size classes as
well as the seasonal diet of combined age
and size classes is seen in Figure 5.
Etheostoma coosae consumed midge
larvae as juveniles and expanded their diet
as adults to include mayflies and caddis-
flies. Midge larvae decreased whereas
crusaceans increased in importance from
spring to winter. Mayflies, caddisflies,
and molluscs were important items during
summer months.
11-20mm
N = 6
21-30nnm
N = 78
31-40mi
N = 143
41-50mi
N = 25
jf, Misc
E|)h
SPRING
N=25
SUMMER
N=97
WINTER
N=34
Figure 5. Stomach contents of Etheostoma coosae collected in Barbaree Creek by size class of darter and
season collected. Seasonal analyses include all size and age classes. Food items are abbreviated as follows:
(Crus)tacea, (Dip)tera, (Eph)emeroptera, (Mis)ellaneous, (Mol)lusca, (Plec)optera, and (Tri)choptera. N
equals sample size.
82
Tulane Studies in Zoology and Botany
Vol. 23
The feeding mode of darters has been
reported to be largely selective in some
species: E. fonticola (Schenck and
Whiteside, 1977), E. nigrum (Roberts and
Winn, 1962), and E. radiosum cyanorum
(Scalel, 1972); and largely opportunistic
in others: E. acuticeps (Bryant, 1979), E.
blennioides (Fahy, 1954), and E. gracile
(Braasch and Smith, 1967). These papers
have illustrated that within the genus
Etheostoma feeding behaviors are quite
variable and complex. Based on the liter-
ature and my own studies, I believe that
feeding behavior is not so restrictive but
rather lies along a dynamic continuum
between selectivity and opportunism.
Species will adapt to prey abundance and
type assuming the most energetically re-
warding feeding response. Prey switching
as a possible behavioral mechanism
involved in feeding is supported by Ihe
studies of Murdoch et al. (1975) on
Poecilia reticulatua and Roberts and
Winn (1962) on the role of visual cues in
the feeding of £. nigrum.
ACKNOWLEDGEMENTS
1 wish to thank Dr. Herbert Boschung,
Dr. Maurice F. Mettee, and John Williams
who read and discussed various parts of
this paper; the USDA Forest Service for a
grant to Boschung from which this study
was funded; and finally Irene Thompson
who performed all typing services on the
manuscript.
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.
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December 30, 1981
84 Tulane Studies in Zoology and Botany Vol. 23
THE TAXONOMIC RELATIONSHIP BETWEEN MALACLEMYSGKM , 1844
AND GRAPTEMYS AGASSIZ, 1857 (TESTUDINES: EMYDIDAE)
JAMES L. DOBIE
Department of Zoology-Entomology
Auburn University, Alabama 36849
Abstract
The turtle genus Graptemys is a distinctive group
clearly separable from Malaclemys on the basis of
external and osteological features. The difference
between the groups indicate that the degree of
genetic relationship is no closer than that resulting
from their both having presumably arisen from a
Pseudemys - like stock or Malaclemys from a
Graptemys stock.
INTRODUCTION
Investigators of Malaclemys and Grapt-
emys have based their taxonomic alloca-
tions on penial, skull, shell, hind limb and
pelvic girdle morphology and on head
patterns. Osteological comparisons, when
indicated, were usually limited to the
skull, and in most cases, head patterns
were used to distinguish taxa. The degree
of evolutionary conservatism and paral-
leUsm exhibited by turtles argues against
the use of external characters (e.g., head
striping), alone in determining taxonomic
and phylogenetic relationships. Thus,
both osteological and surficial features
have been examined in this study.
HISTORICAL REVIEW
The controversy about the relationship
between Malaclemys and Graptemys be-
gan as a resuh of the lumping of Grapt-
emys with Malaclemys by Boulenger
(1889) and the re-establishment of the
genus Graptemys by Baur in 1890. Since
that time, W.P. Hay (1904) and O.P. Hay
(1908) followed Baur in recognizing the
two genera, as did Carr in 1949. Later,
however, Carr (1952) questioned the
validity of separating the two genera and
McDowell (1964), without presenting sup-
porting data, lumped Graptemys with
Malaclemys. Zug (1966, 1971), on the
basis of similiar penial, pelvic girdle, and
hind limb morphology for the two genera
considered them congeneric, and Parsons
(1960, 1968) found the choanal structures
of both genera to be so variable that the
evidence did not particularly support or
refute the congeneric idea. Several other
authors (Ernst and Barbour, 1972;
McKown, 1972; Dundee, 1974; Killebrew,
1979; Dobie and Jackson, 1979; Pritch-
ard, 1979; Vogt, 1978, 1980) have not
supported the synonymy of Graptemys
with Malaclemys; they evidently must
believe that sufficient evidence has not
been presented to lump the two genera
together.
The purpose of this study is to clarify
the generic status of Malaclemys and
Graptemys.
MATERIALS AND METHODS
Representatives of each of the ten ex-
tant Graptemys species (Vogt, 1980) and
their subspecies and individuals of several
subspecies of the monotypic Malaclemys
were examined. External features, includ-
Editorial Committee for this Paper:
Dr. Eugene S. GAFFNEY, Associate Curator, Department of Vertebrate
Paleontology, American Museum of Natural History, New York, New York 10024
Dr. John J. IVERSON, Assistant Professor of Biology, Earlham College,
Richmond, Indiana 47374
85
86
Tulane Studies in Zoology and Botany
Vol. 23
ng scute contracts, plastral patterns, and
striping on the head and leg were analyzed
in juvenile and adult turtles of both sexes.
Skull and shell characters were analyzed
on large sub-adult and adult females.
Skull terminology is that of Gaffney (1972
a); scute and bone terminology is that
used by Zangerl (1969).
The method used to elucidate the rela-
tionship between Malaclemys and Grapt-
emys and to other North American emy-
did turtles is the search for taxa that have
shared derived characters. This method
was described by Hennig (1966), and has
been used by others (Gaffney, 1972 b,
1975; W.E. Clark, 1978) and is called
phylogenetic systematics or cladism.
DIAGNOSTIC CHARACTERISTICS
The diagnostic characteristics of Grapt-
emys, Malaclemys and an outgroup com-
parison of those genera with the other
North American emydid genera are listed
in Table 1 . Each feature is also designated
as either ancestral (primitive) or advanced
(derived).
SIGNIFICANCE OF DIAGNOSTIC
CHARACTERISTICS
The number (s) in a bracket refers to
the number of the diagnostic features in
Table 1.
SKULL FEA TURES
(1) Quadratojugal - maxilla contact. If
the absence of contact between these two
bones represents the primitive state, then
the possession of the derived condition in
three Graptemys species (in one pseudo-
geographica and in all pulchra and bar-
bourij, in M. terrapin, and in some
Pseudemys species suggests that M.
terrapin could have been derived from
one of these Graptemys or Pseudemys
species. Graptemys could have come from
any group lacking contact between the
two bones.
(2) Spoon-shaped symphysis of lower jaw
(Fig. 1). The flattened spoon-shaped
nature of the symphyseal part of the lower
jaw apparently is a derived feature in
Graptemys. The absence of such a struc-
ture in Malaclemys suggests that Grapt-
emys was not ancestral to Malaclemys and
that Malaclemys may have arisen from
some Pseudemys species.
Figure 1. Shape of the symphyseal area of the
lower jaw in mature females of (A) Malaclemys ter-
rapin. (B) Graptemys pseudogeographica, (C) G.
geographica, (D) G. pulchra, (E) G. barbouri, (?) G.
caglei, (G) G. versa, (H) G. ouachitensis sabinensis,
(I) G. o. ouachitensis, and (J) G. flavimaculata (the
shape of the symphysis is the same for
flavimaculata, oculifera, and nigrinoda).
No. 1
Malaclemys- Graptemys Relationship
87
(3) Bones surrounding the foramen pala-
tinum posterius (Fig. 2). The bones sur-
rounding that foramen in Terrapene and
in the species of the Pseudemys rubriven-
tris complex are the same as Graptemys;
the other species of Pseudemys and the
other N.A. emydid genera are like Mala-
clemys. Therefore, Graptemys and Mala-
clemys were possibly derived from differ-
ent species of Pseudemys.
(4) The absence of contact between the
ophisthotic and pterygoid due to the in-
volvement of the exoccipital. If the condi-
tion in Malaclemys and Deirochelys repre-
sents a derived feature, this would strong-
ly suggest that Malaclemys was not the
ancestral stock from which Graptemys
evolved. It could also indicate that a
Graptemys, Deirochelys, or any other
species of North American emydid turtle
could have been ancestral to Malaclemys.
(5) The lack of a notch in the premaxil-
lary bones. The lack of a notch in those
bones in Graptemys and the presence of a
notch in Malaclemys and the rest of the
N.A. emydids, precludes determination
of the possible ancestor for Graptemys
and Malaclemys based on this feature.
SHELL FEA TURES
(6) Flaring of carapace. The presence of
such in Graptemys and to varying degrees
in all other N.A. emydids except Mala-
clemys and some Terrapene, may indicate
that flaring is an ancestral feature. If so,
the upturning of the carapace in the last
two genera would be a derived feature.
This implies that Graptemys did not come
from a Malaclemys stock.
(7) Double notching of some peripherals.
The double notching of some of the per-
ipherals is found only in Graptemys and
in some individuals of Pseudemys scripta
and P. concinna. This could indicate that
Graptemys was not ancestral to Mala-
clemys and that a Pseudemys species was
ancestral to Graptemys.
(8 and 9) The keel and its associated
bosses (Fig. 3). A number of reports have
dealt with the extent and development of
the keel in Malaclemys. The last vertebral
scute is variable with respect to keel devel-
opment. Say (1825) reported that the last
vertebral in M. terrapin centrata was un-
keeled; Wied (1865) noted that all of the
vertebrals of M. t. pileata have a well
developed keel. The keel in Malaclemys t.
centrata was stated by W.P. Hay (1904)
"to be rather low and rounded," whereas
it was "always well developed," in M. t.
macrospilota. A keel is thus not always
present on the last vertebral, and I have
not observed the end of the keel (the fifth
boss area) to extend more than four-fifths
the length of the last vertebral scute. W.P.
Hay's (1904) statement about the keel and
bosses of M. t. littoralis was: "the first
vertebral plate is raised on the middle line
to form a broad, low carina; on the
second plate the elevation is greater, and
stands out as a smooth boss . . . ; the
elevation on the third plate has the form
of a hemispherical button with a well-
marked constriction around the posterior
half of the base . . . ; on the fourth plate
the elevation is raised into a knob-like
protuberance from a base which is con-
stricted all around . . . ; the fifth
vertebral plate is flat or with only a trace
of an elevation." Thus Hay's statement
suggests that four or five bosses are
present on the keel in Malaclemys. This is
not always the case. Auburn University
Museum of Paleontology (AUMP) speci-
men 2179 has only three bosses, and its
shell structures are normal.
Concerning the total number of bosses
on the keel in Graptemys pulchra, Carr
and Coin (1955) said, "the dorsal keel
. . . comprises a boss on each of the first
four centrals, . . . weak to nearly lacking
on the first and completely lacking on the
fifth." A boss on the fifth central (verte-
bral) is not lacking \n pulchra. Although it
is not prominent in G. pulchra or in any
other species of Graptemys, a terminal
boss can be detected in all species. Cagle
(1954), p. 182, Fig. 11) illustrated a
juvenile G. flavimaculata that had five
bosses on the carapace. I have never
examined any specimen of Graptemys,
including G. flavimaculata, in which the
88
Tulane Studies in Zoology and Botany
Vol. 23
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No. 1
Malaclemys- Graptemys Relationship
89
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No. 1
Malaclemys- Graptemys Relationship
91
Figure 2. The location of the foramen palatinum posterius. The foramen is bounded on its mediolateral
and outer lateral sides by the palatine in Graptemys (A) pseudogeographica, (B) geographica, (C) pulchra,
(D) barbouri, (E) eaglet and versa, (F) o. sabinensis, (G) o. ouachitensis and (H) nigrinoda, oculifera, and
flavimaculala. It is bounded on its mediolateral and outer sides by the palatine and maxilla, respectively, in
Malaclemys terrapin (I). Palatine (p). Maxilla (m). Foramen (0- Vomer (v). Pterygoid (pt).
92
Tulane Studies in Zoology and Botany
Vol. 23
fifth boss was located in the position il-
lustrated by Cagle; the fifth boss is always
at the posterior end of the last vertebral
scute. The similar location of each boss in
Graptemys and Malaclemys indicates
their close relationship.
(10) Amount of ventrolateral extension of
the nuchal bone and the costiform process
of the nuchal bone. Graptemys normally
lacks a costiform process; Malaclemys has
one. Even though the nuchal of Grapt-
emys is as wide as the same bone in Mala-
clemys, the distance the nuchal extends
ventrolaterally is less in Graptemys than
in Malaclemys. Therefore, the degree of
such extension must not be solely a func-
tion of the width of the nuchal bone. This
Nu
seems to be the case since the distal width
of the first peripheral is proportionately
greater in Graptemys than in Malaclemys.
Therefore, the presence of a narrower
first peripheral and a costiform process in
Malaclemys results in a greater ventro-
lateral extension of the nuchal in that
genus than in Graptemys.
The other North American emydids
that have a costiform process are Pseud-
emys, Terrapene, some Clemmys and
Deirochelys, and the latter genus is the
only group that has a ventrolateral exten-
sion similar to that of Malaclemys. I think
it unlikely that Deirochelys was ancestral
to Malaclemys; therefore, perhaps some
Pseudemys turtle was the stock from
which Malaclemys arose. The ancestral
stock for Graptemys can not be determi-
ned with respect to this feature.
(11 and 12) The notching of the postero-
lateral borders of the nuchal bone and the
anterior border of the costal bone (Figs. 4
and 5). The presence of such notching in
Graptemys, Terrapene and in most
Clemmys (14 of 16), Pseudemys (29 of
31), and Chrysemys (15 of 20), and not in
Malaclemys (except in one specimen),
Emydoidea, and most Deirochelys sug-
gests that Malaclemys was not ancestral to
Figure 3. The location of the bosses in Graptemys
(A) pulchra, (B) nigrinoda, and (C) Malaclemys
lerrapin and the contact of the eighth costal with the
seventh neural in some G. pulchra due to the loss of
the eighth neural bone. The normal contact is
between eighth costal and eighth neural in Grapt-
emys and eighth costal and seventh and eight neurals
in Malaclemys. Nuchal bone (Nu). Bosses )B 1-5).
Neural bones (N 1-8). Suprapygal bones (S 1-2). Py-
gal bone (P). Costal bones (C 1-8).
Figure 4. Dorsal view of the nuchal bone in
Graptemys (A) pseudogeographica, (B) pulchra and
(C and D) Malaclemys terrapin. Arrows indicate
notches. The position of the anteromedial edge of
the first pleural scute and the anterolateral borders
of the first vertebral scute are not on the nuchal bone
in some Malaclemys (D).
No. I
Malaclemys- Graptemys Relationship
93
Graptemys if the absence of notching is a
derived feature. However, Graptemys
could have given rise to Malaclemys, as
could have Clemmys, Chrysemys, Pseud-
emys, Terrapene, Emydoidea, and Deiro-
chelys. Emydoidea and Deirochelys pre-
sumably would be the best candidates
from which to derive Malaclemys if rela-
tionships are based on the presence of
shared derived features. In spite of the
presence of a shared derived feature
between those genera and Malaclemys, I
do not believe that either one is a good
candidate for being the progenitor of
Malaclemys. Therefore, Graptemys,
Pseudemys, and Chrysemys are
considered to be more likely candidates.
(13 and 14) The amount of pleural scute
overlap on the nuchal bone and first
vertebral scute - nuchal bone relation-
ships. A great deal of pleural scute over-
lap exists in Graptemys, Pseudemys, and
Figure 5. Dorsal view of the first left costal bone in
Graptemys (A) pseudogeographica, (B) pulchra and
(C and D) Malaclemys terrapin. That part of the an-
terior borde: of the costal bone that would adjoin
the nuchal generally is straight and unnotched in
Malaclemys as in (D). Arrows indicate notches.
in some Terrapene and the pleural scute
always contacts the margin of the first
vertebral scute on the nuchal bone in the
first two of the the three above (Dobie
and Jackson, 1979). Malaclemys resem-
bles rriost Chrysemys and some Terra-
pene, Clemmys, and Deirochelys in that
there is little overlap of the pleural scute
on the nuchal and the pleural scute does
not always contact the first vertebral scute
on the nuchal bone (Dobie and Jackson,
1979).
Malaclemys terrapin could have
evolved from Chrystemys in which the
extent of pleural scute overlap was mini-
mal and the margin of the first vertebral
scute did not always meet the pleural scute
on the nuchal bone. If M. terrapin evolved
from any species of Graptemys or Pseud-
emys that had a large amount of pleural
scute overlap and contact between the two
scutes on the nuchal bone, then presum-
ably a reduction in the amount of pleural
scute overlap must have occurred. Grapt-
emys could have arisen from a Pseudemys
stock.
(15 and 16) Amount of nuchal scute over-
lap and underlap and the width-length
relationships of the underlap part of the
nuchal scute (Figs. 6 and 7). The amount
of nuchal scute overlap is small in Mala-
clemys, in some Terrapene, and in all
extant species of Graptemys, except G.
geographica (Dobie and Jackson, 1979).
Both Malaclemys and Graptemys have
smaller amounts of nuchal scute underlap
than any other North American emydid
turtle, and the distal width of the under-
lap part of the nuchal scute is broader
than its length in both of those genera and
in some Pseudemys and Terrapene (Dobie
and Jackson, 1979). Based on these fea-
tures, Malaclemys would seem to be more
closely related to Graptemys than to any
other extant North American emydid
genus.
(17) Contact of the eighth costal. bone
with the seventh and eighth neurals (Fig.
3). The presence of such contacts in Mala-
clemys and the contact of the eighth costal
with only the eighth neural in Graptemys
94
Tulane Studies in Zoology and Botany
Vol. 23
(except for a single population of G.
pulchra) and in all other North American
emydid genera except Terrapene (the
eighth neural is absent in some Terra-
pene } indicates that contact with the
seventh neural is a derived character. The
stock from which Malaclemys was derived
presumably could have been any genus of
North American emydid turtles; Grapt-
emys could have come from Pseudemys
or from any other North American
emydid genus except Malaclemys.
(18) Lateral ridges on undersides of first
and fifth costals (Fig. 8). The lateral
ridges extending toward the midline of the
carapace from the anterior and posterior
ends of the bridge are well developed in
Graptemys in constrast to those of Mala-
clemys and the rest of the North Ameri-
can emydid genera. The functional sig-
nificance of those ridges is not known but
they may serve as supportive units for the
carapace. Malaclemys and Graptemys
presumably could have been derived from
any one of those genera.
(19) Distal widths of the three widest
costal bones. An attempt to indicate the
degree of relationships of Malaclemys to
any other emydid genus on the basis of
this character would be impractical
because of the extremely variable nature
of the widths of the costal bones. The
fairly consistent widths in the species of
Graptemys does indicate that they are
closely related.
(20) Sculpturing on the carapace. The
sculpturing on the carapacial bones in
Graptemys is similar to that of some
species of Pseudemys (P. floridana and P.
concinna) although the degree of sculp-
26|-
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23
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21
20
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o Maiaeltmyt (II)
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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
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75
70
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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
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Tulane Studies in Zoology and Botany
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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-
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DOBIE, J.L. and D.R. JACKSON. 1979. First
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December 30, 1981
ADDENDUM
Tulane Studies in Zoology and Botany Volume 23, number I
The addendum below is a continuation of the left hand paragraph of page 101 in the article
by Dobie. It ends where the right hand paragraph begins LITERATURE CITED.
I'viU -OOL
LIBRARY
OEC 201982
HARVAKD
UNIVIRRSITY
p. stejnegeri: (5) (AUMP 2363, four uncatalogued
specimens).
Rhinodemmys areolata: (1) (AUMP 2111).
R. pukherrima: (2) (AUMP 910, uncatalogued
specimen).
R. unidentified species: (1) (AUMP 2299).
Terrapene ornata: (AUM 10732); (AUMP 122-23,
962, 1939).
T. Carolina: (52) (AUM 551, 1394. 1899, 3909-11,
4998, 5925, 8866, 9414, 11611, 14295, 17634, 20942.
23851, 25096-97); (AUMP 116-20, 124, 128, 130-31,
136-42, 702, 712, 914-16, 2250, 2257, 2312, 2317):
(UF/FSM 7570, 14204, 35023, 38341, 40388,
41508-09, 41518, 41521-22).
Unidentified genus and species: (1) (SDSM & T
59187).
Specimens came from the following collections:
Amerian Museum of Natural History (AMNH);
Auburn University Museum (AUM); Auburn Uni-
versity Museum of Paleontology (AUMP); Carnegie
Museum (CM); Field Mueum of Natural History
(FMNH); University of Kansas Museum of Natural
History (KU); The Vertebrate Museum, Northeast
Louisiana State College (NLSC); South Dakota
School of Mines and Technology (SDSM & T);
Stephen F. Austin State University Vertebrate
Collection (SFA); State University of South Dakota
(SUSD); Texas Cooperative Wildlife Collection,
Texas A&M University (TCWC); Texas Natural
History Collection, Austin (TNHC); Tulane Uni-
versity Museum (TU); University of Florida, Florida
State Museum (UF/FS); Museum of Southwestern
Biology, The University of New Mexico (UNM);
University of South Alabama (USA); United States
Museum of Natural History, Smithsonian Institu-
tion (USNM).
iaai> UU0Z.-O/0Z.
T&-
Volume23 . Number 2 $3.50 ^ ,, .D^eember 15, 1982
OtC2 0ii^82
MARvArxD
I lMI\/irR~lTY
CHANGES IN MELANIN MIGRATION INDUCED BY NORADRENERGIC
AND HISTAMINERGIC AGENTS IN THE FIDDLER CRAB, UCA PUGILA TOR
MUKUND M. HANUMANTE AND MILTON FINGERMAN p. 103
ADDITIONAL TREMATODES OF MAMMALS IN LOUISIANA
WITH A COMPILATION OF ALL TREMATODES REPORTED FROM
WILD AND DOMESTIC MAMMALS IN THE STATE
WESLEY L. SHOOP AND KENNETH C. CORKUM p. 109
COMPARATIVE VISCERAL TOPOGRAPHY OF THE
NEW WORLD SNAKE TRIBE
THAMNOPHIINI (COLUBRIDAE, NATRICINAE)
NITA J. ROSSMAN , DOUGLAS A. ROSSMAN
and
NANCY K. KEITH P- 123
TULANE UNIVERSITY
NEW ORLEANS
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TULANE STUDIES IN ZOOLOGY AND BOTANY
Volume 23, Number 2
December 15, 1982
CHANGES IN MELANIN MIGRATION INDUCED BY NORADRENERGIC
AND HISTAMINERGIC AGENTS IN THE FIDDLER CRAB, UCA PUGILA TOR*
MUKUND M. HANUMANTE AND MILTON FINGERMAN
Department of Biology, Tulane University
New Orleans, Louisiana 70118 U.S.A.
Abstract
The effects of the H, receptor blocker SA-97, the
Hi receptor blocker cimetidine, the tyrosine hydroxy-
lase inhibitor a -methyl-para-tyrosine and the H,
receptor and norepinephrine uptakci blocker diphen-
hydramine on histamine- or 4-methyl histamine-in-
duced inhibition of melanin dispersion in the fiddler
crab, Uca pugilator undergoing a background trans-
fer from white to black were determined. Only cimeti-
dine significantly antagonized the 4-methyl histamine-
evoked decrease in melanin dispersion. a-Methyl-
para-tyroslne by itself significantly diminished where-
as diphenhydramine by itself significantly potentiated
the amount of this centrifugal melanin migration in
the fiddler crabs. None of these drugs affected
melanin migration in vitro. The results are consistent
with the hypotheses that norepinephrine triggers re-
lease of a melanin-dispersing hormone and that H, re-
ceptor activation decreases impulse-mediated nore-
pinephrine release in this crab.
INTRODUCTION
Translocation of the melanin in the
melanophores of the fiddler crab, Uca
pugilator, is regulated by antagonistic
neurohormones, a melanin-dispersing
hormone (MDH) and a melanin-concen-
trating hormone (Carlson, 1935; Sandeen,
1950; Fingerman, 1956). Norepinephrine
(NE) triggers release of MDH in this crab
(Fingerman et al., 1981; Hanumante and
Fingerman, 1981a,b; 1982a,b,c; Hanu-
mante et al., 1981). Recently histamine
(HA) has been shown to inhibit melanin
♦Supported by Grant PCM-8 1-08864 from the
National Science Foundation.
dispersion in a dose-dependent manner
(Hanumante and Fingerman, 1981b). Use
of a variety of histaminergic agonists and
antagonists led to the hypothesis that two
types of HA receptors, called H, and H2,
are present on NE neurons that trigger
MDH release and that HA exerts its inhibi-
tory action by stimulating the H2 recep-
tors. The present investigation was devised
to obtain further support for this
hypothesis. This objective was carried out
by observing the effects of specific mam-
malian histaminergic and noradrenergic
agents not used previously on the inhibi-
tory action of HA and 4-methyl histamine
(4-MeHA; a selective H2 receptor agonist,
Owen et al., 1979; Douglas, 1980; Polanin
et al., 1981) on melanin dispersion in Uca
pugilator transferred from a white to a
black background.
Materials and Methods
Adult male fiddler crabs, Uca pugilator,
from the vicinity of Panacea, Florida,
(Gulf Specimen Company) were used.
Their melanophores were staged according
to the system of Hogben and Slome (1931)
whereby stage 1 .0 represents maximal pig-
ment concentration, stage 5.0 maximal
pigment dispersion and stages 2.0, 3.0, and
4.0 the intermediate conditions. When
intact crabs were used, the melanophores
seen through the cuticle on the anteroven-
tral surface of the second walking leg on
the right side were staged at the time a sub-
EDITORIAL COMMITTEE FOR THIS PAPER:
DR. RAY W. FULLER, Research Advisor, Eli Lilly and Company, Indianapolis,
Indiana 46206
DR. WILLIAM S. HERMAN, Professor and Head, Department of Genetics and
Cell Biology, University of Minnesota, MinneapoHs, Minnesota 55108
103
104
Tulane Studies in Zoology and Botany
Vol. 23
stance was injected and 15, 30, 60, 90, and
120 minutes thereafter. To facilitate com-
parison of the responses of the experimen-
tal and control crabs, mean differences
between the 15 through 120 minute
melanophore stages for the control and ex-
perimental groups were calculated for use
in Table 1 . The depicted data are based on
the mean melanophore stages of 20 intact
crabs (10 experimental and 10 control) or
20 isolated legs (10 experimental and 10
control). When assays were performed on
isolated legs, the melanophores were
staged only at the time the legs were
removed from the crab (at which time the
legs were perfused with the test or control
solution) and 15, 30, 45, and 60 minutes
thereafter. The second and third walking
legs from both sides of the crab were
removed; the legs from the right served as
experimentals and the legs from the left
side received control solution; the melano-
phores on the anteroventral surface of
these isolated legs were observed for
staging. The assays were performed using
isolated legs having initially either maxi-
mally concentrated melanin (stage 1.0) or
maximally dispersed melanin (stage 5.0).
Melanophores in isolated legs of this crab
remain responsive for at least 120 minutes
(Herman and Dallmann, 1975). The statis-
tical significance of the data was deter-
mined using Standard Errors of the Means
(SEM) the Student's t test with significance
set at the 95% confidence interval. None
of the data for isolated legs were statis-
tically significant.
The volume of the solution injected into
each crab or isolated leg was always 0.05
ml. The experiments with intact crabs and
isolated legs were performed at 24 °C under
an illumination of 1190 Ix. 4-MeHA dihy-
drochloride (Smith, Khne and French),
cimetidine (N"-Cyano-N-methyl-N'-{2-
(5-methylimidazol-4-yl) methylthioethyl}
guanidine) (Smith, Kline and French) and
SA-97 (homochlorcyclizine) (Eisai) were
generous gifts. In addition, HA, amethyl-
para-tyrosine (a-MPT) and diphenhydra-
mine HCl (all from Sigma) were used. The
concentration used for each drug, whether
injected alone or in combination, was 20
ug/dose of the free compound. All drugs
except cimetidine were dissolved in
Pantin's physiological saline (Pantin,
1934). Cimetidine was dissolved in acidi-
fied (a drop of 1,2 M HCl) saline. Conse-
quently, a drop of HCl (1.2 M) was added
to control saline for the cimetidine exper-
iments. The rest of the controls received
pure saline.
Results and Discussion
4-MeHA, an H2 receptor agonist,
slowed the rate of melanin dispersion, as
observed earlier by Hanumante and Fin-
german (1981b), in intact crabs transferred
from a white to a black background (Table
1). Cimetidine, which selectively blocks
mammalian H2 receptors (Douglas, 1980;
Polanin and McNeill, 1981) significantly
antagonized the 4-MeHA. On the other
hand, the H, receptor blocker SA-97 not
only did not antagonize the 4-MeHA but
the combination of 4-MeHA plus SA-97
resulted in significantly further inhibition.
None of these drugs affect melanin migra-
tion in vitro nor do SA-97 and cimetidine
by themselves have an effect on the rate of
melanin dispersion in crabs undergoing a
background change from white to black
(Hanumante and Fingerman, 1981b), a
black background fostering melanin dis-
persion (Brown and Hines, 1952) which
will be effected by MDH.
a-MPT selectively inhibits tyrosine
hydroxylase. This enzyme catalyzes the
synthesis of dihydroxyphenylalanine from
tyrosine. At least in mammals this is the
rate-limiting step in the biosynthesis of
NE (Terrasawa et al., 1975; Lofstrom and
Backstrom, 1978). a MPT by itself signi-
ficantly decreased melanin dispersion.
HA by itself, as reported earlier
(Hanumante and Fingerman, 1981b), sig-
nificantly reduced centrifugal melanin
migration in intact crabs transferred from
a white to a black background. However,
in the crabs that were co-administered
either 4-MeHA and a-MPT or HA and
a-MPT (Table 1), 4-MeHA and HA were
not able to produce further, significant
No. 2
Melanin Migration in Crabs
105
reduction of the melanin dispersion.
Diphenhydramine, a blocker of H, recep-
tors and NE uptake, in mammals (Isaac
and Goth, 1965; Fantozzi et al., 1975;
Marco et al., 1980), by itself significantly
enhanced melanin dispersion. However,
when HA was co-administered with
diphenhydramine, the HA-induced inhi-
bition in melanin dispersion was still
evident (Fig. 1).
The present data, in light of our earlier
report (Hanumante and Fingerman,
1981b) and the pharmacological actions
of noradrenergic and histaminergic agents
in mammals, further strengthen the hypo-
thesis that (a) NE serves as a neurotrans-
mitter triggering release of MDH and that
(b) activation of H2 receptors located on
NE neurons which control MDH release
results in a decrement of melanin disper-
sion in Uca pugilator transferred from a
white to a black background. The
observations that cimetidine, a selective
H2 receptor blocker, antagonized the
4-MeHA-induced inhibition in melanin
dispersion, whereas the Hi blocker SA-97
did not, reveal that this effect is mediated
specifically by activation of HA H2 recep-
tors. The marked increase in inhibitory
effect of 4-MeHA when co-administered
with the Hi antagonist SA-97 was
probably due to the fact that excitation of
H, receptors evokes enhanced melanin
dispersion (Hanumante and Fingerman,
1981b), blocking them would prevent any
endogenous Hi stimulation of the crabs.
This would enable 4-MeHA, an agonist of
H2 receptors, to produce an even greater
inhibition of the melanin dispersion. On
the contrary, in the crabs whose H2 recep-
tors were blocked by cimetidine, 4-MeHA
was unable to significantly decrease the
action potential-mediated release of NE,
which in turn resulted in a near normal
quantity of MDH being released into the
hemolymph of these crabs transferred to
the black background. The fact that
metiamide, another H2 receptor blocker,
significantly antagonized the 4-MeHA-
stimulated decrease in centrifugal melanin
migration (Hanumante and Fingerman,
1981b) in vivo further strengthens this
conclusion.
NE has been found (0.51 pg/g) in the
supraesophageal ganglia of male fiddler
crabs (Hanumante and Fingerman,
1982b). Also, we have provided evidence
that Hi and H2 receptors occur on NE
neurons because in fiddler crabs pretreated
with 6-hydroxydopamine (which
presumably destroys NE neuroterminals in
Uca as it does in vertebrates) (Hanumante
and Fingerman, 1982b,c) HA is unable to
significantly reduce further the melanin
dispersion (Hanumante and Fingerman
1981b). We have not determined (i) the
levels of NE in a-MPT injected crabs or (ii)
the exact mechanism of action of a-MPT
in Uca puilator. However, data that we
obtained using noradrenergic and histadre-
nergic agents (Hanuamante and Finger-
man, 1981b) reveal that 20 MPT clearly
interferes with NE neurotransmission.
This probably was either by way of its well-
established (at least in mammals) pharma-
cological NE synthesis-inhibiting effect
(Terraswawa et al., 1975; Lofstrom and
HOURS
Figure 1. Relationships between melanophore stage
and time. Circles with bottom-half darkened, crabs
that received diphenhydramine; circles with top-half
darkened, crabs that received histamine; solid circles,
crabs that received histamine plus diphenhydramine;
open circles, salme-injected controls. Vertical bars
indicate SEM.
106
Tulane Studies in Zoology and Botany
Vol. 23
Table i . The means ( ± SEM) of the differences between the melanophore stages de-
termined at 15, 30, 60, 90, and 120 minutes of the intact crabs that received a
drug versus the saline-injected controls. The minus sign indicates decreased
melanin dispersion relative to the controls. *Statistically significant p^ .05
relative to respective controls.
4-Methyl histamine (4-MeHA)
Cimetidine
4-MeHA plus cimetidine
4-MeHA plus SA-97
a Methyl-p-Tyrosine (a-MPT)
4-MeHA plus a-MPT
Histamine (HA)
HA plus a-MPT
-0.67*(± 0.08)
-0.17 (± 0.01)
-0.39 (± 0.07)
-1.43*(± 0.12)
-1.15*(± 0.15)
-1.44*(± 0.21)
-1.18*(± 0.18)
-1.01*(± 0.12)
Backstrom, 1978; Douglas, 1980) or by
stimulating H2 receptors, thereby leading
to the observed decrement in MDH release
(Table 1). Hence, the melanin of these
a -MPT-treated crabs did not disperse to
the extent it did in the control animals.
As stated above, in the crabs co-injected
with 4-MeHA and a-MPT or HA and
a-MPT, neither 4-MeHA nor HA signifi-
cantly affected the melanin dispersion
compared with that which occurred in re-
sponse to a-MPT alone (Table 1). This pre-
sumably was due to the interference with
NE neurons by a-MPT in such a way that
the impulse-mediated decrement in NE
secretion by the H2 stimulators 4-MeHA
and HA was not large enough to affect
significantly the NE-mediated MDH
release.
The diphenhydramine-evoked increment
in melanin dispersion (Fig. 1) was pre-
sumably due to its blocking action on NE
uptake, (Marco et al., 1980). NE uptake,
inhibitors like nisoxetine (Koe, 1976) have
already been shown to potentiate MDH
release (Hanumante and Fingerman,
1981a). Diphenhydramine antagonizes H,
receptors (Isaac and Goth, 1965; Fantozzi
et al., 1975; Marco et al., 1980) also. How-
ever, because H, receptor blockers do not
significantly abolish HA- or 4-MeHA- (an
H: receptor agonist) mediated inhibition of
melanin dispersion, we suggest that the NE
uptake, blocking action of diphenhydra-
mme is responsible for the potentiation of
melanin dispersion. The observation that
even when HA is co-administered with di-
phenhydramine there is still a decrease in
melanin dispersion (Fig. 1) indicates that
HA does not evoke its effect by stimulating
NE uptake,; uptake, being the major
mechanism of inactivating the postsyn-
aptic actions of monoamines including NE
(Fuller and Wong, 1977). That none of
these drugs affect significantly melanin
migration in isolated legs (Hanumante and
Fingerman, 1981b) is consistent with the
hypothesis that these drugs elicit changes
in melanin dispersion indirectly by inter-
acting with the neuroendocrine system of
Uca pugilator.
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. 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.
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Marco, E.J., G. Balfagon, J. Marin,
B. Gomez, and S. LLUCH. 1980. indirect
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Owen, D.A.A., C.A. Harvey, and R.W.
GRESTWOOD. 1979. Cardiovascular studies with
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PANTIN, C.P.A. 1934. The excitation of crusta-
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T.E. Tenner, jr., and J.H.
McNEILL. 1981. The characterization of cardiac
histaminergic chronotropic receptors in the rabbit.
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SANDEEN, M.I. 1950. Chromatophorotropins in
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108
Tulane Studies in Zoology and Botany Vol . 23
ADDITIONAL TREMATODES OF MAMMALS IN LOUISIANA
WITH A COMPILATION OF ALL TREMATODES REPORTED FROM
WILD AND DOMESTIC MAMMALS IN THE STATE
WESLEY L. SHOOP AND KENNETH C. CORKUM
Department of Zoology and Physiology, Louisiana State University
Baton Rouge, Louisiana 70803
Abstract
The following trematodes were collected from
hunter-trapped mammals in the Atchafalaya basin of
Louisiana during the winters of 1981 and 1982: Alaria
alarioides (Dubois, 1937) Dubois, 1970 from mink,
Mustela vison Schreber, and river otter, Lutra cana-
densis (Schreber); Alaria marcianae (La Rue, 1917)
Walton, 1949 from raccoon, Procyon lotor (Linn.)
and bobcat, Lynx rufus (Schreber); Alaria mustelae
Bosma, 1931 from raccoon and mink; Amphimerus
speciosus (Stiles and Hassal, 1896) Barker, 1911 from
raccoon and the domestic cat. Fells domesticus Linn.;
Baschklrovitrema incrassatum (Dies., 1850) Skrjabin,
1944 from mink and river otter; Brachylaima virgin-
iana Dickerson, 1930 from opossum, Dldelphis vir-
giniana Kerr; Carneophallus basodactylophallus
Bridgman, 1969 from raccoon; Cryptocotyle concava
(Creplin, 1825) Lube, 1899 from mink; Fibricola
cratera (Barker and Noll, 1915) Dubois, 1932 from
mink, opossum, and raccoon; F. lucida (La Rue and
Bosma, 1927) Dubois and Rausch, 1950 from mink
and opossum; Gyrosoma stngulare Byrd, Bogitsh,
and Maples, 1%1 from raccoon and mink; Hasstllesia
texensis Chandler, 1929 from muskrat. Ondatra zibe-
thica (Linn.); Heterobllharzia americana Price, 1929
from mink, raccoon, and bobcat; Isthmiophora mells
(Schrank, 1788) Luhe, 1909 from raccoon and mink;
Linstowiella szldati (Anderson, 1944) Anderson and
Cable, 1950 from opossum and raccoon; Marltremtn-
oides nettae (Gower, 1938) Rankin, 1939 from rac-
coon and mink; Microphallus opacus (Ward, 1894)
Ward, 1901 from raccoon and mink; Paragonlmus
kellkottl Ward, 1908 from opossum; Pharyngosto-
moldes procyonis Harkema, 1942 from raccoon;
Quinqueserialis qulnqueserialis (Barker and Laughlin,
1911) Harwood, 1939 from muskrat; Rhopalias ma-
cracanthus Chandler, 1932 from opossum; and Sella-
cotyle vitellosa Sogandares-Bernal, 1961 from mink.
Alaria alarioides, A. marcianae, Amphimerus spe-
ciosus, Cryptocotyle concava, Isthmiophora mells.
Microphallus opacus, Paragonlmus kellkottl, and
Qulnqueserialis qulnqueserialis have not been pre-
viously reported from Louisiana mammals. Diag-
noses are presented for the species representing state
records along with pertinent notes on the biology of
each. New host records include Heterobllharzia
americana, Cryptocotyle concava, and Maritremi-
noides nettae from mink; Alaria marcianae, Amphi-
merus speciosus, and Linstowiella szldati from rac-
coon; and Hasstllesia texensis from muskrat. A com-
pilation of trematodes previously reported from Loui-
siana mammals is presented.
INTRODUCTION
Recently, we reported some trematodes
collected from mammals in south Louisi-
ana (Shoop and Corkum, 1981a). Since
that time we have continued our examina-
tion of hunter-trapped mammals from the
Atchafalaya basin of Louisiana during the
winters of 1981 and 1982. The following
mammals were examined for trematodes:
42 minks, Mustela vison Schreber; 37 rac-
coons, Procyon lotor (Linn.); seven river
otters, Lutra canadensis (Schreber); five
muskrats, Ondatra zibethica (Linn.); three
bobcats, Lynx rufus (Schreber); four
domestic cats, Felis domesticus Linn.; two
opossums, Dldelphis virginiana Kerr; and
three red foxes, Vulpes fulva (Desmarest).
The red foxes were found uninfected with
trematodes.
Trematodes were fixed in steaming 10%
EDITORIAL COMMITTEE FOR THIS PAPER:
DR. BERT B. BABERO, Professor of Biological Sciences, University of Nevada,
Las Vegas, Las Vegas, Nevada 89154
DR. WALTER E. WILHELM, Associate Professor of Biology, Memphis State
University, Memphis, Tennessee 38152
109
no
Tulane Studies in Zoology and Botany
Vol. 23
formalin and stained in Semichon's aceto-
carmine. All measurements are in micro-
meters unless otherwise stated; means are
followed by the ranges in parentheses. Line
drawings were prepared with the aid of a
microprojector. Representative specimens
of the species for which diagnoses are
given were deposited in the Manter Lab-
oratory, University of Nebraska State
Museum, Lincoln, Nebraska.
Table I lists the trematodes recovered
from the eight species of mammals.
Lumsden and Zischke (1961) reported and
diagnosed Fibricola cratera, F. lucida,
Hasstilesia texensis, Brachylaima virgin-
iana, and Rhopalias macracanthus from
Louisiana mammals. Our specimens agree
in all respects with Lumsden and Zischke's
(1961) diagnoses. Our specimens of Hassti-
lesia texensis from the muskrat represent a
new host record. Shoop and Corkum
(1981a) reported and diagnosed Alaria
mustelae, Baschkirovitrema incrassatum,
Gyrosoma singulare, Maritreminoides
nettae, and Pharyngostomoides procyonis
from Louisiana mammals. In that report
we noted M. nettae in raccoons; it is herein
reported from the mink as well (new host
record). In a more recent note, we (Shoop
and Corkum, 1982) commented further on
the status of G. singulare in this state. He-
terobilharzia americana has been reported
from Louisiana mammals by Malek et al.
(1961) and Kaplan (1964). Our collections
of H. americana from mink represent a
new host record. Carneophallus basodac-
tylophallus was originally described by
Bridgman (1969) from raccoon in Louisi-
ana as was Sellacotyle vitellosa from mink
by Sogandares-Bernal (1961). Lumsden
and Winkler (1962) reported Linstowiella
szidati from opossum. We have found it in
opossum as well as in raccoon. In addition
to these trematodes, we identified eight
other species that have not been previously
reported from Louisiana mammals and
that are of importance from epidemiolo-
gical or zoogeographical standpoints.
Table II compiles all trematodes reported
heretofore from mammals in the state of
Louisiana.
Family DIPLOSTOMIDAE Poirier, 1886
Alaria alarioides (Dubois, 1937)
Dubois, 1970
(Figure 1)
Synonyms: Diplostomum alarioides
Dubois, 1937; Enhydrodiplostomum alar-
ioides (Dubois, 1937) Dubois, 1944.
Hosts: Mustela vison Schreber and Lutra
canadensis (Schreber).
Location: Small intestine.
Locality: Belle River, Assumption Parish,
Louisiana.
Deposition: Univ. Nebraska State Mus.,
Manter Lab. Coll. No. 21367.
Diagnosis (based on ten mature
specimens): Body elongate, distinctly
bisegmented, 1650 (1400-1800) long by 540
(450-650) at the widest point. Forebody
spathulate, 777 (640-940) long by 540
(450-650) wide; pseudosuckers present as
depressions on either side of the oral
sucker, never observed evaginated. Hind-
body claviform, 907 (760-1050) long by
430 (400-480) wide, containing reproduc-
tive organs. Forebody tegument covered
with small spines; hindbody smooth. Oral
sucker terminal, 92 (80-100) long by 106
(90-120) wide; acetabulum weak, spher-
ical, 75 (60-80) long by 76 (60-90) wide,
often covered by the tribocytic organ;
tribocytic organ broadly elliptical when
evaginated, 348 (240-400) long by 280
(240-330) wide, with a longitudinal cleft.
Prepharynx and esophagus extremely short
or absent; pharynx usually in contact with
oral sucker, 77 (70-90) long by 65 (50-80)
wide; paired ceca extend to the posterior
end of body. Testes tandem, not equal;
anterior testis asymmetrical, laterally dis-
posed on either side of midline, 215
(200-250) long by 317 (290-350) wide; pos-
terior testis symmetrical, dumbbell-
shaped, much wider than anterior testis,
218 (190-250) long by 394 (350-410) wide,
with a ventro-median groove to allow pas-
sage of ceca, uterus, and vitellaria; ejacula-
tory duct opens into the genital atrium;
genital atrium opens posterior, subterm-
inally on the dorsal surface. Ovary spher-
ical, located in hindbody just in front of
No. 2
Trematodes of Mammals
111
Table I. Trematodes recovered from hunter-trapped mammals in Louisiana during the winters of 1981
and 1982.
Trematode
Hosts
No.
Examined
No.
Infected
%
Location
Alaria alarioides (Dubois, 1937)
Dubois, 1970
A. marcianae (La Rue, 1917)
Walton, 1949
A. mustelae Bosma, 1931
Amphimerus speciosus (Stiles and
Hassal, 1896) Barker, 1911
Baschkirovitrema incrassatum
(Dies., 1850) Skrjabin, 1944
Brachylaima virginiana
Dickerson, 1930
Carneophallus basodactylophallus
Bridgman, 1969
Cryptocotyle concava (Creplin, 1825)
Luhe, 1899
otter
7
mink
42
raccoon
37
bobcat
3
raccoon
37
mink
42
raccoon
37
domestic cat
4
otter
7
mink
42
opossum
2
raccoon
37
mink
Fibricola cratera (Barker and Noll,
1915) Dubois, 1932
F. lucida (La Rue, and Bosma, 1927)
Dubois and Rausch, 1950
mink
raccoon
opossum
mink
opossum
Gyrosoma singulare Byrd, Bogitsh,
and Maples, 1961
raccoon
mink
Hasstilesia texensis Chandler,
1929
Heterobilharzia americana
Price, 1929
muskrat
raccoon
mink
bobcat
Isthmiophora melis (Schrank, 1788)
Luhe, 1909
raccoon
mink
Linstowiella szidati (Anderson, 1944)
Anderson and Cable, 1950
raccoon
opossum
Maritreminoides nettae (Gower, 1938)
Rankin, 1939
mink
raccoon
Microphallus opacus (Ward, 1894)
Ward, 1901
raccoon
mink
Paragonimus kellicotti Ward,
1908
opossum
Pharyngostomoides procyonis
Harkema, 1942
raccoon
Quinqueserialis quinqueserialis
(Barker and Laughlin, 1911)
Harwood, 1939
muskrat
Rhopalias macracanthus
Chandler, 1932
opossum
Sellacotyle vitellosa Sogandares-
Bernal, 1961
mink
42
42
37
2
42
2
37
42
5
37
42
3
37
42
37
2
42
37
37
42
2
37
2
42
2
24
2
2
1
1
1
1
2
21
1
22
4
12
2
26
2
7
2
1
20
2
1
6
2
1
1
3
6
5
4
1
31
29
57
5
67
3
2
3
25
29
50
50
52
Sm. Int.
Liver
Sm
Int.
10
32
100
62
100
19
5
20
Cecum
54
Mes. Ven
5
33
16
Sm. Int.
5
3
50
7
16
14
10
50
Lungs
84
Sm. Int.
40
Cecum
50
Sm. Int.
5
»
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
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No. 2
Trematodes of Mammals
119
Louisiana.
Deposition: Univ. Nebraska State Mus.,
Manter Lab. Coll. No. 21373.
Diagnosis (based on ten mature
specimens): Body elongate, oval, slightly
attenuated anteriorly, 3850 (3420-4150)
long by 1050 (960-1300) at the greatest
width. Tegument aspinous. Ventral sur-
face with five longitudinal rows of spher-
ical glands. Oral sucker subterminal, 335
(320-350) in diameter; acetabulum absent.
Pharynx absent; esophagus short, paired
ceca extend to posterior end of body.
Testes opposite, highly branched, in pos-
terior end of body, 513 (405-610) long by
305 (260-390) wide; external seminal
vesicle tubular, coursing anteriad to the
base of the cirrus sac; cirrus sac elongate,
claviform, 1277 (1050-1500) long by 145
(125-170) at the greatest width; cirrus often
extruded and much coiled, densely beset
with spines; genital pore median, near
intestinal bifurcation. Ovary deeply lobed,
intertesticular, 334 (300-390) long by 210
(150-250) wide; Mehlis' gland immediately
anterior to ovary; uterus comprised of
transverse loops which may extend beyond
the ceca; metraterm distinct, 775 (700-900)
long; vitellaria pretesticular, in two, extra-
cecal bands. Eggs oval, 17 (16-18) long by
8 (7-9) wide, without polar filaments.
Excretory system not observed.
Discussion: Penn (1942) examined 1,780
muskrats from coastal Louisiana and re-
covered the trematodes Nudacotyle
novicia, Echinochasmus schwartzi, and
Paramonostomum pseudalveatum. Byrd
and Reiber (1942) examined three musk-
rats from the New Orleans area and
reported E. schwartzi and Phagicola nana
( = P. angrense). Because of their declining
numbers, we were unable to obtain a large
series of muskrats from trappers, but we
were successful in obtaining five carcasses.
Two of the muskrat harbored hundreds of
Quinquesehalis quinqueserialis in their
ceca. Although this species is considered a
ubiquitous parasite of muskrats in North
America, this is the first report of it from
Louisiana. Our measurements agree well
with those provided by Kinsella (1971) in
his study of intraspecific variation of Q.
quinserialis. The life-cycle has been eluci-
dated by Herber (1942) and includes the
freshwater snail, Gyraulis parvus, from
which monostome cercariae are released
and encyst on vegetation. The muskrat
becomes infected while grazing on vegeta-
tion containing the cysts.
Incidentally, one muskrat was infected
with thousands of Hasstilesia texensis in
the cecum (new host record). We have
found H. texensis in all of the swamp rab-
bits, Sylvilagus aquaticus, that we have
examined in Louisiana. As all of the speci-
mens from the muskrat were gravid and
showed neither stunting nor any anoma-
lies, we presume that the muskrat may
serve occasionally as a normal, definitive
host for this species.
SUMMARY
The following trematodes were collected
from hunter-trapped mammals in the
Atchafalya basin of Louisiana during the
winters of 1981 and 1982: Alaria alarioides
(Dubois, 1937) Dubois, 1970; Alaria mar-
cianae (La Rue, 1917) Walton, 1949;
Alaria mustelae Bosma, 1931; Amphi-
merus speciosus (Stiles and Hassal, 1896)
Barker, 1911; Baschkirovitrema incras-
satum (Dies. 1850) Skrjabin, 1944; Brachy-
laima virginiana Dickerson, 1930;
Carneophallus basodactylophallus Bridg-
man, 1969; Cryptocotyle concava
(Creplin, 1825) Luhe, 1899; Fibricola cra-
tera (Barker and Noll, 1915) Dubois, 1932;
Fibricola lucida (La Rue and Bosma, 1927)
Dubois and Rausch, 1950; Gyrosoma sin-
gulare Byrd, Bogitsh, and Maples, 1961;
Hasstilesia texensis Chandler, 1929;
Heterobilharzia americana Price, 1929;
Isthmiophora metis (Schrank, 1788) Luhe,
1909; Linstowiella szidati (Anderson,
1944) Anderson and Cable, 1950; Mari-
treminoides nettae (Gower, 1938) Rankin,
1939; Microphallus opacus (Ward, 1894)
Ward, 1901; Paragonimus kellicotti Ward,
1908; Pharyngostomoides procyonis
Harkema, 1942; Quinqueserialis quinque-
serialis (Barker and Laughlin, 1911) Har-
wood, 1939; Phopalias macracanthus
120
Tulane Studies in Zoology and Botany
Vol. 23
(Chandler, 1932; and Sellacotyle vitellosa
Sogandares-Bernal, 1961.
Adult trematodes reported from Louisi-
ana for the first time are: Alaria alarioides,
A. marcianae, Amphimerus speciosus,
Cryptocotyle concava, Isthmiophora
melis, Mircophallus opacus, Paragonimus
kellicotti, and Quinqueserialis quin-
queserialis.
New host records include Heterobilhar-
zia americana, Cryptocotyle concava, and
Maritreminoides nettae from mink; Alaria
marcianae, Amphimerus speciosus, and
Linstowiella szidati from raccoon; and
Hasstilesia texensis from muskrat.
Natural infections of adult Alaria mar-
cianae were found only in juvenile rac-
coons. This substantiates previous experi-
mental work which demonstrated that
adult raccoon are unsuitable for the
maturation of this trematode. The larvae,
however, are able to employ the adult rac-
coon as a paratenic host where they remain
undifferentiated in the subcutaneous fat.
Amphimerus speciosus is a well known
pathogen of dogs and cats in North
America, being herein reported from a
domestic cat and a raccoon. Synonymiza-
tion of the better known A . pseudofelineus
with A. speciosus is corroborated by our
observations.
Whether Isthmiophora is distinct from
Euparyphium has been debated by several
authors. We place our specimens in the
genus Isthmiophora on the basis of body
shape and testicular morphology. We com-
pared our specimens to those of Eupary-
phium beaveri and conclude they are dis-
tinct. This is the first report of /. melis
from a raccoon and, to our knowledge, the
only report of this species from North
America. We consider Euparyphium bea-
veri of Lumsden and Zischke, 1961 to be a
synonym of Baschkirovitrema incras-
satum.
Microphallus opacus is a common para-
site in the mink and raccoon in Louisiana.
Sogandares-Bernal (1965a) stated that the
aquatic snail, Amnicola, probably served
as first intermediate host and that several
species of crayfish served as second inter-
mediate hosts. Therefore, a hypothetical
life-cycle of M. opacus from Louisiana can
be proposed: the first intermediate host is
the aquatic snail, Amnicola; several cray-
fishes serve as second intermediate; and
the raccoon and mink are definitive hosts.
Fishes have yet to be reported with M.
opacus from Louisiana.
The first and second intermediate hosts,
as well as the larval stages, of Paragonimus
kellicotti have been previously reported
from Louisiana. However, this is the first
report from this state of the adult fluke in
a naturally infected definitive host, the
opossum. The locality of the infection is
noteworthy in that it was found in an
upper middle class Baton Rouge residential
area.
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December 15, 1982
COMPARATIVE VISCERAL TOPOGRAPHY OF THE
NEW WORLD SNAKE TRIBE
THAMNOPHIINI (COLUBRIDAE, NATRICINAE)
NITA J. ROSSMAN and DOUGLAS A. ROSSMAN
Museum of Zoology, Louisiana State University
Baton Rouge, Louisiana 70893
NANCY K. KEITH
Dept. of Experimental Statistics, Louisiana State University
Baton Rouge, Louisiana 70893
Abstract
The positions and lengths of a variety of visceral
organs in 631 preserved adult thamnophiine snakes
were determined in terms of ventral scute number and
converted into a per cent of total ventral number; a
mean was calculated for each taxon to allow compar-
ison with other taxa. Dice-Leraas diagrams were then
constructed for the following organ positions and
lengths: posterior end of heart, anterior and posterior
ends of Uver, posterior end of pancreas, anterior and
posterior ends of right and left kidney, Uver length,
right and left kidney lengths, heart-liver interspace,
and kidney overlap. Sexual dimorphism is apparent in
many of the characters examined. Apparently corre-
lated with their need for space to accommodate
developing young, females tend to have their anterior
and midbody organs placed more anteriorly and their
kidneys more posteriorly than those in males.
Stepwise discriminant analysis was performed on
the following four variables in male thamnophiine
snakes: posterior end of heart, anterior end of right
kidney, posterior end of left kidney, and kidney
overlap. The 294 specimens represented 11 groups —
7 genera plus Ruthven's four species groups of
Thamnophis. Two of four linear discriminant
functions were retained as they explain 83.2 l<^o of the
relative variation. Function 1 is generally an anterior
end of right kidney dimension, and function 2 is a
kidney overlap and posterior end of heart dimension.
More than 66% of the specimens were correctly classi-
fied by use of the model. All groups except Clonophis
could be classified with greater success than the 21%
prior probability obtained by placing them all in the
Elegans group of Thamnophis, the numerically
largest sample. The discriminant analysis was able to
distinguish among the seven genera (as well as among
Ruthven's four species groups of Thamnophis) at the
0.05 level except that Clonophis and Tropidoclonion
could not be distinguished from each other.
Although visceral topographic data alone do not
clearly delimit thamnophiine genera nor establish
inter- or intrageneric relationships, some trends are
apparent that serve to support taxonomic conclusions
based on other kinds of characters. Clonophis and
Regina can be distinguished from Nerodia, in which
genus they were formerly included. Thamnophis (less
proximus and sauritus) can also be distinguished from
Nerodia (less erythrogaster and valida). The Sauritus
group of Thamnophis differs markedly from the
other three species groups established by Ruthven in
most visceral topographic features. The ribbon snakes
(Sauritus group) frequently tend to have a posterior
displacement of organs, a condition often occurring
also in the short, semifossorial genera (Clonophis,
Seminatrix, Storeria, Tropidoclonion, Virginia). One
unique feature shared by all of the semifossorial
genera is the possession of a relatively long liver.
Introduction
The technique of determining snake vis-
ceral topography using ventral scutes as re-
ference points has received little attention
since its introduction by Thompson
seventy years ago. Although a moderate
amount amount of descriptive anatomical
work has appeared in print, very little has
EDITORIAL COMMITTEE FOR THIS PAPER:
DR. SAMUEL B. McDOWELL, Professor of Zoology, Rutgers University,
Newark, New Jersey 07102
DR. JAMES S. ROGERS, Associate Professor of Biology, University of New
Orleans, New Orleans, Louisiana 70122
DR. ROBERT A. THOMAS, Director, Louisiana Nature Center, New Orleans,
Louisiana 70127
123
124
Tulane Studies in Zoology and Botany
Vol. 23
been done of a comparative nature that
might be of taxonomic value, and none
using discriminant analysis. The present
study was undertaken to investigate the
possible taxonomic significance of visceral
topography in the tribe Thamnophiini of
the colubrid subfamily Natricinae.
Beddard (1908, 1909) characterized the
position of visceral organs in three genera
of boid snakes in terms of the distance
from the snout to the organ. He also mea-
sured organ length and the distance be-
tween organs. Beddard was convinced that
the position of viscera within the body of
snakes generally had systematic impor-
tance. Subsequent authors who also used
distance measurements were Atwood
(1916, 1918), Bergman (1941 et seq.), and
Brongersma (1951, 1957 a & b). Bergman
expressed the organ positions and lengths
as a per cent of snout-vent length, and
both he and Brongersma also presented
their data diagrammatically.
Thompson (1913a & b, 1914) was the
first to relate the position of the various
visceral organs to the ventral scutes in an
attempt to provide a simple, yet objective,
technique for stating the location of the
organs. The position of an organ was
expressed as a percentage of the total num-
ber of ventrals in order to compensate for
individual, sexual, and geographic varia-
tion in ventral number. This technique has
been utilized subsequently only by Thorpe
(1975), Underwood (1976), and Rasmussen
(1979). Thorpe determined the midpoint of
an organ rather than the anterior and
posterior ends, so his data are not
comparable to ours or to those of other
authors. Inasmuch as one has to ascertain
the anterior and posterior ends in order to
determine the midpoint, the latter would
appear to be an unnecessary complication
and if used alone it also results in a loss of
information.
Garrigues (1962), Bogert (1968), Collins
and Carpenter (1970), and Frenkel and
Kochva (1970) also gave organ positions
and lengths in terms of ventral number,
but they did not express their data as a per
cent of total ventrals. Also, by lumping his
samples for each species, Garrigues failed
to take sexual dimorphism into account.
Valle (1944-45), Bragdon (1953), and
Camazine et al. (1981) used ventral
number to pinpoint the location of various
posterior organs so that surgical proce-
dures could be carried out using the
smallest incisions possible. In each case,
the investigator counted ventral scutes
from the vent forward.
Materials and Methods
We examined 63 1 preserved adult speci-
mens, representing 8 thamnophiine genera
(only Adelophis was omitted because of its
rarity) and 35 species (4 being represented
by two subspecies or populations). Large
subadults were used only if their data fitted
into the range of variation for the taxon
under consideration. Juveniles were
rejected because their values tend to lie
outside the normal range of variation in
adults (see Bergman, 1958a, 1961b).
Only nongravid females or those with
undeveloped eggs were used because of the
distortion caused by developing embryos
(also noted by Bergman, 1961a; CoUins
and Carpenter, 1970; Thorpe, 1975). Be-
cause females tend to have their anterior
organs situated more anteriorly and their
kidneys more posteriorly than those of
males, each sex was considered separately
(see the Sexual Dimorphism section for
further discussion).
Using the Dowling method for counting
ventral scutes, we inserted insect pins in the
20th scute and in every 15th scute there-
after. Several midventral slits were made to
expose the organs being studied. The ven-
tral scute numbers at the anterior and
posterior ends of each organ were re-
corded; to faciUtate inter- and intraspecific
comparisons, a percentage was calculated
by dividing the scute number by the total
number of ventrals. The following organs
were considered where possible: heart,
liver, gall bladder, pancreas, right and left
kidneys. Lungs, thyroid, spleen, and
adrenals were not considered because they
were difficult to locate in many specimens.
Testes and ovaries were not considered
No. 2
Visceral Topography of Snakes
125
because of the varying size depending on
whether the specimens were in a breeding
or non-breeding state (see Matthews and
Marshall, 1956; Manna and Sircar, 1978).
Organ lengths, expressed as the total
number of ventral scales covered, were
also recorded and treated as a percentage
of total number of ventrals. The following
distances were measured and expressed in
the same manner: posterior end of heart to
anterior end of liver, posterior end of liver
to anterior end of gall bladder, distance
between or overlap of the right and left
kidneys. On museum material other than
that in the Louisiana State University
Museum of Zoology (LSUMZ), only the
heart, anterior end of liver, and kidneys
were examined in order to minimize the
number of incisions. Preliminary data on
LSUMZ specimens had indicated that
these organs were the most relevant to the
study.
The statistics used in the Inter- and
Intrageneric Comparisons section con-
sisted of calculating the mean, standard
deviation, and standard error of the mean
for each sex of each taxon, then construct-
ing graphs by the Dice-Leraas method as
discussed in Simpson et al. (1960). This
method presents a graphic representation
of differences between populations, and
the results appear in Figs. 1-19. The 95%
confidence interval of the mean was deter-
mined by dividing the standard deviation
by the square root of the sample size and
multiplying this figure by a value from the
Student's t-test table using n-1 degrees of
freedom (Runyon and Harber, 1968). Be-
cause of the very large confidence interval
generated by a sample of two specimens,
we constructed a Dice-Leraas diagram only
in those cases where we had a minimum
sample of three specimens of the same sex.
The confidence interval results in a plus or
minus figure relative to the mean. Where a
determination of the statistical significance
of the differences between means could not
be obtained from this graphic representa-
tion (using the three general rules on p. 353
in Simpson et al., 1960), then a Student's
t-test was used. When data are stated as
being significantly different in this paper,
it refers to the fact that the differences are
significant at the p< .05 level.
To minimize the possible effects of geo-
graphic variation, we attempted to sample
populations from as restricted an area as
possible. In four instances (Thamnophis
couchii, T. elegans, T. sirtalis, Tropido-
clonion lineatum) we treated different sub-
species or geographically distant popula-
tions as separate taxon samples. Because
enough male and female Thamnophis
eques could not be obtained from one geo-
graphic area, we used females of T. e.
megalops and males of T. e. virgatenuis.
Due to the existence of sexual di-
morphism, data for males and females
could not be combined for discriminant
analysis. We chose to restrict the discri-
minant analysis to the data for males; only
a relatively few confidence intervals could
be shown for females on the Dice-Leraas
diagrams because many of the confidence
intervals exceeded the ranges of variation.
Only those specimens that had data avail-
able for all characters were used. Six
variables (posterior end of heart, anterior
and posterior ends of right kidney, ante-
rior and posterior ends of left kidney, and
kidney overlap) were first run after the
values were standardized at the mean to
allow for comparisons. Because the poste-
rior end of the right kidney and the ante-
rior end of the left kidney were signifi-
cantly correlated, those characters were
eliminated to obtain a four-variable
explanatory and predictive model. The
posterior end of the right kidney and
anterior end of the left kidney values are
reflected in the kidney overlap figures.
Because of the relatively small number
of specimens in each sample, the 294 speci-
mens were placed in the following eleven
groups to achieve greater statistical signi-
ficance of the discriminant values:
1 . Clonophis kirtlandii — 6 specimens
2. Nerodia (cyclopion, erythrogaster,
fasciata, rhombifera, sipedon, valida)
— 51
3. Regina {alleni, grahamii, rigida, sep-
temvittata) — 24
126
Tulane Studies in Zoology and Botany
Vol. 23
4. Seminatrix pygaea — 8
5. Storeria {dekayi, occipitomaculatd)
— 15
Thamnophis (groups from Ruthven,
1908)
6. Sauritus group (proximus, sauritus)
— 14
7. Radix group [brachystoma,^ butleri,
eques { = megalops in Ruthven), mar-
cianus, radix] — 44
8. Elegans group [couchii couchii,^ c.
hydrophilus,' elegans terrestris,^ e.
vagrans,^ melanogaster, nigronucha-
lis,^ ordinoides, rufipunctatus { = an-
gustirostris in Ruthven), scalaris] —
63
9. Sirtalis group [chrysocephalus,^ cyr-
topsis ('eques in Ruthven), godmani,^
sirtalis fitchi, ' s. sirtalis] — 47
10. Tropidoclonion lineatum (Nebraska,
New Mexico, Texas) — 5
11. Virginia {yaleriae, striatula) — 17
Prior probabilities of group membership
were calculated by dividing the number in
any group by the total number in the
study. These prior probabilities are used in
classifying the specimens with the discri-
minant model.
Sexual Dimorphism
Details on sexual dimorphism in this
study appear in Tables I and II and in Figs.
1-19. A comparison of sexual dimorphism
data from this study with other studies
appear in Table III.
Anterior organ positions
The posterior end of the heart and the
anterior end of the liver in males are
located posteriorly to those positions in fe-
males in 11^0 and 81 % of the taxa, respec-
tively. Male Clonophis, Seminatrix, Stor-
eria, Tropidoclonion (for heart only), and
Virginia have the posterior end of the heart
and the anterior end of the liver located
posteriorly to those positions in females in
all species. In Nerodia, Regina, and
Thamnophis there is interspecific variabi-
Uty in both features. Male Thamnophis
'taxon described since Ruthven (1908)
Haxon not recognized by Ruthven (1908)
have the posterior end of the heart situated
posteriorly to that of females in 74% of the
taxa; male Nerodia in 67<^o; male Regina in
67% . The anterior end of the liver in males
lies posteriorly to that of females in 8O070
of the species of Nerodia, 73% of the taxa
of Thamnophis, and in the only species of
Regina for which data are available.
Midbody organ positions
Sexual dimorphism of the midbody
organ positions is not pronounced. Males
have the posterior end of the liver located
posteriorly to that of females in 50% of the
taxa, the posterior end of the gall bladder
posteriorly to that of females in 69%. The
posterior end of the liver is more poste-
riorly placed in males in 60% of the species
of Nerodia, both species of Storeria, and
in the one species of Virginia examined.
Males have the posterior end of the gall
bladder located more posteriorly than do
females in 60% of the species of Nerodia,
70% of the taxa of Thamnophis, and in
the one species of Storeria examined.
Posterior organ positions
In contrast to most of the preceding
characters, the kidneys exhibit marked
sexual dimorphism in many of their fea-
tures. The anterior ends of the right and
left kidneys in males are anterior to those
of females in all taxa, as are the posterior
ends of the right and left kidneys in 86%
and 73% of the taxa, respectively. The
posterior end of the right kidney in males is
situated anteriorly to that of females in all
species of Regina, Seminatrix, Tropido-
clonion, and Virginia, and 95% of the taxa
of Thamnophis. In Nerodia the posterior
end of the right kidney of males is situated
posteriorly to that of females in 67% of the
species. In both species of Storeria and in
half the species of Nerodia, the posterior
end of the left kidney of males is situated
posteriorly to that of females. The poste-
rior end of the left kidney of males is
anterior to that of females in 95% of the
taxa of Thamnophis, 67% of the species of
Regina, one population of Tropidoclo-
nion, and in both species of Virginia.
No. 2
Visceral Topography of Snakes
127
Table 1. Sexual dimorphism in certain thamnophiine snakes.
Character
post
ant .
post
post
ant .
post
ant .
post
, heart^
liver
. liver
i gall bladder
r. kidney
• r. kidney
1 . kidney
1. kidney
35
27
18
16
37
37
37
37
Position in eft? Position in 99
posterior to posterior to
that in 99 (or that in cfcC (or
o" organ longer') 9 organ longer")
7 77. 17%
81% 15%
50% 44%
69% 31%
0% 100%
14% 86%
0% 100%
24% 73%
liver length 17
gall bladder length 16
r. kidney length 3 7
1. kidney length 37
post, heart-ant.
liver interspace 26
kidney overlap 3 7
76%
69%
5%
0%
2 7%
13%
-a' significantly
cfd=99 different (pS05)
from 9?
6%
4%
6%
0%
0%
0%
0%
3%
6%
0%
3%
0%
0%
3%
:>7%
41%
22%
25%
81%
49%
86%
32%
18%
6%
46%
73%
12%
22%
means of the taxa were used in computing the figures in this table
Organ lengths and interspaces
The liver and gall bladder of females are
longer than those of males in 76% and
69% of the taxa, respectively. However,
the right and left kidneys of males are
longer than those of females in 92% and
100% of the taxa, respectively, probably
due to the presence of a hypertrophied
sexual segment in males (Matthews and
Marshall, 1956; Prasad and Reddy, 1972).
The male heart-liver interspace is longer
than that of females in 73% of the taxa, as
is the male kidney overlap in 84% of the
taxa. In 80% of the taxa of both Nerodia
and Thamnophis, males have a shorter
liver than do females; the hver is also
shorter in male Storeria dekayi (in S. occi-
pitomaculata the Hver shows no sexual
dimorphism). Data were available for both
sexes in only one species of Regina and one
of Virginia. Males have a shorter gall blad-
der than do females in 80% of the species
of Nerodia, in 70% of the taxa of Tham-
nophis, and in Storeria dekayi. Males of
Clonophis, Nerodia, Regina, Seminatrix,
Storeria, Tropidoclonion, and Virginia
have longer right and left kidneys than do
females. In all taxa of Thamnophis, males
have a longer left kidney than do females;
in 86% of those taxa, males also have a
longer right kidney. In all species of
Clonophis, Regina, Seminatrix, Storeria,
and Virginia, males have a longer heart-
Hver interspace than do females, as is the
case for 60% of the species of Nerodia and
64% of the taxa of Thamnophis. In all
species of Clonophis, Nerodia, Regina,
Seminatrix, Tropidoclonion, and Virginia,
males have a more extensive kidney
overlap than do females, as is the case for
76% of the taxa of Thamnophis.
Asymmetry of kidney lengths
In 76% of the taxa, females have the
right kidney longer than the left (24%
differ significantly). On the other hand,
males have the left kidney longer than the
right in 55% of the taxa (5% differ signi-
ficantly). In all species of Nerodia, the
right kidney is the longer one in both sexes.
Summary and conclusions
In general, the anterior and midbody
organs are placed more posteriorly in
males than in females, whereas the kidneys
of males are positioned more anteriorly
than those of females. This more anterior
positioning of the anterior organs and
more posterior positioning of the posterior
organs in females would allow greater
space for the developing young.
Inter- and Intrageneric Comparisons
To facilitate comparisons, each set of
128
Tulane Studies in Zoology and Botany
Vol. 23
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No. 2
Visceral Topography of Snakes
129
Table III. Data on sexual dimorphism reported In the literature. X indicates that the organ is longer or
located more caudally in sex indicated; ND that there is no appreciable dimorphism.
^
^
^"■^"--^^^ Character
^
u
^
c
c
Xi
^ >
>.
r.
•o £
TJ X
4J
■M U
Taxon ^^■^'^^
M OO
J^ 00
Ul >
tfl T3
*j -a
Ul T3
CU C
> c
c
O tu
C -H
c ■-<
0 ^
C -H
O'-H
<u fu
D. j:
« M
D.^
O. 00
n M
O.M
(D M
C1.M
^ i-H
u ^
^ -H
<S 9
a 9
d 9
d d
d 9
d 9
d 9
d 9
d 9
d 9
rf 9
d 9
Colubrldae, Natrlcinae ^
Natrix (=Phabdophis) ahrysarga
X
X
N. (=Rhabdcphis) subminiata^
X
X
X
X
X
X
X
X
X
It. ( =Sinonatrix ) trianguligera
N. (=Xenochi'ophis) vittata
X
X
X
X
X
Colubrldae ^
Ablabee (=aongiilosoma) baliodeira
X
X
X
X
X
X
X
X
ND
X
X
Calamapia multipunotata^
X
X
X
Coluber melanums (=Elapke
flavolineata)^
X
X
X
X
X
X
X
Dendrophis (=Dendrelaphis) pictus
ND
ND
X
X
Colubrldae, Homalopslnae
Enhydris plwnbea°
X
X
ND
ND
X
X
X
X
X
X
Fordonia leucobatia
X
X
X
Homalopsis bucaata^
ND
ND
TO
Hypsirhina (=Enhydris) altemans
X
X
X
ND
ND
ND
ND
ND
ND
ND
ND
Acrochordldae
Acroahordus gi-anulatus
A. Javaniaus'-^
X
X
X
X
X
X
X
ND
ND
ND
X
X
X
X
X
X
X
ND
Anllildae
Cylindrophis rufus
ND
ND
ND
ND
ND
ND
ND
ND
ND
ND
ND
Boldae
Xenopeltis icniaolor
ND
ND
ND
ND
ND
ND
ND
ND
ND
Elapldae
Hydrophis fasaiatus
X
Vlperidae
AgkistTodon pisaivorus
Tpimeresurus gramineus
X
ND
ND
ND
ND
X
X
X
X
X
X
X
X
X
Bergman (1959a); Bergman (1956b); Bergman (1950); Bergman (1963); Bergman (1965); Bergman (1961a); Bergman (1955b);
^Bergman (1960); ^Bergman (1951); ^°Bergman (1958a); ^^Bergman (1953); ^^Bergman (1955a); ^^Bergman (1962a);
l^ColUns and Carpenter (1970); 15Bergman (1961b).
Dice-Leraas diagrams (Figs. 1-19) was
divided into three equal triads, using the
highest and lowest individual values as
outer parameters. In these figures each ver-
tical bar represents the mean, each hori-
zontal line the range expressed as a per
cent, each black rectangle the 95% confi-
dence interval, each number the quantity
of specimens examined in that taxon, each
horizontal dotted line the separation
between genera, and each vertical dashed
Une the boundary between two triads. See
Table IV for a comparison of taxa assigned
to the lowest, middle, and highest triads.
Table V shows the degree to which each
taxon differs significantly from other taxa
in this study. The results of this study are
compared with those of other workers in
-Appendix A.
Posterior End of Heart. — Assuming,
strictly for the sake of comparison, that
the middle triad represents the normative
condition, there is a clear tendency for the
heart to be situated more posteriorly than
the norm in both sexes of Nerodia rhom-
bifera, Regina alleni, Seminatrix pygaea,
and Thamnophis sauritus (T. melano-
gaster, T. proximus, and T. rufipunctatus
exhibit similar tendencies, but to a lesser
degree). On the other hand, Tropidoclo-
nian lineatum and almost half the taxa of
Thamnophis (including some representa-
tives from three of Ruthven's species
groups) tend to have the heart displaced
anteriorly relative to the norm.
Anterior End of Liver. — The anterior
end of the liver lies markedly farther poste-
riorly than the norm in both sexes of
Regina alleni and Seminatrix pygaea. It
appears to extend slightly more anteriorly
than the norm in about half the taxa of
Thamnophis (the same ones having an
anteriorly displaced heart) and in female
Clonophis kirtlandii, Nerodia erythrogas-
ter, Storeria, Tropidoclonion, and
Virginia. The most posterior placement in
Nerodia is again found in TV. rhombifera,
and in Thamnophis again found in T.
130
Tulane Studies in Zoology and Botany
Vol. 23
Table IV. A comparison of various taxa in terms of the triads into which they
fall. L represents the lowest triad, M the middle triad, and H the highest triad.
^^■~--,..^^ Character
Taxon ^~~~^-,^^^
• u
0 OJ
u
• (U
4-' >
C -H
CO M
• u
■U QJ
m >
O -H
OJ
• >-i
•U U
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o «
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T3
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• J^
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c
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>
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x:
QJ 00
> c
•H Q>
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c
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c
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1 tn
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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
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Visceral Topography of Snakes
133
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134
Tulane Studies in Zoology and Botany
Vol. 23
melanogaster (except females), T. proxi-
mus, T. rufipunctatus, and T. sauritus.
Posterior End of Liver. — The poste-
rior end of the liver extends markedly
farther posteriorly than the norm in both
sexes of Virginia valeriae. In general,
Thamnophis other than T. proximus and
T. sauritus (and female T. melanogaster)
tend to have the posterior end of the liver
lying farther anteriorly than in any other
thamnophiines save Nerodia erythrogaster
and N. valida.
Posterior End of Pancreas. — In males
the posterior end of the pancreas extends
farthest posteriorly in Thamnophis prox-
imus and the two species of Storeria; in
females it extends farthest posteriorly in
Regina rigida, Thamnophis proximus, T.
sauritus, Tropidoclonion lineatum, and
the two species of Virginia. About half the
taxa of Thamnophis tend to have the pan-
creas located more anteriorly than in any
of the other thamnophiines except female
Nerodia valida; this condition is most
prounced in male T. eques. Unfortunately,
the absence of data for one of the sexes in
12 of the taxa greatly reduces the value of
the pancreas comparisons.
Anterior End of Right Kidney. — In all
Thamnophis except T. proximus and T.
sauritus, the right kidney in males lies
anterior to the position of that organ in all
other thamnophiines except Nerodia
erythrogaster. There is a similar tendency
in females, but it is neither as marked nor
as consistent. On the other hand, there is
marked posterior displacement from the
norm in both sexes of Seminatrix pygaea,
Thamnophis proximus, T sauritus, and
Virginia striatula, and a similar but slightly
less pronounced tendency in both sexes of
Clonophis kirtlandii and V. valeriae and in
females of Storeria occipitomaculata and
Tropidoclonion lineatum.
Posterior End of Right Kidney. — The
pattern of variation here is generally simi-
lar to that described in the preceding
account. The most notable difference,
however, is that only Thamnophis proxi-
mus, T. sauritus, and female Seminatrix
pygaea show a pronounced extension pos-
teriorly. A similar but less pronounced
trend appears in males of Clonophis kirt-
landii, all species of Nerodia (except N.
erythrogaster and N. valida), Regina
grahamii, and Seminatrix pygaea.
Anterior End of Left Kidney. — This
position lies posterior to the norm in both
sexes of Clonophis kirtlandii, Seminatrix
pygaea, Tropidoclonion lineatum, both
species of Storeria, Thamnophis proximus,
T. sauritus, and both species of Virginia,
and in females of Regina grahamii. Nero-
dia erythrogaster and about half the taxa
of Thamnophis show a slight tendency
toward anterior displacement from the
norm (in most cases this tendency is better
developed in males).
Posterior End of Left Kidney. — The
end of the left kidney extends more poste-
riorly than the norm in both sexes of
Storeria dekayi, Thamnophis proximus,
and T. sauritus, and to a lesser degree in
males of Storeria occipitomaculata and fe-
males of Regina grahamii, Seminatrix
pygaea, and Thamnophis radix. Males of
about half the taxa of Thamnophis show a
tendency toward anterior displacement
from the norm, as do females of T. nigro-
nuchalis.
Liver Length. — The liver is relatively
long in the genera Storeria, Tropidoclo-
nion, and Virginia. Unfortunately we have
no data for males of the latter two genera
or for female Seminatrix. Male Seminatrix
have an even longer liver than is found in
the other three genera. Two male and two
female Clonophis, although not shown on
the Dice-Lerras diagram because of the
small sample size, also have a relatively
long liver (mean values of 26.3 and 26.0,
respectively).
Right Kidney Length. — The right kid-
ney is relatively short in both sexes of Tro-
pidoclonion lineatum and in both species
of Virginia, and in females of Storeria oc-
cipitomaculata and males of Seminatrix
pygaea. In males there is a tendency
toward a greater length than the norm in
the species of Nerodia and about half the
taxa of Thamnophis; the same tendency is
present in females but it is developed to a
No. 2
Visceral Topography of Snakes
135
lesser degree. Notably, Clonophis and
Regina separate completely from Nerodia
on this character.
Left Kidney Length. — The left kidney
is relatively short in both sexes of Semina-
trix pygaea, Thamnophis sauritus, Tropi-
doclonion lineatum, and the two species of
Virginia, and in females of Clonophis kirt-
landii, Storeria occipitomaculata, and
Thamnophis proximus. The tendencies
seen with regard to left and right kidney
lengths are generally similar, but the dis-
tinction between Nerodia and Clonophis-
Regina is less clearly defined in the left
kidney length of males.
18 20 22 24 26
28 30 32
34 36 38
c.
kirtlandi i
cyclopion
erythrogaster
fasciata
rhombifera
sipedon
al leni
grahamii
rigida
septemvittata
pygaea
dekayi
occipitomaculata
chrysocephalus
couchii A
couchii B
cyrtopsis
elegans A
elegans B
eques
godmani
marcianus
melanogaster
nigronuchalis
proximus
radix
rufipunctatus
sauritus
scalaris
sirtalis A
sirtalis B
lineatum B
striatula
valeriae
K-
\- 5 {
N.
1
1
1
— 9 1
9
-6
R
1
1
1
1
1
1
i^-l
1 c
! ^
Se.
r
i
-^ 7
St.
Th.
— ■■■- 7
1 +5
■ 6 1
5 1
- 3
— 9
1
1 —.
iiHm^
-his
1 -^
1 ^
1±:.:i..
1^ — ^1
I
Tr.
1
1
1
■ .1
V.
• ^J ^ '
~1 ""^
^^M '
Figure 2. Location of the anterior end of the liver in male thamnophiine snakes (expressed as a % of total
ventrals).
136
Tulane Studies in Zoology and Botany
Vol. 23
Heart-Liver Interspace. — Both sexes
of Seminatrix pygaea and males of Regina
alleni have a relatively long interspace, a
tendency that is also seen in males of
Nerodia sipedon and Regina septemvit-
tata, and in females of Nerodia cyclopion,
Tropidoclonion lineatum, and three
species of Regina (no data available for
female septemvittata). The interspace is re-
latively short in females of Clonophis kirt-
landii.
Kidney Overlap. — The greatest degree
of kidney overlap occurs in Nerodia and a
few Thamnophis {cyrtopsis, male
18 20 22 24 26 28
30 32 34
36 38
c.
kirtlandii
cyclopion
erythrogaster
fasciata
±A 1
N.
»!' ,
1
-7\
rhombifera
sipedon
al leni
1 -^'0 1
R.
1
1 -f-
6
grahaniii
-H-^
rigida
septenivi ttata
pygaea
dekayi
occipitomaculata
chrysocephalus
couchii A
1 h
— 4
Se.
T
i
' 1
1 1
St.
-4'° 1
Th.
IV :
i'
1
couchii B
cyrtopsis
elegans A
elegans B
eques
godmani
marcianus
melanogaster
nigronuchal is
proximus
radix
rufipunctatus
' -1-
4 1
8 j
3 1
sauritus
r*-^
scalaris
sirtalis A
sirtalis B
lineatum B
striatula
valeriae
....'...^.
I
Tr.
f-l-6 1
V.
~^'" \
Figure 3. Location of the anterior end of the liver in female thamnophiine snakes (expressed as a % of total
ventrals).
No. 2
Visceral Topography of Snakes
137
marcianus, melanogaster, radix, scalaris).
The least amount of overlap occurs in
Storeria occipitomaculata and Virginia
striatula. Clonophis-Regina again separate
completely from Nerodia.
Liver-Gall Bladder Interspace. —
McDowell (1979) reported that the most
striking visceral feature of all Acrochordus
is the close proximity of the gall bladder to
the liver. In Acrochordus granulatus the
gall bladder usually lies behind the liver,
but is separated from it by less than one
gall bladder length; in A. arafurae the gall
bladder lies immediately behind the liver;
and in A. javanicus the gall bladder is
usually overlapped by the posterior end of
the liver. McDowell stated that Acrochor-
dus seems to be the only snake genus
known to have the gall bladder so near the
liver, and he noted that having the gall
bladder displaced far behind the liver is
often cited as a distinctive feature of
snakes.
A survey of Bergman's many studies
(1950-1965) on the visceral topography of
a wide variety of snakes reveals that the
condition described by McDowell (1979) is
somewhat more widespread than he had
thought and that this feature exhibits
sexual dimorphism in a number of species.
Bergman's findings can be summarized as
follows:
1. No interspace, liver overlaps gall
bladder: Colubridae, Homalopsinae — fe-
male Enhydris enhydris (\955e), Homa lop-
sis buccata (1951), male Hypsirhina { = En-
hydris) alternans (1960); Acrochordidae —
Acrochordus javanicus (1958a).
2. Interspace less than one gall bladder
length: Colubridae, Homalopsinae —
male Enhydris enhydris (1955e), female
Cerberus rhynchops (1955c), Hypsirhina
( = Enhydris) plumbea (1960); Acrochordi-
dae — Acrochordus granulatus (1958a);
Elapidae — Enhydrina schistosa (1955d).
3. Interspace one to two times gall blad-
der length: Colubridae, Natricinae —
male Matrix {- Sinonatrix) trianguligera
(1959b), female Matrix (^Xenochrophis)
vittata (1950); Colubridae, Homalopsinae
— female Hypsirhina ( = Enhydris) alter-
nans (1960), male Cerberus rhynchops
(1955c); Elapidae — female Hydrophis
fascia tus (1962a), female Thalassophis
anomalus (1954); Viperidae — Ancistro-
don ( = Calloselasma) rhodostoma
( 1 96 1 b) , Trimeresurus gramineus ( 1 96 1 b) .
4, Interspace more than twice gall blad-
der length: Colubridae, Natricinae — Ma-
trix {^ Rhabdophis) chrysarga (1959a), TV.
i^Rhabdophis) subminiata (1956b),
female TV. { = Sinonatrix) trianguligera
(1959b), male TV. { = Xenochrophis) vittata
(1950); Colubridae, Homalopsinae —
Eordonia leucobalia (1960); other Colubri-
bae — Ablabes { = Gongylosoma)
baliodeira (1963), Calamaria
multipunctata (1965), Coluber melanurus
( = Elaphe flavolineata) (1961a), C.
( = Elaphe) radiatus (1961a), Dendrophis
( = Dendrelaphis) pictus (1955b), Dryophis
(=Ahaetulla) prasinus (1956a), Elapoides
fuscus (1956-58), Ptyas korros, P. mucosa
(1952); Aniliidae — Cylindrophis rufus
(1953); Boidae — Xenopeltis unicolor
(1955a); Elapidae — Bungarus candidus,
B. fasciatus, male Hydrophis fasciatus,
Maja tripudians (1962b), male
Thalassophis anomalus (1954).
We found the Thamnophiini to be highly
variable in this character although the
majority of individuals do have an inter-
space greater than one gall bladder length
(see Table VI for details). Noteworthy
exceptions are the females of Thamnophis
melanogaster and Virginia valeriae, in
which the mean values are 0.9 and 0.2,
respectively. In general, the interspace
tends to be relatively short in most
Merodia, Storeria, and Virginia, and rela-
ti'ely long in Regina, most Thamnophis,
and Tropidoclonion. By far the greatest in-
terspace/gall bladder values occur in
Thamnophis proximus and T. sauritus, but
this reflects unusually short gall bladders
rather than exceptionally long interspaces
in these animals.
Asymmetry of Kidney Lengths
In only 11 taxa are the differences in
length between the right and left kidneys
statistically significant. The left kidney is
longer than the right in male Thamnophis
138
Tulane Studies in Zoology and Botany
Vol. 23
c. couchii (difference between means 1 .6,
significantly different at p<.01) and female
T. nigronuchalisilA, p<.02). The right
kidney is longer than the left in male
Thamnophis sauritus (1.2, p<.01) and
female T. cyrtopsis {\ .1 , p <.02), T. radix
(1.9, p<.01), T. sirtalis fitchi (2.0, p<01),
Nerodia cyclopion (1.4, p<.01), N.
rhombifera (1.7, p<.05), N. sipedon (1.2,
p<.02), Regina alleni (1.5, p<.02), and R.
grahamii (1.2, p<.05).
Discriminant Analysis
In an effort to ascertain which, if any,
characters could be used taxonomically to
separate genera and other groups, stepwise
discriminant analysis was performed using
the Statistical Package for the Social
JQ 42
56 58
N. cyclopion
erythrogaster
fasciata
rhombifera
sipedon
valida
R. grahamii
rigida
septemvittata
St. dekayi
occipitomaculata
Th,
Tr,
V.
couchii A
couchii B
cyrtopsis
elegans A
elegans B
eques
marcianus
melanogaster
nigronuchalis
proximus
radix
sauritus
sirtalis A
sirtalis B
lineatum B
striatula
valeriae
Figure 4. Location of the posterior end of the liver in male thamnophiine snakes (expressed as a % of total
ventrals).
No. 2
Visceral Topography of Snakes
139
Sciences (SPSS) (Nie et al., 1975; Hull and
Nie, 1979). Only adult male specimens
were used in this part of the study (see
Materials and Methods). Elsewhere in this
paper under Materials and Methods we
have discussed the details of how the speci-
mens were treated for the discriminant
analysis.
In stepwise dicriminant analysis, the
variable that best discriminates among the
groups enters the model first, then the next
best discriminating variable enters, etc.
The process terminates when there are no
more variables that contribute significantly
to discrimination among the groups. In
this analysis four variables were found to
discriminate among the groups. In the
rior right kidney, (3) posterior heart, and
(4) kidney overlap. In the four variable
model, all groups but Clonophis and Tro-
pidoclonion were significantly different
(p<.05) (see Table VII).
Eleven groups were used in the dis-
criminant analysis, and four linear discri-
minant functions were computed. How-
ever, only the first two were retained as
they explain 83.21 % of the relative varia-
tion (function 1 accounts for 59.82% and
function 2 accounts for 23.39%). Function
1 is generally a right kidney anterior di-
mension. Function 2 is a kidney overlap
and heart posterior dimension. The all-
groups scatterpoint diagram with two dis-
criminant functions appears in Fig. 20. On
dimension 1, we see that Seminatrix, the
Sauritus group of Thamnophis, and
Virginia are separated widely from the
Elegans, Sirtalis, and Radix groups of
Thamnophis. On dimension 2, we see that
Nerodia is the most widely separated group
from Tropidoclonion and Storeria. Ap-
parently, as the right kidney anterior mea-
sure increases, the specimens are more
likely to belong to Seminatrix, the Sauritus
group of Thamnophis, and Virginia.
Similarly, as right kidney anterior de-
creases, specimens are more likely to
belong to the Elegans, Sirtalis, and Radix
groups of Thamnophis. Also, as kidney
overlap and heart posterior measurements
increase, the specimens are more likely to
belong to Nerodia. Similarly, as these mea-
Table VII. F statistics and significance between pairs of taxa in the four variable model (df=4, 280).
Taxon
1 Clonophis
2
3
4
5
6
7
8
9
10
2 Nerodia
16.557
0.0000*
3 Regina
7.4392
0.0000
15.827
0.0000
4 Seminatrix
11.479
0.0000
26.348
0.0000
9.4137
0.0000
5 Storeria
6.7379
0.0000
52.616
0.0000
20.963
0.0000
20.288
0.0000
6 Thamnophis
(Sauritus group;
6.5426
0.0000
27.357
0.0000
11.059
0.0000
5.1882
0.0005
14.861
0 . 0000
7 Thamnophis
(Radix group;
8.6467
0.0000
38.890
0.0000
20.513
0.0000
37.208
0.0000
24.534
0.0000
37.326
0.0000
8 Thamnophis
(Elegans groupj
24.343
0.0000
70.264
0.0000
40.509
0.0000
52.621
0.0000
58.937
0.0000
74.748
0.0000
24.394
0.0000
9 Thamnophis
(Sirtalis group,)
16.569
0.0000
43.839
0.0000
27.789
0.0000
44.918
0.0000
42.816
0.0000
55.380
0.0000
7.5158
0.0000
4.7066
0.0011
LO Tropidoclonion
1.2101
0.3067
23.372
0 . 0000
11.002
0.0000
14.360
0.0000
8.1301
0.0000
11.953
0.0000
10.465
0.0000
22.652
0.0000
16.875
0.0000
LI Virginia
7.9233
0.0000
57.601
0.0000
15.848
0.0000
7.0630
0.0000
28.365
0.0000
16.768
0.0000
50.209
0.0000
76.698
0.0000
62.552
0.0000
6.9183
0.0000
* significance leve
!l
140
Tulane Studies in Zoology and Botany
Vol. 23
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No. 2
Visceral Topography of Snakes
141
surements decrease, the specimens are
more likely to belong to Tropidoclonion
and Storeria.
The model was used to classify the 294
original specimens. The classification
matrix indicates how specimens were class-
ified by the model (see Table VIII). Over
66% of the specimens were correctly class-
ified. The Elegans group of Thamnophis,
which had the largest number of speci-
mens, had the highest prior probability of
21.4%. In the order of highest percentage
to lowest, Virginia was classified correctly
88.2% of the time, Nerodia 86.3%, the
Sauritus group of Thamnophis 85.7%, the
Elegans group of Thamnophis 76.2%,
Seminatrix 75.0%, Storeria 66.1 ^q, the
Radix group of Thamnophis 63.6%,
Tropidoclonion 60.0%, Regina 58.3%, the
Sirtalis group of Thamnophis 29.8%, and
36 38 40 42 44 46
%
48
50 52 54 56 58 60
N. cyclop ion
erythrogaster
fasciata
rhombifera
sipedon
val ida
R. grahamii
rigida
septemvittata
St. dekayi
occipitomaculata
Th. couchii A
couchii B
cyrtopsis
elegans A
elegans B
eques
marcianus
melanogaster
nigronuchalis
proximus
radix
sauritus
sirtalis A
sirtalis B
Tr. lineatum B
V. striatula
valeriae
4-^
— 4
Figure 5. Location of the posterior end of the liver in female thamnophiine snakes (expressed as a % of
total ventrals).
142
Tulane Studies in Zoology and Botany
Vol. 23
Clonophis 16.7<^o (less than chance). Vir-
ginia has the highest percentage correctly
classified, the Sauritus group of Thamno-
phis the third highest, and Seminatrix the
fifth highest, a notable finding inasmuch
as these taxa ranked only sixth, eighth, and
ninth, respectively, in terms of the number
of specimens per group.
The discriminant analysis was able to
distinguish among the eight genera (as well
as among Ruthven's four species groups of
Thamnophis) at the 0.05 level ex'^ept that
Clonophis and Tropidoclonion could not
be distinguished from each other. All
groups except Clonophis could be classi-
fied by the model with greater success than
the 21% prior probability obtained by
placing them all in the Elegans group of
Thamnophis, the numerically largest
sample. Thus the visceral topographic data
are remarkably concordant with the other
kinds of morphological data that have
4i 46 46 50 52 54
%
56
N. cyclopion
erythrogaster
fasciata
rhombifera
sipedon
valida
R. grahamii
rigida
St. dekayi
occipitomaculata
Th. couchii A
couchii B
cyrtopsis
elegans A
elegans B
eques
marcianus
mel an og aster
nigronuchal is
proximus
radix
sauritus
sirtalis A
sirtalis B
Tr. lineatum B
V. striatula
valeriae
5 8 60 62 64 66 68
Figure 6. Location of the posterior end of the pancreas in male thamnophiine snakes (expressed as a % of
total ventrals).
No. 2
Visceral Topography of Snakes
143
been used to generate the existing
classification of thamnophiine snakes.
Within the genus Thamnophis, all of
Ruthven's species groups except the 5/>-
talis group are distinguishable from each
other. In the classification matrix (Table
VIII), members of the Sirtalis group are
misclassified as members of the Elegans
group much more frequently (42. 6*^0) than
they are correctly classified (29.8%); they
also are often misclassified (21.3%) as
members of the Radix group. One of the
most interesting results of the discriminant
analysis is the wide separation of the
Sauritus group from the other three groups
(see Fig. 20).
Conclusions
Visceral topographic data alone do not
clearly delimit thamnophiine genera nor
establish inter- or intrageneric relation-
ships. Nevertheless, some trends are
%
cyclop ion
erythrogaster
fasciata
rhombifera
sipedon
valida
R, grahamii
rigida
St. dekayi
occipitomaculata
Th. couchii A
couchii B
cyrtopsis
elegans A
elegans B
eques
marcianus
melanogaster
nigronuchal is
proximus
radix
sauritus
sirtalis A
sirtalis B
Tr. lineatum B
V. striatula
valeriae
46 48 50 52 54 56 58 60 62 64 66
Figure 7. Location of the posterior end of the pancreas in female thamnophiine snakes (expressed as a % of
total ventrals).
144
Tulane Studies in Zoology and Botany
Vol. 23
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 0 2 i 6
% Males
10 12 -4 2 0 2 4 6 8 10 12 14
C. kirtlandii
N. cyclopion
erythrogaster
fasciata
rhombifera
sipedon
val ida
R. alleni
grahamii
rigida
septemvittata
Se. pygaea
St. dekayi
occipi tomaculata
Th. brachystoma
butleri
chrysocephalus
couchii A
couchii B
cyrtopsis
elegans A
elegans B
eques
godmani
marcianus
melanogaster
nigronuchalis
ordinoides
proximus
radix
rufipunctatus
sauritus
scalaris
sirtalis A
sirtalis B
Tr. lineatum A
lineatum B
V. striatula
valeriae
Figure 19. Kidney overlap in thamnophiine snakes (expressed as a <Vo of total ventrals).
No. 2
Visceral Tonography of Snakes
157
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158
Tulane Studies in Zoology and Botany
Vol. 23
groups designated by Ruthven (1908) do
not appear to be distinguishable from one
another solely on the basis of visceral topo-
graphy.
As was implied above, in many cases the
small, semifossorial thamnophiines tend to
have a posterior displacement of organs, a
condition they share frequently with the
ribbon snakes (Thamnophis proximus, T.
sauritus) and occasionally with some
species of Nerodia and Regina. Posterior
displacement is a general trend, not an in-
variable phenomenon, and both inter- and
intrageneric variation occur from one
character to the next. The semifossorial
genera also show a definite trend toward
Table IX. Significance of Ruthven' s Thamnophis groups compared as four se irate
populations. NS indicates difference not significant at p>.05.
Character
Sex
Sauritus-
Radix
Sauritus -
Elegans
Sauritus -
Sirtalis
Radix-
Elegans
Radix-
Sirtalis
Elegans -
Sirtalis
post, heart
9
.001^
.01
.001
.001
.001
.001
NS
NS
NS
NS
NS
NS
ant. liver
d
9
.001
.001
.001
.001
.001
.001
.05
.05
.05
.001
NS
NS
post, liver
d
9
.001
.001
.001
.001
.001
.001
NS
NS
NS
NS
NS
.05
post, gall bladder
d
9
.001
.001
.001
.001
.001
.001
NS
NS
NS
NS
NS
NS
post, pancreas
cf
9
.001
.001
.001
.001
.001
.001
NS
NS
NS
NS
NS
NS
ant. r. kidney
d
9
.001
.01
.001
.001
.001
.001
.001
.001
.001
.01
.01
NS
post . 1. kidney
d
9
.001
.001
.001
.001
.001
.001
.001
.001
.PI
.dOl
NS
NS
ant. 1 kidney
d
9
.001
.001
.001
.001
.001
.001
.001
.001
.00-
.001
.01
NS
post 1. kidney
d
9
.001
.001
.001
.001
.001
.001
.001
.001
.001
.001
.001
NS
liver length
d
9
NS
NS
.02
NS
.01
NS
NS
NS
NS
.02
NS
NS
r. kidney length
d
9
.01
.001
.01
.001
.01
.001
NS
.01
NS
NS
NS
NS
1. kidney length
d
9
.001
.001
.001
.001
.001
.001
NS
NS
NS
NS
NS
NS
heart-liver
interspace
cC
9
.02
NS
.01
NS
NS
NS
NS
NS
VS
NS
NS
NS
kidney overlap
d
9
NS
.001
NS
.01
NS
.001
NS
NS
NS
K:
NS
NS
Significance levels determined using 2-tailed Student's t-test.
No. 2
Visceral Topography of Snakes
159
having relatively short kidneys, but the
data for Storeria are equivocal and the
characteristic is not unique to those
genera. One unique feature that is shared
by all of the semifossorial genera is the
possession of a relatively long liver. We do
not know why small snakes would possess
a proportionally longer liver than large
snakes, but perhaps there are physiological
constraints that prevent the mutual reduc-
tion of body and of liver from being
directly proportional — perhaps a mini-
mum quantity of liver tissue is required for
the proper functioning of that organ.
Acknowledgments
For the loan of specimens and for other
courtesies, we are indebted to the follow-
ing curators: Charles W. Myers and
Richard G. Zweifel (American Museum of
Natural History); Harry A. Shankman
(Arizona State University); Douglas C.
Cox (Brigham Young University); C.J.
McCoy, Jr. (Carnegie Museum); Hymen
Marx (Field Museum of Natural History);
Walter Auffenberg and Peter Meylan
(Florida State Museum); Kenneth CHffer
and Philip J. Regal (James Ford Bell
Museum of Natural History); Joseph T.
Collins and William E. Duellman (Uni-
versity of Kansas Museum of Natural
History); Gloria Z. Wurst and David B.
Wake (Museum of Vertebrate Zoology,
University of California at Berkeley);
Harold A. Dundee (Tulane University); T.
Paul Maslin (University of Colorado
Museum of Natural History); Gary Brei-
tenbach and Arnold G. Kluge (University
of Michigan Museum of Zoology); James
F. Jackson (University of Southwestern
Louisiana); Jonathan A. Campbell (Uni-
versity of Texas at Arlington); and Robert
G. Webb (University of Texas at El Paso).
We are also grateful to Darrel R. Frost,
Mark S. Hafner, Dominique G. Homber-
ger, and Randy H. Vaeth for helpful sug-
gestions at various stages in the develop-
ment of this manuscript.
Specimens Examined'
Clonophis kirtlandii. ILLINOIS, Christian Co.:
LSUMZ 40065; Cook Co.: FMNH 23166, 25437;
/Gross Pt.y: FMNH 2989; Will Co.: FMNH 55562,
65902. INDIANA, Delaware Co.: FMNH 64670;
Porter Co.: FMNH 42069; /Orange Co.?y FMNH
3060. KENTUCKY, Jefferson Co.: FMNH 25535.
OHIO, Hamilton Co.: LSUMZ 7445, 13539.
Nerodia cyclopion. LOUISIANA, Ascension Par.:
LSUMZ 13703; Calcasieu Par.: LSUMZ 12150;
Cameron Par.: LSUMZ 18671-2; Iberville Par.:
LSUMZ 18286, 20703, 24669; Jefferson Par.:
LSUMZ 8670, 13704; Lafourche Par.: LSUMZ
13557, 19183; St. Bernard Par.: LSUMZ 9280; St.
Charles Par.: LSUMZ 18757, 29355; St. James
Par.: LSUMZ 18293, 19174; St. Tammany Par.:
LSUMZ 34308; Vermilion Par.: LSUMZ 24025,
33939.
Nerodia erythrogaster. LOUISIANA, Acadia Par.:
LSUMZ 20310; Cameron Par.: LSUMZ 20344;
East Baton Rouge Par.: LSUMZ 17321, 17702,
19175, 20312, 20723, 22909, 24028; Iberville Par.:
LSUMZ 18287, 22558-9; Jefferson Par.: LSUMZ
18716; Livingston Par.: LSUMZ 28812; St. Ber-
nard Par.: LSUMZ 8992; St. John the Baptist Par.:
LSUMZ 23864; St. Tammany Par.: LSUMZ
12983, 20279; Vermilion Par.: LSUMZ 34295;
Washington Par.: LSUMZ 12540; West Baton
Rouge Par.: LSUMZ 11887; West FeHciana Par.:
LSUMZ 18758.
Nerodia fasciata. LOUISIANA, Ascension Par.:
LSUMZ 17698; Cameron Par.: LSUMZ 12731,
17315, 20281, 28666; Jefferson Par.: LSUMZ 8947,
8953; Natchitoches Par.: LSUMZ 30410; Plaque-
mines Par.: LSUMZ 8653; Pointe Coupee Par.:
LSUMZ 20274; St. Charles Par.: LSUMZ 7142,
7527; St. Landry Par.: LSUMZ 18113, 18122; St.
Martin Par.: LSUMZ 19171, 19173.
Nerodia rhombifera. LOUISIANA, East Baton
Rouge Par.: LSUMZ 17687, 17794, 17945, 20799,
23662, 28008-10; Iberville Par.: LSUMZ 13756; St.
Charles Par.: LSUMZ 9216.
Nerodia sipedon. ALABAMA, Jackson Co.:
LSUMZ 36375; Pickens Co.: LSUMZ 36399,
36400. ILLINOIS, Jackson Co.: LSUMZ 27610;
Pope Co.: LSUMZ 27599. MISSISSIPPI, Greene
Co.: LSUMZ 36379, 36381-3, 36385, 36387,
36390-3, 36396-7; Lauderdale Co.: LSUMZ
36403-4; Wilkinson Co.: LSUMZ 28712.
MISSOURI, Lawrence Co.: LSUMZ 9107.
Nerodia valida. MEXICO, Colima: LSUMZ 7876;
Nayarit: LSUMZ 33099, 36266, 36268; Sinaloa:
AMNH 36269, 84077, 84080-2, 87575, 87577,
88889-90, 88892; Sonora: AMNH 84074-6.
Regina alleni. FLORIDA, Alachua Co.: FSM 2476,
2498, 6634, 6637, 7171, 9096, LSUMZ 13618-9;
Collier Co.: LSUMZ 28992; Dade Co.: FSM
42527; Dixie Co.: LSUMZ 7473; Hillsborough Co.:
FSM 42529; Indian River Co.: FSM 42524-6,
160
Tulane Studies in Zoology and Botany
Vol. 23
42530; Polk Co.: FSM 1868; Sumter Co.: FSM
11157.
Regina grahamii. LOUISIANA, East Baton Rouge
Par.: LSUMZ 17947, 33460, USL 7623; Iberville
Par.: LSUMZ 20271; Lafayette Par.: USL 20945;
St. Landry Par.: LSUMZ 28665, USL 15936,
23236, 23414, 23427; St. Martin Par.: USL 22953,
24432; Terrebonne Par.: LSUMZ 36484-7; Vermi-
lion Par.: USL 10687, 17353. TEXAS, Chambers
Co.: LSUMZ 33462.
Regina rigida. NO DATA: USL 6067, 8820. LOUISI-
ANA, Iberville Par.: LSUMZ 22556; Lafayette
Par.: USL 24245; Natchitoches Par.: LSUMZ
12988; Orleans Par.: LSUMZ 8982-3; Sabine Par.:
USL 24453; St. Charles Par.: LSUMZ 8680; St.
Landry Par.: USL 15930, 17620; St. Martin Par.:
USL 14365, 19471, 22425, 24433; Terrebonne Par.:
LSUMZ 36483.
Regina septemvittata. ALABAMA, Baldwin Co.:
LSUMZ 15783. NORTH CAROLINA, Orange
Co.: LSUMZ 14353^. OHIO, Montgomery Co.:
LSUMZ 24476, 30184-5. TENNESSEE, Clay Co.:
LSUMZ 34795; Jackson Co.: LSUMZ 34798.
Seminatrix pygaea. FLORIDA, Alachua Co.: FSM
9813 (-6,-12), 14146 (-4), 14147 (-1,-7), 14215 (-4),
14216 (-2,-4,-9), 14217 (-3,-5,-7), 14218 (-4,-6);
Dade Co.: LSUMZ 6530, 7405, 24582.
Storeria dekayi. LOUISIANA, Ascension Par.:
LSUMZ 18776; Cameron Par.: LSUMZ 2764,
12196, 18168-70, 24038, 28819-20, 28822, 29977,
32649; Iberia Par.: LSUMZ 2771; Iberville Par.:
LSUMZ 12229, 23877; St. Landry Par.: LSUMZ
18665, 20074; Vermilion Par.: LSUMZ 24733.
Storeria occipitomaculata. LOUISIANA, Claiborne
Par.: LSUMZ 24658; East Feliciana Par.: LSUMZ
16686; Natchitoches Par.: LSUMZ 24745, 33077-8;
West Feliciana Par.: LSUMZ 12602, 17898.
Thamnophis brachystoma. PENNSYLVANIA, Clar-
ion Co.: CM 28292-3, 28295, 28297-9, 28302-3,
28306-9, 28311, 28313, 28317-8, 28320-1.
Thamnophis butleri. CANADA, Ontario: UMMZ
90193. INDIANA, Noble Co.: UMMZ 132822.
OHIO, Lucas Co.: UMMZ 68864, 99627(3).
MICHIGAN, Sanilac Co.: UMMZ 98774; Wash-
tenaw Co.: UMMZ 465234; Wayne Co.: UMMZ
89519. WISCONSIN, Waukesha Co.: AMNH
76178-80.
Thamnophis chrysocephalus. MEXICO, Guerrero:
AMNH 72500-1, 72503; Oaxaca, AMNH 91094-5,
93235, 97855-6, 97865-6, 97868-9, 97871.
Thamnophis couchii couchii. CALIFORNIA, Ama-
dor Co.: LSUMZ 16530, 16544; Kern Co.: LSUMZ
16549; Shasta Co.: LSUMZ 22938, 34587-8, 34590,
MVZ 18824-5; Tehama Co.: LSUMZ 16550;
Tulare Co.: LSUMZ 16547; Tuolumne Co.:
LSUMZ 34585.
Thamnophis couchii hydrophilus. CALIFORNIA,
Humboldt Co.: LSUMZ 34578; Shasta Co.:
LSUMZ 1655M; Trinity Co.: LSUMZ 34594-5.
OREGON, Jackson Co.: LSUMZ 16560-4, 16567.
Thamnophis cyrtopsis. ARIZONA, Coconino Co.:
LSUMZ 29940, 30062, 30083, 30088; Gila Co.:
LSUMZ 30061; Maricopa Co.: LSUMZ 30063,
30081; Pima Co.: LSUMZ 30066, 30090; Santa
Cruz Co.: LSUMZ 10035, 30072, 30076-7; Yavapai
Co.: LSUMZ 29943, 29945-6, 29948, 30064-5,
30067-8.
Thamnophis eiegans terrestris. CALIFORNIA,
Mendocino Co.: LSUMZ 34378, 34380; San Mateo
Co.: LSUMZ 7922, 16502-3, 16507, 34371, 34373;
Sonoma Co.: LSUMZ 34368-9, 34374-5; Sonoma-
Mendocino Co.: LSUMZ 34367.
Thamnophis eiegans vagrans. NO DATA: LSUMZ
20747-50. ARIZONA, Coconino Co.: LSUMZ
29957, 29959-62. COLORADO, Conejos Co.:
LSUMZ 11571, 11609, 11611, 11615, 30051, 30055;
Costilla Co.: LSUMZ 7985, 11603-5, 11607, 11614,
11618, 13929, 13931-2, 30050; Rio Grande Co.:
LSUMZ 30056.
Thamnophis eques megalops. MEXICO, Chihuahua:
AMNH 104471, 104772, BYU 22701; San Luis
PotosK LSUMZ 4374-5, 4879.
Thamnophis eques virgatenuis. MEXICO, Durango:
AMNH 102521, LSUMZ 16424-6, 16429-30.
Thamnophis godmani. MEXICO, Oaxaca: AMNH
89604, 91101-2, 91105, 97853, 97873-4, 97884,
97888, 103090, 103092-5, 103101, 103103, 103105,
103113, 104394, 106993, 106995-8, 107002-5,
718170.
Thamnophis marcianus. TEXAS, Bexar Co.
LSUMZ 10315; Duval Co.: LSUMZ 23239, 23243
Hartley Co.: LSUMZ 10407; Jeff Davis Co.
LSUMZ 29608; McMuUen Co.: LSUMZ 23248
Moore Co.: LSUMZ 10365; Presidio Co.: LSUMZ
23255; San Patricio Co.: LSUMZ 23249, 23252;
Webb Co.: LSUMZ 30929; Zavala Co.: LSUMZ
23254.
Thamnophis melanogaster. MEXICO, Jalisco:
LSUMZ 16434; Michoaca'n: LSUMZ 14489-90,
14492-3, 16435, 34346, 36277, 36279-80, 36282-6.
Thamnophis nigronuchalis. MEXICO, Durango:
LSUMZ 11637, 16448, 16450-5, 16459-60, UTEP
3386-7.
Thamnophis ordinoides. CALIFORNIA, Del Norte
Co.: MVZ 30276-7, 30279. OREGON, Clatsop
Co.: MVZ 34265-8, 36848; Polk Co.: MVZ 24808;
TUlamook Co.: MVZ 47856. WASHINGTON,
Clark Co.: MVZ 34259; King Co.: MVZ 38653,
38655, 38657, 38670, 38674; Lewis Co.: MVZ
70366; Pacific Co.: MVZ 34262.
Thamnophis proximus. LOUISIANA, Acadia Par.:
LSUMZ 17899; Cameron Par.: LSUMZ 33964;
No. 2
Visceral Topography of Snakes
161
Claiborne Par.: LSUMZ 33966; East Baton Rouge
Par.: LSUMZ 16912, 18714, 20254; Iberia Par.:
LSUMZ 18077; Iberville Par.: LSUMZ 20255,
20316, 22548; Livingston Par.: LSUMZ 7960,
18974; Pointe Coupee Par.: LSUMZ 20220; St.
Tammany Par.; LSUMZ 7934; Vermilion Fax.:
LSUMZ 24052. TEXAS, Hidalgo Co.: LSUMZ
18621-3.
Thamnophis radix. NO DATA: LSUMZ 20735^0,
20742-5. COLORADO, Denver Co.: LSUMZ
7465; Larimer Co.: UC 31837^0, 31842-3, 31847,
31851, 31873, 31888. ILLINOIS, Iroquois Co.:
LSUMZ 8126. NEW MEXICO, San Miguel Co.:
LSUMZ 7942, 7944, 7972.
Thamnophis rufipunctatus. ARIZONA, Coconino
Co.: LSUMZ uncatalogued, LSUMZ 36815.
MEXICO, Chihuahua: AMNH 4342, 68286, ASU
17042, 5304-5, 5335, UTEP 2043, 2262, 3657.
Thamnophis sauritus. FLORIDA, Alachua Co.: FSM
14183, 14550 (-1), 14550 (-2), 14550 (-3), 14550 (-4),
14550 (-7), 14550 (-8), 14550 (-9), 14551 (-2), 14551
{,-4), 39197; Collier Co.: FSM 39198, 39200-2; Dade
Co.: FSM 22874, 39204-5; Franklin Co.: LSUMZ
21805-6, 21810; Pasco Co.: LSUMZ 22003. LOUI-
SIANA, St. Tammany Par.: LSUMZ 8302, 23770.
Thamnophis scalaris. MEXICO, Distrito Federal:
AMNH 75934; JaUsco: UTA R-4932, R^949,
5991, 5993; Mexico: AMNH 71315 (2), 94714;
Michoacan: AMNH 88724.
Thamnophis sirtalis fitchi. CALIFORNIA, Amador
Co.: LSUMZ 16486-8, 16489-92; Mendocino Co.:
LSUMZ 16493; Modoc Co.: LSUMZ 8215; Plumas
Co.: LSUMZ 16477-8, 16481-2; Shasta Co.:
LSUMZ 16496-8.
Thamnophis sirtalis sirtalis. INDIANA, Allen Co.:
LSUMZ 7988. MINNESOTA, Carlton Co.: JFBM
1115, Cass Co.: LSUMZ 7991, 7996; Clearwater
Co.: JFBM 2644-5, 2651-2, 2657, 2659; Isanti Co.:
LSUMZ 23229, 23232, 23234, 24461-2; Pine Co.:
LSUMZ 23230.
Tropidoclonion lineatum. NEBRASKA, Jefferson
Co.: KU 45252-65, 45267-8; Richardson Co.: KU
52228. NEW MEXICO, San Miguel Co.: LSUMZ
29998-9, 30096-7. TEXAS, Travis Co.: LSUMZ
20078-9.
Virginia striatula. NO DATA: USL 5395, 15841.
LOUISIANA, Acadia Par.: LSUMZ 12091; Ascen-
sion Par.: LSUMZ 12087, 18777; Caddo Par.:
LSUMZ 20210; East Baton Rouge Par.: LSUMZ
1598, 1604-5, 2786, 17348, 18712, 23536, 23745;
East Feliciana Par.: LSUMZ 2779; Lafayette Par.:
USL 11179, 22890; Livingston Par.: LSUMZ
12126; Sabine Par.: LSUMZ 20193; St. Helena
Par.: LSUMZ 18360; St. Landry Par.: USL 18277;
St. Tammany Par.: LSUMZ 2773.
Virginia valeriae. FLORIDA, Alachua Co.: FSM
42545; Leon Co.: FSM 1942, 34858; Liberty Co.:
FSM 42531-2, 42534-5; Wakulla Co.: FSM 32991.
GEORGIA, Chattahoochee Co.: FSM 42546.
LOUISIANA, Bossier Par.: LSUMZ 24656; Caddo
Par.: LSUMZ 12094; East Baton Rouge Par.:
LSUMZ 12147, 17671; East Feliciana Par.:
LSUMZ 15536; Livingston Par.: LSUMZ 20256;
St. Helena Par.: TU 5957; St. Tammany Par.: TU
1 1844, 14238, 18395; Webster Par.: LSUMZ 12142;
West Feliciana Par.: LSUMZ 17901. MISSISSIP-
PI, Hancock Co.: TU 14304, 15056, 17681.
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Appendix A
Comparative data on non-thamnophiine
snakes obtained from the literature fell
within the outer parameters of the thamno-
phiine data sets generated by our study
except for the following taxa whose organ
positions lie more posteriorly or which
have longer organs or interspaces.
Posterior end of heart — non-natricine
and non-homalopsine Colubridae:
Boiga,^ Chamaetortus,^ Coluber
( = Gonyosoma) oxycephalus, ^
Dipsadoboa, ' and male Zamenis
rhodorhacis;^ Acrochordidae: Acro-
chordus arafurae,^ A. granulatus,^ A.
javanicus;^ Boidae: male Bolyeria/ male
Corallus,* rnale Eunectes,* mdXt Licha-
nura,* male Loxocemus,* male Xenc-
peltis;* Viperidae: C<2W5W5 rhombeatus.^
Anterior end of right kidney — non-natri-
cine and non-homalopsine Colubridae:
Coluber ( = Gonyosoma) oxycephalus,^
female Philothamnus semivariegatus,^
male Psammophis sibilans,^ male
Zamenis florulentus,^ Z. rhodorhacis?
Posterior end of right kidney — non-natri-
cine and non-homalopsine Colubridae:
Coluber ( = Gonyosoma) oxycephalus, ^
male Leptodira ( = Crotaphopeltis)
hotamboeia,^ female Philothamnus
semivariegatus,^ male Psammophis sibi-
lans;^ Viperidae: male Causus
rhombeatus.^
Anterior end of left kidney — non-natri-
cine and non-homalopsine Colubridae:
male Coluber { = Gonyosoma) oxyce-
phalus,^ female Philothamnus semi-
variegatus,^ male Psammophis sibilans,^
male Zamenis florulentus.^
Posterior end of left kidney — non-natri-
cine and non-homalopsine Colubridae:
male Coluber { = Gonyosoma) oxyce-
phalus,^ female Philothamnus semiva-
riegatus,^ male Psammophis sibilans.^
Right kidney length — Viperidae: Causus
rhombeatus.^
Heart-liver interspace — The following
taxa had an overlap — Tropidophiidae:
Trachyboa gularis,^ Tropidophis;^
Viperidae: Causus rhombeatus.^
Kidney overlap — all taxa reported in the
literature have an overlap, but Causus
164
Tulane Studies in Zoology and Botany
Vol. 23
rhombeatus^ (Viperidae) is the only one
to have a greater overlap than any of the
Thamnophiini.
The following taxa have an organ posi-
tion lying more anteriorly or have shorter
organs than any of the Thamnophiini.
Posterior end of left kidney — Tropido-
phiidae: female Exiliboa placata.'
Liver length — non-natricine and non-
homalopsine Colubridae: female
Philothamnus semivariegatus,^
Heart-liver interspace — Colubridae, Na-
tricinae: male Natrix ( = Amphiesma)
vibakari^ from Japan.
Kidney asymmetry — In the present study
males in 64°7o of the taxa have the left
kidney longer than the right but the dif-
ference is significant in only 5"7o. How-
ever, the literature reveals that in the
'Rasmussen (1979)
Thompson (1914)
'McDowell (1979)
'Underwood (1976)
'Thompson (1913b)
'Brongersma (1951)
'Bogert (1968)
'Bergman (1959b)
'Bergman (1955e)
'"Bergman (1956-58)
"Bergman (1960)
males of most taxa the right kidney is
longer than the left. The following are
the taxa in which the left kidney is
longer: Colubridae, Natricinae — Natrix
( = Sinonatrix) trianguligera;* Colu-
bridae, Homalopsinae: Enhydris enhy-
dris;" other Colubridae: Coluber
( = Gonyosoma) oxycephalus,^
Elapoides fuscus.'" Females in 76% of
the thamnophiines have the right kidney
longer than the left (28% significantly
different) as do the females of all taxa
reported in the literature except:
Colubridae, Natricinae — Natrix
(=Amphiesma) vibakari;^ Colubridae,
Homalopsinae: Hypsirhina { = Enhy-
dris) plumbea;'' other Colubridae:
Elapoides fuscus. ' "
December 15, 1982
087