Chelonian Conservation and Biology, 2024, 23(1): 000–000

doi:10.2744/CCB-1589.1
Ó 2024 Chelonian Research Foundation

Morphometric Variation in the Red-Cheeked Mud Turtle (Kinosternon cruentatum) and its
Taxonomic Implications

JOHN B. IVERSON1,* AND JAMES F. BERRY2

1
Department of Biology, Earlham College, Richmond, Indiana 47374 USA [johni@earlham.edu];
2
Department of Biology, The Citadel, Charleston, South Carolina 29409 USA

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*Corresponding author

ABSTRACT. – The Scorpion Mud Turtle (Kinosternon scorpioides) has been recognized by most recent
authors to be a single species (3 subspecies) ranging from Tamaulipas, Mexico, to northern
Argentina. However, recent molecular analyses have demonstrated that it is not monophyletic, but
rather paraphyletic relative to other Neotropical Kinosternon. Based on extensive genetic sampling
across the range of this species, a recent paper elevated the 3 subspecies (albogulare, cruentatum, and
scorpioides) to species status, confirmed the divergence of Pacific and Atlantic versant populations of
K. cruentatum, and recommended that both be given species status. However, the type locality of
K. cruentatum is imprecise, making the allocation of the name problematic. Our study sought to deter-
mine the provenance of the type using morphometric analysis of specimens from across the range.
That analysis demonstrates unequivocally that the type of K. cruentatum was collected on the Atlantic
versant, likely from an eastern population. Furthermore, the analysis also revealed that the type of K.
mexicanum, previously synonymized with K. cruentatum, was collected from along the Pacific versant.
Hence, the name K. cruentatum should be restricted to populations in Atlantic drainages, and the
name K. mexicanum should be restricted to Pacific drainages. Our analysis also indicated diver-
gence among the 3 allopatric Atlantic versant populations, but future genetic work will be needed
to determine whether they merit taxonomic recognition.
KEY WORDS. – Kinosternidae; Kinosternon mexicanum; Kinosternon scorpioides: Mexico; taxonomy

Over more than 50 yrs, the Scorpion Mud Turtle
(Kinosternon scorpioides) has been considered by most
authors to be a single species ranging from Tamaulipas,
Mexico, to northern Argentina, the greatest distribution of
any living turtle (Turtle Taxonomy Working Group [TTWG]
2021). Fifteen synonyms of K. scorpioides (sensu lato [s.l.])
were subsequently described, but only 4 have been recog-
nized as subspecies in the recent literature, based on Berry’s
(1978) morphometric analyses: K. s. scorpioides from Pan-
ama to Argentina; K. s. albogulare from Panama to Guate-
mala; K. s. abaxillare from the Central Valley of Chiapas,
Mexico, and western Guatemala; and K. s. cruentatum
from northern El Salvador, Guatemala, and Belize to
Oaxaca and Tamaulipas, Mexico (Berry and Iverson
2001; Legler and Vogt 2013; TTWG 2021; Fig. 1).
However, some authors have recognized cruentatum
and/or albogulare as full species (Schmidt 1941; Davis
1953; Duellman 1965; Wermuth and Mertens 1977;
McCranie 2018), but without substantive morphometric
or genetic data to support those actions.
Based on 3 mitochondrial and 3 nuclear DNA
markers, Iverson et al. (2013) discovered that K. scor-
pioides (s.l.) was not monophyletic, and because K.
s. abaxillare was allopatric relative to the other subspe-
cies, as well as morphometrically and genetically dis-
tinct, they recommended its elevation to species level
(as proposed earlier by Wermuth and Mertens 1961 and
Alvarez del Toro 1973, 1982). That analysis also found the
remaining 3 subspecies to be paraphyletic, but made no
other taxonomic recommendations for the group pend-
ing further research. Two subsequent molecular phylo-
genetic analyses relying on 14 to 17 nuclear markers
(Spinks et al. 2014; Thomson et al. 2021) did not
include abaxillare among their samples, but also found
K. scorpioides to be paraphyletic, although they also
made no taxonomic recommendations.
More recently, Hurtado-Gómez et al. (2024) undertook
another molecular analysis of the genus Kinosternon using 3
mitochondrial and up to 17 nuclear markers, but with exten-
sive geographic sampling within K. scorpioides (and
Kinosternon leucostomum). That study clearly revealed
that K. scorpioides (s.l.) was polytypic, and supported
the recognition of K. cruentatum, albogulare, abaxillare,
and scorpioides as full species. Hurtado-Gómez et al.
(2024) also identified several additional lineages in
South and Central America that may eventually merit
taxonomic recognition.
Their analysis also revealed that populations of both
K. cruentatum and K. albogulare on the Pacific versant were
highly divergent from those on the Atlantic (e.g., 2.9% uncor-
rected p distance for the mitochondrial cytochrome b frag-
ment for the K. cruentatum populations). Hurtado-Gómez
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Figure 1. Distribution of Kinosternon cruentatum (sensu stricto; red shading) and Kinosternon mexicanum (blue shading) in
Mexico and Central America, demonstrating the allopatry of the Pacific, eastern Atlantic, Veracruz, and Tampico embayment
populations. Open circles are verified locality records (TTWG 2021). Green star is the restricted type locality of both K. mexi-
canum (and K. cruentatum sensu lato; see text); yellow star is our newly restricted type locality of K. cruentatum.

et al. (2024) made no taxonomic recommendations regarding
lineages within K. albogulare, but recommended that the
Atlantic and Pacific populations of K. cruentatum were
divergent enough to warrant species recognition. Interest-
ingly, Legler and Vogt (2013) recently mentioned that K. s.
cruentatum from Pacific populations have larger, and rela-
tively wider and higher shells than those on the Atlantic.
Nevertheless, if the recommendation of Hurtado-Gómez
et al. (2024) to split Atlantic populations of K. cruentatum
from the Pacific population is confirmed by morphometric
analysis, this will result in a complication of nomenclature.
Two of its current synonyms (K. mexicanum and K. trilira-
tum) lack precise type localities, a situation made even
more complex because Smith and Taylor (1950a, 1950b)
restricted the type localities of all 3 taxa to San Mateo del
Mar, Oaxaca, Mexico, without published justification.
Hence, the appropriate names for the Pacific and Atlantic
populations hinge on determining on which versant the
types were collected.
The purposes of this study were 1) to examine mor-
phometric variation within K. s cruentatum to determine
whether the Pacific and Atlantic populations differed
significantly (i.e., do these populations corroborate the
genetic data of Hurtado-Gómez et al. 2024); and if so, 2)
to identify taxonomic characters that diagnose the 2 pop-
ulations (proposed species); and 3) to attempt to estab-
lish the versant origin of the types of K. cruentatum and
K. mexicanum and thus restrict the correct names to the
appropriate population(s).
METHODS
The following measurements for specimens of K. (s.)
cruentatum from Berry’s (1978) doctoral dissertation
 IVERSON AND BERRY — Kinosternon cruentatum Morphometrics 0

et al. 1975,), but concordance of the statistical results

Table 1. Comparisons of body size (carapace length [CL]) by
sex for subpopulations of Kinosternon cruentatum (sensu lato) based on ratios vs. size-standardized residuals on these
based on data from Berry (1978) and additional specimens very data was established by Berry (1978).
measured by Iverson (J.B.I., unpubl. data). Based on the lack of significant morphometric varia-
Mean CL 6 Range tion across Atlantic drainages revealed by Berry (1978),
Versant Sex n 1 SD (mm) (mm) all Atlantic data were initially lumped into a single sam-
Pacific Male 46 122.0 6 8.5 102–143 ple for our initial analyses. Similarly, all Pacific drainage
Pacific Female 71 122.9 6 9.3 97–145 data were lumped into a single sample. These prelimi-
East Atlantic Male 80 111.7 6 6.6 94–134 nary analyses revealed substantial variation across the 3
East Atlantic Female 111 114.3 6 9.9 94–156

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Veracruz Male 5 121.0 6 9.9
Veracruz Female 11 120.8 6 7.4
Tampico Male 10 130.5 6 9.9
Tampico Female 12 126.5 6 21.9
were examined in this study: maximum carapace length,
maximum plastron length, maximum carapace width,
maximum shell height, bridge length, gular length, inter-
humeral seam length, interpectoral seam length, interab-
dominal seam length, interfemoral seam length, and
interanal seam length. Most measurements were taken to
the nearest millimeter by J.F.B. (see Berry 1978 for pre-
cise methods of measurement), although those for an
additional 25 specimens from the Atlantic versant were
measured by J.B.I. (listed in Supplemental Data S1; all
supplemental material is available at http://dx.doi.org/
10.2744/CCB-1589.1.s1) and included in our analyses.
Our analysis did not include data from populations of
the Rio Lempa basin in San Salvador, Honduras, and
Guatemala that Berry (1978) considered as potential
intergrades with albogulare. The full morphometric
data sets used in this study are included in the Supple-
mental Data S1. The latter 10 variables were standard-
ized for size by dividing each by carapace length. We
are well aware of the controversy surrounding the use of
ratios in statistical analyses (e.g., Atchley 1976; Atchley
110–135 allopatric Atlantic subpopulations of K. cruentatum (Fig.
107–131 1): the Yucatan region (from Belize across the Yucatan
120–151
93–148 peninsula), the central Veracruz region, and the Tampico
embayment. Our subsequent analyses examined morpho-
metric variation among these 3 subpopulations.
Population comparisons of character ratios were
made with t-tests and Mann-Whitney U-tests (i.e., Pacific
vs. Atlantic) and employed the STATVIEWtm statistical
package (Abacus Concepts, Berkeley, CA). These statisti-
cal analyses were used primarily to identify taxonomically
useful characters that demonstrated statistically significant
differences among populations. Mean ratios (6 1 SD) were
calculated by sex and subpopulation. Hence, initial vari-
ables (character states) identified as differing between
Pacific and Atlantic populations were recombined into
new character-state ratios that were useful in diagnosing
the 2 populations. Bivariate plots of these character
states by sex were produced to clarify the population
differences and facilitate the taxonomic diagnosis of
future specimens.
Discriminant function analyses (DFA; SPSS BASE
v.10.0) were performed (males and females separately) on
the 10 size-standardized ratio variables at 3 levels. First,
the Pacific population was compared to the 3 Atlantic
populations collectively; second, all 4 allopatric subpopu-
lations identified above were compared, and third, only
the 3 Atlantic versant subpopulations were compared.

Table 2. Character comparisons of Pacific and Atlantic populations of female Kinosternon cruentatum.

Pacific populations East Atlantic populations Veracruz population Tampico population

(n ¼ 70) (n ¼ 111) (n ¼ 11) (n ¼ 12)
Ratioa Mean 6 1 SD Range Mean 6 1 SD Range Mean 6 1 SD Range Mean 6 1 SD Range
PL/CL 0.953 6 0.022 0.902–1.008 0.972 6 0.031 0.908–1.103 0.971 6 0.021 0.941–1.000 0.978 6 0.024 0.994–1.028
GL/CL 0.163 6 0.021 0.117–0.228 0.162 6 0.017 0.101–0.208 0.168 6 0.015 0.139–0.187 0.165 6 0.018 0.139–0.200
IH/CL 0.145 6 0.023 0.092–0.194 0.139 6 0.020 0.083–0.180 0.143 6 0.024 0.106–0.189 0.143 6 0.018 0.093–0.166
IP/CL 0.023 6 0.018 0.0–0.072 0.020 6 0.015 0.0–0.064 0.015 6 0.013 0.0–0.039 0.037 6 0.013 0.020–0.062
IAB/CL 0.264 6 0.019 0.208–0.299 0.283 6 0.016 0.243–0.330 0.290 6 0.013 0.269–0.314 0.288 6 0.016 0.259–0.320
IF/CL 0.045 6 0.020 0.008–0.088 0.040 6 0.021 0.0–0.134 0.044 6 0.018 0.025–0.072 0.057 6 0.016 0.033–0.084
IAN/CL 0.313 6 0.018 0.268–0.367 0.328 6 0.018 0.244–0.375 0.324 6 0.015 0.302–0.350 0.313 6 0.021 0.274–0.356
SH/CL 0.477 6 0.024 0.417–0.532 0.457 6 0.036 0.367–0.546 0.496 6 0.022 0.466–0.532 0.479 6 0.025 0.439–0.531
BL/CL 0.309 6 0.027 0.231–0.364 0.307 6 0.016 0–0.134 0.320 6 0.012 0.293–0.336 0.310 6 0.014 0.288–0.334
CW/CL 0.734 6 0.022 0.656–0.777 0.671 6 0.023 0.244–0.375 0.718 6 0.031 0.662–0.775 0.715 6 0.033 0.647–0.768
CW/PL 0.771 6 0.028 0.692–0.819 0.691 6 0.030 0.620–0.756 0.740 6 0.030 0.694–0.788 0.732 6 0.040 0.654–0.793
IAB/SH 0.554 6 0.039 0.438–0.648 0.622 6 0.052 0.492–0.793 0.586 6 0.032 0.540–0.643 0.602 6 0.023 0.576–0.654
IAN/SH 0.656 6 0.041 0.545–0.773 0.720 6 0.054 0.603–0.864 0.655 6 0.048 0.584–0.748 0.654 6 0.047 0.601–0.745
a
CL ¼ maximum carapace length; PL ¼ maximum plastron length; CW ¼ maximum carapace width; SH ¼ maximum shell height; BL ¼ bridge length;
GL ¼ gular length; IH ¼ interhumeral seam length; IP ¼ interpectoral seam length; IAB ¼ interabdominal seam length; IF ¼ interfemoral seam length;
IAN ¼ interanal seam length.
0 CHELONIAN CONSERVATION AND BIOLOGY, Volume 23, Number 1 – 2024

Table 3. Character comparisons of Pacific and Atlantic populations of male Kinosternon cruentatum. See Table 2 footnote for char-
acter abbreviations.

Pacific populations East Atlantic populations Veracruz population Tampico population

(n ¼ 46) (n ¼ 80) (n ¼ 5) (n ¼ 10)
Ratio Mean 6 1 SD Range Mean 6 1 SD Range Mean 6 1 SD Range Mean 6 1 SD Range
PL/CL 0.919 6 0.023 0.884–0.984 0.939 6 0.026 0.890–1.032 0.944 6 0.037 0.896–1.000 0.954 6 0.022 0.921–0.987
GL/CL 0.154 6 0.015 0.120–0.190 0.158 6 0.018 0.109–0.194 0.151 6 0.024 0.119–0.186 0.159 6 0.015 0.128–0.175
IH/CL 0.146 6 0.027 0.064–0.236 0.131 6 0.019 0.086–0.173 0.132 6 0.022 0.097–0.154 0.144 6 0.017 0.111–0.172
IP/CL 0.025 6 0.015 0.0–0.061 0.025 6 0.011 0.0–0.048 0.038 6 0.010 0.027–0.052 0.031 6 0.015 0.00–0.055

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IAB/CL 0.242 6 0.019 0.205–0.296 0.263 6 0.011 0.246–0.309 0.265 6 0.016 0.248–0.285 0.265 6 0.011 0.254–0.287
IF/CL 0.061 6 0.018 0.032–0.112 0.049 6 0.016 0.010–0.103 0.059 6 0.017 0.037–0.083 0.057 6 0.017 0.039–0.087
IAN/CL 0.291 6 0.018 0.256–0.333 0.302 6 0.016 0.266–0.346 0.303 6 0.019 0.274–0.325 0.307 6 0.017 0.280–0.334
SH/CL 0.428 6 0.028 0.342–0.483 0.405 6 0.024 0.361–0.486 0.473 6 0.043 0.430–0.545 0.431 6 0.016 0.397–0.461
BL/CL 0.286 6 0.026 0.214–0.355 0.291 6 0.014 0.255–0.324 0.301 6 0.043 0.292–0.317 0.296 6 0.012 0.284–0.319
CW/CL 0.685 6 0.030 0.630–0.759 0.643 6 0.022 0.587–0.697 0.678 6 0.018 0.663–0.697 0.665 6 0.014 0.638–0.685
CW/PL 0.745 6 0.037 0.687–0.853 0.685 6 0.029 0.611–0.744 0.719 6 0.042 0.697–0.740 0.698 6 0.025 0.657–0.732
IAB/SH 0.568 6 0.067 0.446–0.822 0.650 6 0.053 0.538–0.733 0.562 6 0.033 0.522–0.604 0.616 6 0.037 0.577–0.700
IAN/SH 0.683 6 0.053 0.567–0.789 0.768 6 0.063 0.618–0.884 0.643 6 0.042 0.597–0.690 0.713 6 0.039 0.653–0.786

Data from the holotypes of K. mexicanum and K. cruenta-
tum (measured by J.B.I.) were included in all sex-appro-
priate analyses, but without a drainage designation, for a
posteriori classification. The type of K. triliratum has
been lost (Malnate 1971), and hence could not be
included in this study.
RESULTS
Character ratio analyses (Tables 1–5) demonstrated
considerable difference between the Pacific and Atlantic
populations. Turtles from Pacific slope populations
have, on average, longer carapace and plastron lengths,
greater carapace widths and height, and shorter interab-
dominal and interanal seams than turtles from Atlantic
slopes (Tables 1–5; Figs. 2–4). Furthermore, this analy-
sis indicated that the female type of K. cruentatum was
nested among Atlantic coast females (Table 5; Figs. 2–
3), and that the male type of K. mexicanum was nested
among Pacific coast males (Table 4; Fig. 2–3).
For the initial Pacific vs. Atlantic DFA analyses of
both males and females, 100% of the variance was
explained by the first canonical axis. The canonical plots
clearly demonstrated that populations of K. cruentatum
from the Pacific and Atlantic versants differed signifi-
cantly in morphometric hyperspace (Fig. 5). The DFA for
males grouped the male type of K. mexicanum with turtles
from the Pacific slope with a probability of 100%, and the
DFA for females grouped the female type of K. cruenta-
tum with turtles from the Atlantic slope with the same
probability.
The second set of DFA analyses (comparing the 4
allopatric populations) again demonstrated the morpho-
metric divergence between Pacific and Atlantic popula-
tions, but also revealed substantial variation across the
Atlantic versant populations (Figs. 6–7). With 100%
probability the DFA again placed the male type of K.
mexicanum with Pacific coast turtles, and the female
type of K. cruentatum with the Atlantic coast turtles,
specifically within the eastern Atlantic subpopulation.
Finally, the third set of DFA analyses of just the 3 allo-
patric Atlantic subpopulations revealed that the eastern
Atlantic subpopulation was the most divergent of the 3
(especially for females; Figs. 8–9), and again placed the
holotype of K. cruentatum among the eastern Atlantic
specimens.
Bivariate plots of the most discriminating charter
ratios (Fig. 10–11) also suggest that the Veracruz and

Table 4. Character ratios most useful in discriminating Pacific versus Atlantic populations of male Kinosternon cruentatum, along
with measurements of the male holotype of K. mexicanum. Comparisons made with Student’s t-test (t) and Mann-Whitney U test
(Z). See Table 2 footnote for character abbreviations.

Pacific populations (n ¼ 46) Atlantic populations (n ¼ 96)

Ratio mexicanum type Mean 6 1 SD Mean 6 1 SD t/Z p
CW/PL 0.723 0.746 6 0.036 (0.687–0.853) 0.689 6 0.028 (0.610–0.743) 9.5/7.6 , 0.0001
IF/CL 0.077 0.061 6 0.018 (0.032–0.112) 0.051 6 0.016 (0.010–0.103) 3.5/3.3 0.0002/0.001
IAN/CL 0.286 0.291 6 0.018 (0.256–0.333) 0.311 6 0.016 (0.266–0.346) 6.3/5.6 , 0.0001
SH/CL 0.459 0.428 6 0.028 (0.342–0.483) 0.412 6 0.030 (0.361–0.545) 3.1/3.7 0.0012/0.0002
IAB/SH 0.559 0.568 6 0.067 (0.446–0.822) 0.644 6 0.045 (0.523–0.734) 8.0/6.8 , 0.0001
IAN/SH 0.459 0.683 6 0.053 (0.567–0.789) 0.758 6 0.062 (0.596–0.884) 7.0/6.1 , 0.0001
 IVERSON AND BERRY — Kinosternon cruentatum Morphometrics 0

Table 5. Character ratios most useful in discriminating Pacific vs. Atlantic populations of female Kinosternon cruentatum, along
with measurements of the female holotype of K. cruentatum. Comparisons made with Student’s t-test (t) and Mann-Whitney U-test
(Z). See Table 2 footnote for character abbreviations.

Pacific populations (n ¼ 70) Atlantic populations (n ¼ 135)

Ratio cruentatum type Mean 6 1 SD Mean 6 1 SD t/Z p
CW/PL 0.683 0.771 6 0.028 (0.693–0.819) 0.698 6 0.035 (0.619–0.793 14.9/10.4 , 0.0001
IF/CL 0.042 0.045 6 0.020 (0.008–0.110) 0.042 6 0.021 (0–0.134) 1.1/1.0 0.03/0.32
IAN/CL 0.322 0.313 6 0.018 (0.268–0.367) 0.326 6 0.018 (0.244–0.375) 4.8/4.9 , 0.0001
SH/CL 0.498 0.477 6 0.024 (0.417–0.532) 0.462 6 0.036 (0.367–0.546) 3.2/3.0 0.0009/0.003

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IAB/SH 0.595 0.553 6 0.039 (0.439–0.649) 0.617 6 0.049 (0.492–0.793) 9.4/8.3 , 0.0001
IAN/SH 0.647 0.656 6 0.041 (0.545–0.772) 0.708 6 0.058 (0.571–0.863) 6.7/6.4 , 0.0001

Tampico embayment subpopulations are morphometrically
similar (despite being separated by the eastern terminus
of the Transverse Volcanic Arc in north-central Vera-
cruz), but that they are distinct from the eastern Atlantic
populations.
DISCUSSION
Our morphometric results support the recognition of
separate species for the Pacific and Atlantic populations
previously referred to under the name K. cruentatum, as
recently proposed by Hurtado-Gómez et al. (2024). In
addition, our analyses also identified the female holotype
of K. cruentatum as originating from the Atlantic versant
and that of the male K. mexicanum as originating from
Pacific drainages. Given the genetic data provided by
Hurtado-Gómez et al. (2024) showing the reciprocal
monophyly of turtles from those 2 versants, an estimated
divergence time of 2.4 million years, and an uncorrected
p distance of 2.9% for the cytochrome b marker, the spe-
cies recognition of K. cruentatum from the Atlantic and
K. mexicanum from the Pacific is strongly supported.
Future phylogenetic work will be necessary to evaluate
whether K. cruentatum (sensu stricto) exhibits subspecific
variation in the Atlantic versant. However, because Vera-
cruz has been a major trade center at least since it was
founded by Hernán Cortez in 1519, market specimens
should not be included in those analyses (as some were in
this study).

Figure 3. Bivariate plot of character ratios useful in discrim-

Figure 2. Bivariate plot of character ratios useful in discrim-
inating Pacific from Atlantic populations of Kinosternon
cruentatum (males above; females below). Abbreviations are
IAN (interanal seam length), SH (shell height), CW (cara-
pace width), and PL (maximum plastral length). Male type
(0.72, 0.62) is Kinosternon mexicanum (see also Table 4);
female type (0.68, 0.65) is K. cruentatum (see also Table 5).
inating Pacific from Atlantic populations of Kinosternon
cruentatum (males above; females below). Abbreviations
are IAB (interabdominal seam length), SH (shell height),
CW (carapace width), and PL (maximum plastral length).
Male type (0.72, 0.56) is Kinosternon mexicanum (see also
Table 4); female type (0.68, 0.60) is K. cruentatum (see also
Table 5).
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Figure 4. Photographs of the plastron of a typical female Kinosternon cruentatum from the Atlantic versant (Manuel,
Tamaulipas; upper left; photo by J.B.I.), that of the holotype of K. cruentatum (MNHN 1759; upper right; courtesy of Roger
Bour), that of a typical male Kinosternon mexicanum (Suciate, Chiapas; lower left; courtesy of Eduardo Reyes-Grajales), and
that of the type of K. mexicanum (ANSP 90; lower right; photo by J.B.I.). Note the relatively longer interabdominal and
interanal seams of K. cruentatum (see text).

The collector and date of collection of the type of
K. cruentatum (Museum national d’Histoire naturelle, Paris,
MNHN-RA-0.1759, a female) are unknown, and its type
locality was reported as “Amerique septentrionale” (north-
ern America, presumably in contrast to South America;
Berry and Iverson 2001; Bour 2004). Hence, as indicated
above, the name cruentatum should be restricted to Atlantic
populations (the Atlantic Red-cheeked Mud Turtle). This
action invalidates the type locality restriction of cruentatum
to San Mateo del Mar, Oaxaca, by Smith and Taylor
(1950a, 1950b). We propose that the type locality of
K. cruentatum be restricted to Pueblo Nuevo X-Can,
Quintana Roo, Mexico (20 520 900 N, 87 360 1100 W), from
which at least 20 specimens are known (University of
Utah 9563–80, 11874, and American Museum of Natu-
ral History 93244).
The type of K. mexicanum (Academy of Natural
Sciences Philadelphia 90, a male) was collected by Wil-
liam S. Pease on an unspecified date in “Mexico”
(LeConte 1854). However, it was likely collected or
 IVERSON AND BERRY — Kinosternon cruentatum Morphometrics 0

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Figure 5. Distribution of canonical discriminant function (DF) scores on the first canonical axis (representing 100% of the var-
iance) for female (above) and male (below) specimens of Kinosternon cruentatum from the Pacific and Atlantic drainages of
Mesoamerica, including the holotype of K. cruentatum among the males, and the holotype of Kinosternon mexicanum among
the females (both in green). Holotypes were unassigned a priori and classified a posteriori. DF equation for males:
Score ¼ 3 3 PL/CL þ 13 3 GL/CL þ 8 3 IH/CL þ 27 3 IP/CL þ 30 3 IAB/CL þ 15 3 IF/CL þ 47 3 IAN/CL  20 3 SH/CL þ 17 3 BL/
CL  24 3 CW/CL  10.605. DF equation for females: Score ¼ 24 3 PL/CL þ 23 3 GL/CL þ 25 3 IH/CL þ 25 3 IP/CL  12 3 IAB/
CL þ 1 3 IF/CL  18 3 IAN/CL þ 15 3 SH/CL  4 3 BL/CL þ 36 3 CW/CL  6.707. CL ¼ maximum carapace length; PL ¼
maximum plastron length; CW ¼ maximum carapace width; SH ¼ maximum shell height; BL ¼ bridge length; GL ¼ gular
length; IH ¼ interhumeral seam length; IP ¼ interpectoral seam length; IAB ¼ interabdominal seam length; IF ¼ interfemoral
seam length; IAN ¼ interanal seam length.
0 CHELONIAN CONSERVATION AND BIOLOGY, Volume 23, Number 1 – 2024
Figure 6. Distribution of canonical discriminant function scores
on the first 2 canonical axes (representing 83.8% and 13.5% of
total variation, respectively) for male Kinosternon cruentatum
from the Pacific (1; red), eastern Atlantic (2; green), Veracruz
(3; dark blue), and Tampico drainages (4; magenta). Group cen-
troids (large yellow squares) are numbered. The male type of
Kinosternon mexicanum (light blue square) was ungrouped and
classified a posteriori with the Pacific turtles with 100% probability.
purchased in 1847, when, on behalf of the Philadelphia
Academy of Natural Sciences, Pease accompanied the
US military march from Veracruz to Jalapa and on to
Mexico City during the Mexican–American War (Cassin
Figure 7. Distribution of canonical discriminant function scores
on the first 2 canonical axes (representing 66.3% and 31.8% of
total variation, respectively) for female Kinosternon cruentatum
from the Pacific (1; red)), eastern Atlantic (2; green), Veracruz
(3; dark blue), and Tampico drainages (4; magenta). Group cen-
troids (large yellow squares) are numbered. The female type of
K. cruentatum (light blue) was ungrouped and classified a poste-
riori with the Pacific turtles with 100% probability (and falls on
the plot with cluster 2; coordinates, 0.92, 0.72).
Downloaded from http://meridian.allenpress.com/ccb/article-pdf/doi/10.2744/CCB-1589.1/3362993/10.2744_ccb-1589.1.pdf by Earlham College, John B. Iverson on 12 May 2024
Figure 8. Distribution of canonical discriminant function
scores on the first 2 canonical axes (representing 88.7% and
11.3% of total variation, respectively) for male Kinosternon
cruentatum from the eastern Atlantic (2; red), Veracruz (3;
green), and Tampico drainages (4; dark blue). Group centroids
(large magenta squares) are numbered.
1848–1849; Pease 1848–1849, 1849). There is no evi-
dence that the expedition traveled to the Pacific coast,
but the holotype could have been purchased in a mar-
ket. Nevertheless, based on our morphometric data,
and pending genetic data from both holotypes, we recom-
mend recognizing Pacific populations as K. mexicanum
(the Pacific Red-cheeked Mud Turtle), and retaining the
restricted type locality designated by Smith and Taylor
(1950a, 1950b).
Figure 9. Distribution of canonical discriminant function
scores on the first 2 canonical axes (representing 91.1% and
8.9% of total variation, respectively) for female Kinosternon
cruentatum from the eastern Atlantic (2; red), Veracruz (3;
green), and Tampico drainages (4; dark blue). The female type
of K. cruentatum was ungrouped and classified a posteriori
with the eastern Atlantic turtles with 100% probability (coor-
dinates, 0.92, 0.72). Group centroids (large light blue
squares) are numbered.
 IVERSON AND BERRY — Kinosternon cruentatum Morphometrics
Figure 10. Bivariate plot of character ratios useful in discrimi-
nating among Atlantic populations of male Kinosternon cruen-
tatum. Abbreviations are IAB (interabdominal seam length),
IAN (interabdominal seam length), SH (shell height), CW
(carapace width), and PL (maximum plastral length).
Figure 11. Bivariate plot of character ratios useful in discrimi-
nating Atlantic populations of female Kinosternon cruentatum.
Abbreviations are IAB (interabdominal seam length), IAN
(interabdominal seam length), SH (shell height), CW (cara-
pace width), and PL (maximum plastral length). Female holo-
type of K. cruentatum falls in the east Atlantic cluster (see
also Table 5).
0
As noted by Hurtado-Gómez et al, (2024), the biogeo-
graphic history of the subgenus Kinosternon (including albo-
gulare, cruentatum, oaxacae, chimalhuaca, and integrum)
has been primarily along the Pacific versant of Central
America and Mexico. This suggests that the common
ancestor of K. cruentatum and K. mexicanum likely dis-
persed from the Pacific to the Atlantic versant, probably
across the relatively low Isthmus of Tehuantepec. From
there it apparently migrated northward to the Tampico
Downloaded from http://meridian.allenpress.com/ccb/article-pdf/doi/10.2744/CCB-1589.1/3362993/10.2744_ccb-1589.1.pdf by Earlham College, John B. Iverson on 12 May 2024
embayment and eastward as far as the Yucatan penin-
sula and Belize. That extensive Atlantic range was
apparently subsequently fragmented (Fig. 1), perhaps
by the eastward orogeny of the Transverse Volcanic Arc
in north central Veracruz, and the Tabasco lowlands,
where it appears to be absent.
ACKNOWLEDGMENTS
Iverson’s research was funded by the American
Philosophical Society, the American Museum of Natural
History, the Joseph Moore Museum of Earlham College,
and the National Science Foundation. The Paris Museum
and the Academy of Natural Sciences of Philadelphia
generously loaned J.B.I. the type specimens over 40 yrs ago.
Berry’s research was funded by the University of Utah and
Elmhurst College. Curators of 19 herpetological collections
provided access to the material used in this study (full
details in Berry 1978). Anders Rhodin generously produced
the map of the current range of these populations, updated
from TTWG (2021). Eduardo Reyes Grajales and Roger
Bour provided photographs.
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Received: 27 April 2023
Revised and Accepted: 23 October 2023
Published Online: 25 April 2024
Handling Editor: Jeffrey E. Lovich