2004

Asiatic Herpetological ResearchVol. 10, pp. 53-109

A Review of the Comparative Morphology of Extant Testudinoid Turtles(Reptilia: Testudines)
WALTER G. JOYCE1 AND CHRISTOPHER J. BELL2
1Department of Geology and Geophysics, Yale University, New Haven, CT 06511

E-mail: walter.joyce@yale.edu
2Department of Geological Sciences, The University of Texas at Austin, Austin, TX 78712

E-mail: cjbell@mail.utexas.edu
Abstract. - With an expansive geographic distribution, an excellent fossil record, and over 140 recognized extant
species, testudinoid turtles constitute one of the most diverse and widespread clades of turtles. The current understanding of the distribution of morphological characters among testudinoid turtles is poor. Improved knowledge will
help to facilitate accurate identification of fossil remains, and to provide a reliable morphological data set for phylogenetic analyses. We provide a critical review of skeletal and scute characters commonly utilized in previous systematic analyses of Testudinoidea. Description and illustration of character states, discussion of their distribution within
Testudinoidea, and polarity determinations for 93 characters are provided. Our preliminary results indicate that ontogenetic changes in skeletal structure are an important source of variation within Testudinoidea. Sexual variation, ontogenetic variation, and intra- and inter-population variation are inadequately documented for most testudinoid taxa.
Furthermore, data matrices of morphologic characters in the existing literature must be carefully reconsidered.
Previously published morphologic data provide reasonably strong support for the monophyly of 'Testudinidae.' Strong
morphologic support for a monophyletic 'Emydidae' is lacking, and 'batagurid' monophyly has not been rigorously
tested in the literature. Because a new research cycle centered on testudinoid phylogeny is now under way, it is essential to critically re-examine the underlying assumptions and working hypotheses that have governed this field of study
over the last 20 years.
Key words. - Testudines, Testudinoidea, Testudinidae, Emydidae, Bataguridae, Geoemydidae, morphology, systematics.

Introduction
Pond turtles and land tortoises (collectively,
Testudinoidea) form one of the largest and most widespread clades of living turtles, with more than 140 extant
species and an almost worldwide distribution. The discovery and description of many new fossil testudinoids
in the last half century, combined with the emergence
and ascendancy of molecular techniques in systematics,
provide new opportunities to explore the evolutionary
history of the group in unprecedented detail.
Concomitant with the appearance of these new data sets
and analytical techniques comes an increasing appreciation for conservation efforts to preserve these turtle lineages and help to secure their future in the face ofincreasing human predation and habitat encroachment.
This is true especially for the Asian representatives ofthis clade (e.g., van Dijk et al., 2000) but also is relevant
at a more generalized and inclusive level (e.g., Rhodin,
2000).
Our recent attempts to diagnose fossil testudinoids
reliably and to place them within a phylogenetic context
led to the recognition that a critical re-evaluation of morphological data and purported synapomorphies for the© 2004 by Asiatic Herpetological Researchsubclades of testudinoid turtles is desirable. A morethorough understanding of morphological data sets will
provide not only a means by which molecular trees may
be independently assessed, but also will form an essential foundation for diagnosing and interpreting fossil
specimens. This in turn will facilitate the integration offossil taxa into future systematic analyses, and will
enhance our understanding of the paleobiogeography
and divergence times of extant lineages.
The recent flurry of published works appears to represent the beginning of a new research cycle (sensu
Kluge, 1991) in testudinoid systematics. We suggest thatan important part of this cycle will be a critical re-examination of the working hypotheses that have governed
testudinoid systematics since the publication of
McDowell’s (1964) seminal work on the group. A key
component of this will be the assessment of fundamental, often unstated, assumptions that underlie current
hypotheses of relationship. Our contribution to thisresearch cycle is the first critical reappraisal of morphological characters applied to testudinoid systematics
since the work of Hirayama (1985). The emerging
improvement in our understanding of testudinoid relationships based on molecular sequence data will certain-

Vol. 10, p. 54

Asiatic Herpetological Researchly result in numerous new questions (e.g., regarding
paleobiogeography, the timing of sequence and evolutionary divergences, and the evolution of morphological
adaptations) that will demand a clearer understanding of
testudinoid morphology.
The purpose of this paper is to present a preliminary
revision and discussion of the morphological characters
previously utilized in investigations of testudinoid systematics. Our goal here is not to produce a phylogenetic
hypothesis (indeed, we deliberately eschew such a production), but rather to evaluate the morphological datathat have been, and will be, used to generate such
hypotheses. To enhance our discussion and facilitate
improved communication about testudinoid morphology, we provide illustrations of all characters states we
discuss.
Abbreviations. - Institution and collection abbreviations: CAS, California Academy of Sciences, San
Francisco, California; CM, Carnegie Museum of Natural
History, Pittsburgh, Pennsylvania; FMNH, Field
Museum of Natural History, Chicago, Illinois; IVPP,
InstituteofVertebrate
Paleontology
andPaleoanthropology, Beijing, China; KU, The Universityof Kansas Natural History Museum, Lawrence, Kansas;
LMNH, Louisiana Museum of Natural History, Baton
Rouge, Louisiana; MCZ, Museum of Comparative
Zoology,
Harvard
University,
Cambridge,
Massachusetts; TNHC, Texas Natural History
Collections, Texas Memorial Museum, Austin, Texas;
TUMNH, Tulane University Museum of Natural
History, New Orleans, Louisiana; YPM, Yale Peabody
Museum, New Haven, Connecticut.
Abbreviations used in figures: AB, abdominal
scute; af, articular facet; an, angular; bo, basioccipital;CE, cervical scute; co, costal bone; ent, entoplastron;
epi, epipterygoid; fdm, foramen dentofaciale majus; fr,
frontal; fpp, foramen palatinum posterius; HU, humeralscute; hyo, hyoplastron; hypo, hypoplastron; ju, jugal;
MA, marginal scute; mx, maxilla; ne, neural bone; pa,parietal; pal, palatine; PEC, pectoral scute; pf, prefrontal; PL, pleural scute; pm, premaxilla; po, postorbital; pt, pterygoid; qj, quadratojugal; qu, quadrate; VE,
vertebral scute; vf, vomerine foramen; vo, vomer.

Material and Methods
We examined 309 testudinoid specimens representing
93 species, but focused our efforts on 46 representative
species. The list of specimens examined is provided inAppendix 1. Turtle shell nomenclature follows Zangerl(1969) and cranial nomenclature follows Gaffney(1972). Of the 46 focal species, most were recognized as

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valid species by Ernst and Barbour (1989), with the
exception of texana, which they placed within ‘concinna.’ Generic allocations for testudinoid species varied
widely over the last 50 years and are subject to differing
opinions today, particularly because the monophyly ofmany testudinoid genera remains untested. We consequently suppress the use of generic names wherever possible and use species epithets only. This procedure also
has the advantage of precisely associating observations
with species only instead of higher taxonomic categories. Most extant turtles have distinct species names,
but among those turtles discussed in this review thespecies epithets insculpta, nelsoni, oculifera, ornata,and platynota each appear twice (insculpta underGlyptemys and Carettochelys; nelsoni under Pseudemys
and Terrapene; oculifera under Graptemys andPsammobates; ornata under Pseudemys and Terrapene;
platynota under Geochelone and Notochelys). For clarity in these instances, we indicate our usage with a single-letter generic abbreviation. A complete list of all currently recognized testudinoid species and all outgroupspecies used herein is provided in Appendix 2 together
with a list of their various generic assignments used in
the last 50 years.
Our use of the classic higher categories is always
restricted to their phylogentic crown. ‘Emydinae’ (sensu
McDowell, 1964) are also referred to as ‘Emydidae,’‘emydids,’ or North American pond turtles;‘Batagurinae’ (sensu McDowell, 1964) as ‘Bataguridae,’‘batagurids’ or Asian pond turtles; and ‘Testudinidae’(sensu McDowell, 1964) as ‘testudinids’ or (land) tortoises. We make no a priori assumptions of monophyly
for any of these categories, and retain single quotations
around these names throughout the text to emphasize
our uncertainty.
We attempted to examine most significant morphological characters commonly utilized in systematic studies of testudinoids, but the majority of our observations
concern the skeletal system and scute characters. Almost
all characters were derived from the literature. Major
sources for each category were: ‘Batagurinae’(Hirayama, 1985; McCord et al., 1995; Yasukawa et al.,2001); ‘Testudinidae’ (Crumly, 1982, 1985, 1994); and‘Emydinae’ (Gaffney and Meylan, 1988; Burke et al.,1996). Additional characters were also found in
Mlynarski (1976), Shaffer et al. (1997), and othersources cited in the character discussions. With few
exceptions, morphological features were examined on
specimens themselves; evaluations based on previously
published literature are indicated where applicable.
Sexual dimorphism, ontogenetically influenced polymorphisms, and geographic variation in morphology are
not well explored in testudinoid turtles. These areas arein need of much more research. A full exploration of2004

Asiatic Herpetological Researchsuch variation is beyond the scope of this work, but weare able to make some preliminary observations regarding morphological change through ontogeny in someanatomical systems.
Polarizing characters with the help of outgroups
proved to be a difficult task, mostly because all relevant
extant sister taxa are highly specialized after more than
65 million years of independent evolution. Furthermore,
hypotheses of the systematic relationships of the majorgroups of cryptodires reveal a highly unstable picture(e.g., Bickham, 1981; Gaffney, 1975, 1985; Gaffney etal., 1991; Shaffer et al., 1997) making it impossible to
make any a priori decisions regarding the succession ofoutgroups. We consequently assessed polarity for mostcharacters by examining select outgroup taxa and theingroup taxa. Where polarity is not clear from outgroupcomparison, we sometimes relied upon ingroup commonality. To allow full transparency, we discuss every
polarity decision at the end of each character description.
Outgroup taxa include the cryptodires caretta,
odoratus, serpentina, and spinifera, and the pleurodiresgibba, siebenrocki, subglobosa, and subrufa. For a number of characters, especially of the shell, neither ingroup
nor outgroup analysis of extant taxa proved useful. In
these instances, polarity was based on literature descriptions of the “lindholmemydid” taxa Gravemys,
Lindholmemys, and Mongolemys. These Cretaceous,
Asian, fossil taxa are not well described in the literature,but sufficient material and description exists to use thesetaxa to help polarize character states (e.g., Khosatzky
and Mlynarski, 1971; Sukhanov, 2000; Danilov andSukhanov, 2001). The group may not be monophyletic,
but putative members currently are hypothesized to sit
along the phylogentic stem of Testudinoidea (Danilov
and Sukhanov, 2001). We purposefully did not use thefossil taxon ‘Echmatemys’ as an outgroup taxon(Hirayama, 1985), because its phylogenetic position outside of Testudinoidea or even ‘Batagurinae’ is not sufficiently demonstrated.
All figures were produced using digital photography and processed using Adobe Photoshop. Images weredigitally enhanced using the burn and burnish tools andthe unsharp mask filter option.
Taxonomic and Systematic Background. - Despite theincreased attention directed towards testudinoids by scientists, hobbyists, and nonprofessional enthusiasts in thelast thirty-five years, our collective conceptualization of
the higher-level (beyond the specific and generic) systematics within this clade remained virtually unchanged
since the work of McDowell (1964). The various taxonomies in current use owe their existence in large partto historical contexts that are not well appreciated byVol. 10, p. 55

many authors. A brief summary is given here.
During the second half of the 19th century, a number of attempts were made to work out higher-level testudinoid relationships and to apply taxonomic conventions that were designed (to a greater or lesser extent) tocommunicate conceptualizations of these relationships.
In his synopsis on the turtles of North America, Agassiz(1857) united all pond turtles into the Emydoidae andsubdivided this group into a three monotypic subfamilies (Deirochelyoidae for reticularia, Evemydoidae forblandingii, Cistudinina for T. ornata and carolina) followed by the subfamilies Clemmydoidae (for G. insculpta, guttata, marmorata, and muhlenbergii) and
Nectemydoidae (for those species currently placed in thegenera Pseudemys, Trachemys, Graptemys, Malaclemys
and Chrysemys). Most land tortoises were placed in
Testudinidae by Theobald (1868); he also included all‘leaf turtles and tortoises’ (e.g., amboinensis, emys, dentata, grandis, tricarinata) in Geoemydidae, and an
eclectic group of aquatic turtles, including megacephalum, serpentina, kinosternids, and all remaining
testudinoids, in Emydidae. Subsequently, all land tortoises (including emys and impressa) were united in the
Testudinidae by Gray (1870). Those species currently
placed in Pseudemys and Trachemys were assigned byGray (1870) to the Pseudemydae; the Asian taxa baska,
borneoensis, thurjii, kachuga, and ocellata wereassigned to the Bataguridae, and all hinged pond turtles,‘true terrapins,’ and ‘snail-eating pond turtles’ to theHolarctic families Cistudinidae, Emydidae, andMalaclemmydae, respectively.
Despite these early attempts, most subsequentauthors (e.g., Boulenger, 1889; Siebenrock, 1909;
Lindholm, 1929; Smith, 1931; Bourret, 1941) ignored(or were unaware of) these works and simply divided all
testudinoid turtles into two speciose subgroups: tortoises (Testudinidae or Testudininae) and pond turtles(Emydidae or Emydinae). This situation remained static
for nearly 100 years until the comprehensive and influential work of McDowell (1964). He not only divided all
known pond turtles into several species complexes(‘Hardella,’ ‘Batagur,’ ‘Orlitia,’ ‘Geoemyda,’‘Chrysemys,’ ‘Deirochelys,’ and ‘Emys’ complexes), butalso concluded that pond turtles can be divided clearly
into two subgroups, the predominantly North American‘Emydinae’ and the Asian and central American‘Batagurinae.’ Furthermore, McDowell (1964) reasoned
that tortoises are not the sister group of pond turtles, butrather were likely derived from a ‘batagurine’ ancestor.
These conclusions were later corroborated by the first,
and to date only, comprehensive morphological cladistic
analysis of ‘batagurine’ systematics (Hirayama, 1985).
The influence of McDowell’s (1964) work is best
understood when considering its continuous impact on

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Asiatic Herpetological Researchsubsequently proposed phylogenies. Despite differencesof opinion regarding generic- and species-level systematic arrangements, virtually all major synthetic works in
the last thirty years followed McDowell’s (1964) subdivision of pond turtles into the ‘Batagurinae’ and‘Emydinae’ (e.g., Mlynarski, 1976; Pritchard, 1979;
Ernst and Barbour, 1989), even though ‘Batagurinae’
may best be regarded as a paraphyletic taxon(McDowell, 1964; Hirayama, 1985). The fundamental
division proposed by McDowell is also reflected in more
recent studies centered on using various molecular techniques to elucidate phylogeny, the majority of which
dealt with treatments of in-group relationships within
one or the other of McDowell’s groups (Sites et al.,
1984; Bickham et al., 1996; Carr and Bickham, 1986;
Wu et al., 1999; McCord et al., 2000; Feldman andParham, 2002; Honda et al., 2002; Iverson et al., 2002;
Stephens and Wiens, 2003).
Admittedly, the list of autapomorphic characterscompiled by Crumly (1985) for the ‘Testudininae’ compellingly corroborates the hypothesis of tortoise monophyly. However, most characters that currently unite‘Emydinae’ or ‘Batagurinae + Testudininae’ seem tosupport these groupings weakly, because the derived
states typically are not found within all species of the
ingroup and commonly also are observed in species of
the alleged sister group (e.g., Hirayama, 1985; Gaffney
and Meylan, 1988). In addition, several of the charactersthat purportedly distinguish ‘Batagurinae +Testudininae’ from the ‘Emydinae’ probably should beconsidered primitive for the entire group (Gaffney andMeylan, 1988). Even if some characters do successfully
unite a group, monophyly is not established until the
involved characters are demonstrated to be derived within Testudinoidea. Furthermore, the simple demonstration of monophyly for a given group does not automatically imply that it must be the sister to the remainingtaxa. For instance, ‘Emydinae’ may be monophyletic,
but monophyly does not necessarily demand that
‘Emydinae’ be regarded as the sister to ‘Batagurinae +Testudininae.’ It is at least plausible that ‘Emydinae’ issituated within ‘Batagurinae,’ a possibility that is notadequately explored and tested in the literature.
Similarly, most of the groupings considered byGray (1855, 1870) and Agassiz (1857) were not discussed in recent literature, even though they might bevalid. For instance, given the considerable list of morphological similarities that are shared by hinged turtlesof the New and Old World (e.g., development of a plastral hinge, reduction of posterior neural elements, fusionof the femoral trochanter, great reduction of the temporal arch) perhaps Gray (1870) was truly visionary inuniting these turtles as the ‘Cistudinidae.’ Only a global
cladistic analysis with no a priori assumption regarding
internal relationships can evaluate these alternatives and2004

produce testable results. It is toward this end that weoffer our critical reappraisal of morphological charactersin testudinoids.

Results and Discussion
Cranium(1) Shape of the fissura ethmoidalis; 0 = narrow or
closed, keyhole-shaped, Fig. 1; 1 = very wide, Fig. 2(modified from Crumly, 1982, 13; Hirayama, 1985, 1;
McCord et al., 1995, 5).
The general configuration of the fissura in ‘emydids’ and ‘batagurids’ is keyhole-shaped (McDowell,1964). Different proportions and widths are apparent(especially in the ventral part of the fissura), and werescored by Hirayama (1985) and Crumly (1982) as discrete character states. Our survey of many taxa reveals
morphological intermediates, and the expression of various states appears to have an ontogenetic component inwhich younger individuals exhibit a relatively larger fissura, which corresponds to a less-ossified nasal cavity.
However, a rather significant morphological gap can beobserved between tortoises and pond turtles. For the
purpose of this review, we lumped Hirayama’s (1985)
states into our state 0, and Crumly’s (1982) into our state
1. The scoring presented by Hirayama (1985) and
McCord et al. (1995) permitted phylogenetic resolution
within ‘batagurids’, and that of Crumly (1982) within
tortoises. Our revised scoring permits support only forthe hypothesis of a monophyletic ‘Testudinidae.’
Polarity: Pleurodires lack a defined fissura ethmoidalis. A keyhole-shaped fissura ethmoidalis is present in spinifera, odoratus, caretta, serpentina, andMongolemys, and this condition is considered primitive
for testudinoids.
(2) Medial inflection of the inferior descending processes of the frontal; 0 = absent, or very small, Fig. 3; 1 =present, well-developed, medial contact present oralmost present, Fig. 4 (modified from Hirayama, 1985,
2).
In most turtles, a gutter (the sulcus olfactorius) isformed along the ventral surface of the frontals. Thisgutter transmits the olfactory nerve. The lateral rims of
the sulcus sometimes form processes that descend ventromedially to surround the nerve from below(McDowell, 1964). According to Hirayama (1985) theseprocesses are well-developed, or are in contact medially,in ocellata and hamiltonii. We confirm the presence ofwell-developed processes in these taxa and add petersi,
N. platynota, and all sampled tortoises to the list. We
recommend that this character not be subdivided into
additional character states, because the descending
processes of the frontals grow larger through ontogeny.

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Asiatic Herpetological Research

Polarity: A medial inflection is absent in all outgroups and the vast majority of the ingroup. We consider its presence to be derived.
(3) Frontal contribution to the orbital rim; 0 = present,
no prefrontal/postorbital contact on dorsal surface, Fig.
5; 1 = absent, frontal excluded from orbital rim by prefrontal/postorbital contact, Fig. 6 (modified from
Crumly, 1982, 17; Hirayama, 1985, 3; Shaffer et al.,1997, 97; Yasukawa et al., 2001, 1).
Three states for this character were scored by
Hirayama (1985) and Yasukawa et al. (2001): frontal
contribution always or usually present, frontal sometimes excluded from orbital rim, and frontal always
excluded from orbital rim. Our sample size for manytaxa does not permit a reliable assessment of intraspecific variation in this character, and thus our initial scores
differed for some taxa from those of Hirayama (1985).
We add petersi and N. platynota to the list of taxa inwhich the frontal appears always to be excluded. For
those taxa that sometimes exclude the frontals, we confirm this polymorphic condition in crassicollis, and add
agassizii and annulata. Our sample was too small to
confirm the reported polymorphic condition inamboinensis, and pulcherrima by Hirayama (1985) andYasukawa et al. (2001), but we scored these taxa as
polymorphic based on their observations. We also followed Crumly (1982) by coding pardalis as polymorphic, even though we were not able to observe this in our
sample. Our coding differs from that of Shaffer et al.
(1997) for Heosemys and reevesii. In their analysis, theyused spinosa (in which the frontal contributes to theorbital rim), but we used grandis (in which it does not).
In both specimens of reevesii available to us, the frontal
clearly does not participate in the orbital margin. Given
the contrary statement by Shaffer et al. (1997), reevesii
may be polymorphic for this character.
Polarity: The frontal participates in the orbital rim
in pleurodires and spinifera, it is excluded in odoratus
and caretta, and it is polymorphic in serpentina. No pattern is apparent within the ingroup. Given that thefrontal clearly contributes to the orbital rim in
Mongolemys, we consider its absence to be derived.
(4) Contact between jugal and pterygoid; 0 = present,
medial process of jugal well-developed and touching thepterygoid, Fig. 7; 1 = absent, medial process reduced,
Fig. 8 (modified from Hirayama, 1985, 11, 12; McCord
et al., 1995, 3; Burke et al., 1996, 23; Yasukawa et al.,2001, 4, 5).
The jugal of most testudinoid turtles is expanded at
its ventral end to form a medial process that contacts thepterygoid medially (McDowell, 1964). Presence orabsence of the medial process, and presence or absence

Vol. 10, p. 57

of a medial contact with the pterygoid were treated as
two characters by Hirayama (1985) and Yasukawa et al.
(2001). The scoring for the two characters appears to be
redundant and we followed the recommendation ofGaffney and Meylan (1988) by combining them.
We confirm the loss of a medial contact between thejugal and the pterygoid in galbinifrons, flavomarginata,and mouhotii (Hirayama, 1985; Yasukawa et al., 2001),
but we found this condition to be polymorphic in spengleri (also reported by McCord et al., 1995), and in trijuga. Our observations are concordant with those of Burkeet al. (1996).
Polarity: A contact between the jugal and pterygoid
in present in spinifera, odoratus, and serpentina, but isabsent (although the two bones closely approach one
another) in caretta. We conclude that the contact
between the two bones is the primitive condition for testudinoids and that their separation is derived, a conclusion also reached by Hirayama (1985). Our polarity
determination is opposite that used by McCord et al.
(1995), who mistakenly claimed to have derived theirpolarity assessment from Hirayama (1985).
(5) Contact between jugal and palatine; 0 = absent, Fig.
9; 1 = present, Fig. 10 (Gaffney and Meylan, 1988,
F5.4).
The presence of a contact between the medial
process of the jugal and the palatine was used previously in support of a monophyletic Deirochelyinae(Gaffney and Meylan, 1988). We confirm the formerly
observed distribution of this character within ‘emydids’
with the exception of reticularia, which does not exhibit a contact. A contact is present in numerous‘batagurids,’ such as borneoensis, reevesii, and hamiltonii, but was absent in all examined members of‘Testudinidae.’
Polarity: A contact between the jugal and the palatine is present in caretta, odoratus, and pleurodires, but
is absent in spinifera and serpentina. A contact is absent
in Mongolemys. We consequently consider its presence
to be derived.
(6) Contact of the epipterygoid with the jugal; 0 = clearly absent, Fig. 11; 1 = present, or almost present,
epipterygoid forms a long lateral process that approaches the jugal, Fig. 12 (Gaffney and Meylan, 1988, F8.1;
Shaffer et al., 1997, 106).
According to Gaffney and Meylan (1988) the
epipterygoid and the medial process of the jugal
approach one another or are in contact in reticularia andthe various species they included in Pseudemys andTrachemys. They also noted a contact between these twoelements in species they classified in Graptemys, but thecondition in those taxa was interpreted to be a result of

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the medial expansion of the jugal and not a lateralexpansion of the epipterygoid, and consequently was
regarded as non-homologous (Gaffney and Meylan,
1988). We confirm the contact or near contact of these
two elements in reticularia, decorata, scripta, alabamensis, P. nelsoni, rubriventris, texana, flavimaculata,
geographica, kohnii, nigrinoda, G. oculifera, ouachitensis, and versa, and also report it in picta and terrapin.
Contact was clearly absent in the specimens of barbouri,
ernsti, and gibbonsi we examined. We made no assessments of homology, but accept any contact between
these two elements as the derived state (as was done by
Shaffer et al., 1997). Among ‘batagurids,’ we also founda close approach in reevesii.
Polarity: A contact, or near contact, between the
epipterygoid and the jugal is absent in all outgroups andthe vast majority of the ingroup. We consider its presence to be derived.
(7) Contact of the inferior process of the parietal withthe medial process of the jugal; 0 = absent, Fig. 13; 1 =present, Fig. 14 (Hirayama, 1985, 13).
Our coding differs significantly from that of
Hirayama (1985). In reevesii, N. platynota, and bealei
we found no contact between the parietal and jugal,
although these were the only three taxa in which
Hirayama (1985: table 2) scored it to be present.
However, we found a pronounced contact between these
two elements in subtrijuga, a species scored by
Hirayama (1985: table 2) as lacking such a contact, but
shown on his tree (Hirayama, 1985: fig. 2) as a unique‘batagurid’ feature convergent with some ‘emydids.’
Among ‘emydids,’ a well-developed contact occurs inbarbouri and other broad-headed species currently classified in Graptemys.
Polarity: There is no contact between the inferior
process of the parietal and the medial process of thejugal in all outgroups and the vast majority of the
ingroup. A contact is considered to be the derived condition.
(8) Contact of the inferior process of the parietal withthe maxilla; 0 = absent, Fig. 13; 1 = present, Fig. 14(Hirayama, 1985, 14).
Our coding differs from that of Hirayama (1985).
According to his character matrix (table 2) a contact
should be present between the inferior process of theparietal and the maxilla in reevesii and mouhotii, but his
cladogram (fig. 2) indicated that the presence of a contact should be regarded as a uniquely derived autapomorphy of subtrijuga. We found no contact in reevesii or
mouhotii. Of the testudinoid species we examined, subtrijuga is the only one that shows this feature.
Polarity: There is no contact between the inferior

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process of the parietal and the maxilla in all outgroups.
Its presence is considered to be derived.
(9) Extent of quadratojugal; 0 = quadratojugal welldeveloped, firmly attached to jugal, Fig. 15; 1 = quadratojugal present, contact lost with jugal, Fig. 16; 2 =quadratojugal so heavily reduced that it appears to beabsent in many skeletal specimens, Figs. 17, 18 (modified from Hirayama, 1985, 16; Shaffer et al., 1997, 47;
Burke et al., 1996, 21; McCord et al., 1995, 6; Yasukawa
et al., 2001, 7, 8).
Variation in the structure of the temporal region of
turtles was discussed in detail by Zdansky (1924) andcomments specific to testudinoids were provided byZangerl (1948) and McDowell (1964). We originallyscored the reduction of the quadratojugal as three different characters: loss of contact with the jugal, loss of contact with the squamosal, and the apparent loss of thequadratojugal. All five logically possible combinations
were observed, but in most testudinoid turtles the temporal arch is so slender that the contact between thequadratojugal and squamosal is commonly reduced to asliver that would have to be scored as ‘just barely present’ or ‘just barely absent.’ We therefore abandoned ourefforts to evaluate the contact between the quadratojugal
and squamosal. Our observations generally agree withthose of McDowell (1964), Hirayama (1985), Burke etal., (1996), McCord et al. (1995), and Yasukawa et al.
(2001).
We purposefully avoid addressing the apparent lackof a quadratojugal in many species as an absence,
because previous work by Zdansky (1924) showed thatthe quadratojugal of some ‘batagurids’ is so poorly ossified and connected to the surrounding elements that it
tends to be lost in skeletal specimens (Figs. 17, 18). An
example of this problem can be found among the many
conflicting statements made regarding the presence of
this element in N. platynota (e.g., Smith, 1931; Bourret,
1941; McDowell, 1964; Ernst and Barbour, 1989).
Polarity: The quadratojugal is present and firmly
attached to the jugal in all outgroups, with the exceptionof chelids. Its reduction is considered to be derived.
(10) Contribution of jugal to the rim of upper temporalemargination (Hirayama, 1985, 15); 0 = absent, Figs.
19, 20; 1 = present, Fig. 21.
Participation of the jugal in the rim of the upper
temporal emargination was reported previously inhamiltonii and ocellata (Hirayama, 1985). We confirm
its presence in both species, but in one of the hamiltonii
specimens we examined (MCZ 120333) the jugal forms
a significant part of the rim only on one side of the skull;on the other side, which appears abnormal and likely
represents a teratology, it does not. In all other species

2004

Asiatic Herpetological Researchavailable to us, the jugal does not participate in the rim.
In subtrijuga, the jugal is excluded from the upper temporal emargination by narrow extensions of the postorbital and quadratojugal (Fig. 20).
Polarity: The jugal participates in the upper temporal rim of spinifera, but it is excluded in odoratus, caretta, serpentina, Mongolemys, and most pleurodires. We
consider the participation of the jugal in the rim of theupper temporal emargination to be the derived condition
within testudinoids.
(11) Contact between the quadratojugal and the articular facet of the quadrate; 0 = absent, Fig. 22; 1 = present, quadratojugal sends a process ventrally along therim of the cavum tympani and touches the lateral edgeof the articular facet, Fig. 23 (modified from Hirayama,
1985, 17).
The original character definition (Hirayama, 1985,
character 17) is inappropriate, because the jugal does notcontact the articular surface of the quadrate in any turtle
except for madagascariensis and dumerilianus (Gaffney
and Meylan, 1988). However, because Hirayama indicated in his tree that the only ‘batagurid’ taxon to exhibit this character is subtrijuga, we assume that he wasreferring to a contact between the quadratojugal and thearticular facet of the quadrate, a characteristic of subtrijuga only among testudinoids. A contact between thetwo elements was reported previously for reevesii in thecharacter matrix published by Hirayama (1985), but weconclude that this must be a publishing error, because it
stands in conflict with his tree. In the specimens ofreevesii available to us, there is no contact. It is possiblethat the scoring for subtrijuga and reevesii were flipped,
at least in part, in the Hirayama (1985) matrix (in which
the taxa were listed next to one another).
Polarity: A contact between the quadratojugal andthe articular surface of the quadrate is absent inspinifera, but present in odoratus, caretta, and serpentina, and consequently could be considered primitive.
However, based on ingroup commonality and theabsence of a contact in Mongolemys, we consider a contact to be derived for Testudinoidea.
(12) Contact between quadratojugal and maxilla; 0 =absent, Fig. 22; 1 = present, Figs. 23, 24 (Hirayama,
1985, 18).
According to Hirayama (1985), among ‘batagurids’
a contact between the quadratojugal and maxilla is only
present in subtrijuga and reevesii. We did not find acontact in our specimens of reevesii, but confirm its
presence in subtrijuga.
Gaffney and Meylan (1988) listed a contact
between the quadratojugal and maxilla as a synapomorphy for Platysternina {megacephalum + ‘Chelydropsis’}Vol. 10, p. 59

and as an independently evolved synapomorphy for
Kinosternidae, whereas Shaffer et al. (1997) noted acontact to be present in Kinosternidae, C. insculpta, andmegacephalum. Our observations confirm the presenceof a contact in all of these extant groups. Among testudinoids, subtrijuga is unique in having an extensive contact in lateral view (Fig. 23). In several ‘emydids,’ a contact is present on the inside of the temporal arch (barbouri, and nigrinoda; polymorphic in geographica, G.
oculifera, and texana; Fig. 24). For now, we scored all
taxa as present, regardless of whether the contact is visible in lateral view, medial view, or both. In several other‘emydid’ taxa, the bones closely approach one another,but do not actually meet, on the inside of the temporalarch (alabamensis, ernsti, flavimaculata, gibbonsi,
kohnii, P. nelsoni).
Polarity: A contact is present between the quadratojugal and maxilla in odoratus, but absent in spinifera,
caretta, serpentina, and Mongolemys. We consider thepresence of a contact to be derived for Testudinoids.
(13) Medial contact of the maxillae along the anterior
margin of the jaw; 0 = absent, Figs. 25, 26; 1 = present,
Fig. 27 (modified from Hirayama, 1985, 20; McCord etal., 1995, 2; Yasukawa et al., 2001, 10).
In most testudinoids, the anteromedial ends of themaxillae are separated medially by the premaxillae
along the anterior margin of the jaw (Fig. 25). Hirayama(1985) noted that the maxillae have a medial contact insome ‘batagurids,’ which was confirmed by McCord etal. (1995) and Yasukawa et al. (2001) for spengleri andseveral other species that they included in the genus
Geoemyda. We found a broad medial contact of the maxillae in spengleri and annulata (Fig. 27). In somespecies, the maxillae approach one another along theventral rim of the nasal opening (e.g., amboinensis,
mouhotii, pulcherrima, crassicollis), but a well-developed contact is never present (Fig. 26).
Polarity: The maxillae do not meet medially along
the anterior margin of the jaw in odoratus, caretta, serpentina, and Mongolemys. A medial contact is present inspinifera but only along the ventral border of the external nares. We consider a medial contact along the anterior margin of the jaw to be the derived condition within Testudinoidea.
(14) Size of the foramen orbito-nasale; 0 = small, lessthan 1/6 of orbit length, Figs. 28, 29; 1 = large, morethan 1/6 of orbit length, Fig. 30 (modified from
Hirayama, 1985, 33; Gaffney and Meylan, 1988, F9.3,
F10.2, G10.3, H11.1, H16.3; Crumly, 1982, 25; Crumly,
1994, 12).
We were cautious when first approaching this character due to the inconsistent usage and definition of

Vol. 10, p. 60

Asiatic Herpetological Research‘small’ and ‘large’ by various authors. However, after
assessing the size of this foramen based on its size relative to the length of the orbit, we were surprised to seethat we were able to reproduce Hirayama’s (1985) scoring for the ‘batagurids’ without too many difficulties. In
contrast, our initial observations of tortoises were instark contrast to those of Crumly (1982, 1985, 1994) and
Gaffney and Meylan (1988). This may be due to the thinnature of the palatine of many tortoises, and the relative
ease with which that part of the palate can be damaged
during skeletal preparation and handling. Furthermore,
the foramen becomes progressively more closed withincreased ontogenetic age (Crumly, 1982). We encountered similar problems in attempting to score reticularia
and blandingii. Because we deem this character to bepotentially useful for helping to resolve phylogeny within ‘batagurids’ and ‘emydids,’ we decided to score all
testudinoids with delicate palatines (i.e., all tortoises,
reticularia, and blandingii) as ‘unknown.’ We acknowledge that our redefinition of the character is still subjective and somewhat problematic, but using this definition
we were able to unambiguously score all the ingrouptaxa we examined.
Polarity: The foramen orbito-nasale is large in serpentina, odoratus, and spinifera, but small in caretta.
We consider presence of a large foramen orbito-nasale tobe the derived condition within testudinoids, because theforamen is small in Mongolemys.
(15) Contact between maxilla and vomer; 0 = present,
Fig. 31; 1 = absent, vomer separated from the maxillaby the premaxilla, Fig. 32 (Hirayama, 1985, 31; Crumly,
1982, 21; Yasukawa et al., 2001, 14).
We generally agree with previous scorings for thischaracter (Hirayama, 1985, Crumly, 1982, Yasukawa etal., 2001). We confirm the absence of a contact inamboinensis and pulcherrima, but our scoring differs
slightly for those taxa that Hirayama (1985) coded as‘intermediate apomorphic,’ a character state that weinterpret as polymorphism. Of those taxa that Hirayama(1985) and Yasukawa et al. (2001) scored as intermediate (flavomarginata, caspica, annulata), our sample size
is too small to confirm whether both character states arepresent. We consequently follow these authors by scoring those taxa as polymorphic.
Polarity: A contact between the maxilla and vomer
is present in all outgroups. The loss of this contact isderived for testudinoids.
(16) Size of the foramen palatinum posterius; 0 = large,
Fig. 33; 1 = small, Fig. 34 (modified from Hirayama,
1985, 22; Gaffney and Meylan, 1988, F2.2, F6.1;
McCord et al., 1995, 4; Yasukawa et al., 2001, 12).
Our characters 16 and 17 were published originally

2004

by Hirayama (1985) as one character that combined twomorphological features: the size of the foramen palatinum posterius (f.p.p.) and participation of the pterygoid
in the margin of the f.p.p. Although four possible combinations of these features are logically possible, only
two were originally included (participation present,
f.p.p. large; participation absent, f.p.p. small). Gaffney
and Meylan (1988) also used this character within ‘emydids,’ but their character applied only to the exclusion of
the pterygoid from the f.p.p. We decided to subdivide
Hirayama’s (1985) character into one character that
describes the size of the f.p.p. and a second that addresses the position of the pterygoid relative to the f.p.p.
We found no difficulty in identifying the f.p.p. as‘large’ or ‘small,’ (Figs. 33 and 34, respectively), and no
ambiguous condition was encountered. Because ourcharacter definition only includes two character states,
our scorings do not reflect those of other workers with amore limited target group (e.g., McCord et al., 1995). In
juveniles the f.p.p. tends to be larger, but during later
ontogenetic stages the f.p.p. is slowly reduced in size.
Polarity: The f.p.p. is small in odoratus, spinifera,
and most pleurodires; it is absent in caretta, but is large
in serpentina and Mongolemys. We consider a small
f.p.p. to be the derived state.
(17) Position of the pterygoid relative to foramen palatinum posterius (f.p.p.); 0 = pterygoid situated posterior
to the f.p.p., Fig. 33; 1 = pterygoid situated posterior to
the f.p.p., but sends a process anterior and lateral to thef.p.p., Fig. 34.
Our survey of testudinoids indicated that reliable
assessment of participation of the pterygoid in the f.p.p.
may be difficult because many species show an ontogenetic change in configuration of this part of the palate. In
juveniles, the f.p.p. typically includes the pterygoid in its
posterior margin. During later ontogenetic stages thepterygoid is excluded. In spite of this, it appears that therelative position of the pterygoid tends to stay constantduring ontogeny.
Polarity: The anterior end of the pterygoid is situated posterior to the f.p.p. in odoratus, serpentina, mostpleurodires, and Mongolemys, but is situated lateral to
the f.p.p. in spinifera. We consider a posterior position to
be primitive for Testudinoidea.
(18) Epipterygoid participation in the trigeminal foramen; 0 = absent, Fig. 35; 1 = present, epipterygoid
clearly separates the parietal and pterygoid in lateralview, Fig. 36.
The anteroventral rim of the trigeminal foramen of
most testudinoid turtles is formed by the parietal andpterygoid. The epipterygoid commonly comes close to
the foramen, but does not form part of it. In spengleri

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Asiatic Herpetological Research

and mouhotii, the epipterygoid consistently participatesin the margin of the trigeminal foramen thus separating
the parietal and pterygoid, at least in lateral view.
Polarity: The epipterygoid forms part of theanteroventral rim of the trigeminal foramen in the
majority of outgroups, with the exception of pleurodiresthat lack a definitive ossified epipterygoid (Gaffney,1979). The condition is unclear for Mongolemys.
However, within the ingroup we found this character
only in the seemingly rather specialized turtles spengleri
and mouhotii. We consequently consider its presence tobe secondarily derived.
(19) Vomerine foramen; 0 = absent, Fig. 37; 1 = present, Fig. 38 (Gaffney and Meylan, 1988, H4.1; Crumly,
1994, 15).
The vomerine foramen (= anteromedial vomerine
aperture of Crumly, 1982 and 1994 [in part]) is a small
opening that pierces the vomer along the midline justposterior to the foramen praepalatinum (Bramble, 1971).
The presence of a vomerine foramen was noted in agassizii and berlandieri by Bramble (1971), and was usedby Gaffney and Meylan (1988) to unite various species
currently placed in Gopherus as a clade. Its irregularpresence in elegans, elongata, chilensis, and radiata
was reported by Crumly (1982). Specimens in our sample enable us to confirm its presence in agassizii,
berlandieri, and chilensis.
Polarity: The vomerine foramen occurs in only afew ‘testudinids’ and it is absent in all outgroups. Itspresence is considered derived for Testudinoidea.
(20) Development of the foramen praepalatinum as acanal (canalis praepalatinum) that is concealed by abony secondary palate in ventral view; 0 = absent, Fig.
39; 1 = present, Fig. 40 (modified from Hirayama, 1985,
24).
In most testudinoids, the foramen praepalatinum isa small opening that connects the nasal cavity with theroof of the oral cavity (Fig. 39). However, in a numberof taxa with extensively developed secondary palates,
the anterior nasal artery passes through an elongatedcanal that is concealed in ventral view by the bony secondary palate (e.g., baska, tentoria, petersi; Fig. 40). We
refer to this structure as the canalis praepalatinum. Our
scorings are fully consistent with those of Hirayama(1985).
Polarity: The foramen praepalatinum is absent inspinifera (Gaffney, 1979) and caretta (Nick, 1912), andis developed as a true foramen that is exposed in ventral
view in odoratus, serpentina, and Mongolemys. The
development of a canalis praepalatinum is considered
the derived condition within Testudinoidea.

Vol. 10, p. 61(21) Contact between pterygoid and basioccipital; 0 =present, Fig. 41; 1 = absent, Fig. 42 (modified from
Gaffney and Meylan, 1988, F1.1, F10.3, H18.3; Crumly,
1994; Shaffer et al., 1997, 103).
Two of the most often-cited characters that purportedly help to distinguish the ‘Emydidae’ from the‘Bataguridae’ are the batagurine process and the contact
between the pterygoid and the basioccipital (McDowell,1964). Both traits are commonly combined as one character (e.g., Gaffney and Meylan, 1988) and even seem tohave been confused with one another (e.g., Mlynarski,
1976; Shaffer et al., 1997). The batagurine process is apoorly-defined feature that, in McDowell’s originalusage (1964) appears to consist of a lateral process ofthe basioccipital that floors the recessus scalae tympani.
Many testudinoid species (including non-‘batagurines’)
have a laterally-projecting process of the basioccipital; it
may or may not floor the recessus scalae tympani, but it
is often obscured from view in articulated specimens.
When disarticulated material is examined, a broader distribution of this feature (assuming it is interpreted as wehave done above) across testudinoids is revealed.
Within Testudinoidea, the pterygoid commonly
sends a process posteriorly and contacts the basioccipital just lateral to the basisphenoid. This character
appears to be absent in most ‘emydids,’ but is present interrapin and those species that are currently attributed to
Graptemys (Gaffney and Meylan, 1988). We noticed astrong ontogenetic component to this character within‘emydids.’ The pterygoid is commonly rather short during younger ontogenetic stages, but finally reaches thebasioccipital in later stages. For instance, among our
specimens of terrapin, orbicularis, and texana, thepterygoid does not contact the basioccipital in younger
individuals, but a clear contact is present in adults. We
score such species as polymorphic, but note that ontogenetically influenced polymorphisms are not well
explored in turtles.
Among ‘batagurids,’ we noted a similar pattern.
Contrary to general belief we were not able to observe acontact between the pterygoid and the basioccipital in all
taxa traditionally classified in this group (e.g., trijuga,
pulcherrima, sinensis). As with the ‘emydids,’ there
seems to be an ontogenetic effect in which younger
specimens tend not to have a contact. Where recognized,
we scored these species as polymorphic.
We found much conflicting data regarding the distribution of this character among tortoises (Crumly,
1982, 1985, 1994; Gaffney and Meylan, 1988). Amongthe specimens we examined, we note the complete
absence of a contact only in graeca; polymorphisms
were observed in polyphemus and horsfieldi. Again, an
ontogenetic component is apparent.
Polarity: A contact between the pterygoid andVol. 10, p. 62

Asiatic Herpetological Researchbasioccipital is present in all outgroups. Its absence isconsidered to be derived.
(22) Contact of the pterygoid with the articular facet ofthe quadrate; 0 = absent, Fig. 43; 1 = present, Fig. 44(Hirayama, 1985, 38).
According to the data matrix published by
Hirayama (1985) he only observed this contact inreevesii, however, in his tree (fig. 2) the contact ismapped as an autapomorphy of subtrijuga. We confirm
the presence of a contact between the posterior processof the pterygoid and the articular surface of the quadrate
in subtrijuga. It is the only taxon we examined that displays the derived condition.
Polarity: A contact between the pterygoid and thearticular surface of the quadrate is absent in all outgroups and the vast majority of the ingroup. We consider its presence to be derived.
(23) Closure and depth of the incisura columella auris;
0 = absent, incisura is open, Fig. 44; 1 = present,
incisura closed, Fig. 45 (Crumly, 1985; Gaffney andMeylan, 1988, H1.3).
The incisura columella auris is a notch that isformed by the quadrate and that holds the stapes andeustachian tube (Gaffney, 1972). In a number of turtles,
the incisura closes to fully surround the stapedial shaft(Gaffney and Meylan, 1988). Within testudinoids, theincisura evidently is closed in most tortoises (Crumly,1985). We are able to confirm the presence of such aclosed incisura in all ‘testudinids’ we examined with the
exception of one specimen of kleinmanni (CAS228431), the smallest of the species now classified in
Testudo. In a number of ‘batagurids’ and ‘emydids’ theincisura commonly is very narrow and even appears to
be closed, however, a closer look under the microscope
combined with a probing needle reveals that this apparent closure is produced by dry tissues that remain in thisarea in many articulated skulls. The only ‘batagurid’ forwhich we sometimes found a closed incisura is N.
platynota; in that species the quadrate does not fuse
together forming a solid ring behind the incisura, but thisis also the case for many tortoises (e.g., some belliana,
emys, some homeana, some kleinmanni). In some cases,
the polymorphism we noted (e.g., in belliana and homeana) appears to be a result of ontogenetic age, with older
individuals displaying a greater degree of fusion at theposterior part of the incisura.
Polarity: The polarity of this character is somewhat
ambiguous, because the incisura columella auris is
closed in serpentina and spinifera, but open in caretta
and odoratus. We conclude that its presence is derived
within testudinoids because it is absent in Mongolemys.

2004

Mandible(24) Angular contribution to the sulcus cartilaginis
Meckelii; 0 = present, the angular contributes to the sulcus and is as long or longer than the prearticular, Fig.
47; 1 = absent, the angular is shorter than the prearticular, Fig. 48 (modified from Gaffney and Meylan, 1988,
F1.4).
A broad contact of the angular with Meckel’s cartilage was used by McDowell (1964) to characterize the‘Emydinae’ and later used by Gaffney and Meylan(1988) as a synapomorphy to unite the same grouping.
As it was originally worded, this character is difficult toobserve in its literal sense for most museum specimens,
because the Meckel’s cartilage usually is not present inmodern and fossil skeletal specimens. We suggest thatthe spirit of McDowell’s (1964) character can be evaluated by examining the participation of the angular in thesulcus cartilaginis Meckelii. We confirm that a small tobroad angular contribution is present in all ‘emydids’
with the exception of rubriventris. In most ‘batagurids,’
the angular is a short bone that does not participate in thesulcus and barely spans half the distance the prearticulardoes. However, a small but clear contribution to the sulcus is present in an eclectic group comprised of baska,
dentata, thurjii, punctularia, and some grandis. We were
not able to carefully evaluate potential polymorphisms
in these taxa.
Polarity: An angular contribution to the sulcus cartilaginis Meckelii is present in all cryptodiran outgrouptaxa. We consider is absence to be derived.
(25) Contact between surangular and dentary; 0 =simple contact, Fig. 49; 1 = strongly interdigitated suture,
Fig. 50 (Crumly, 1982, 12; Crumly, 1985; Gaffney andMeylan, 1988, H6.1).
In most testudinoids, the surangular and dentary
meet along the lateral side of the mandible in a simple,
overlapping contact. According to Crumly (1982, 1985),
this contact is stabilized through a finger-like process ofthe surangular that interdigitates with the dentary in all
tortoises except emys, impressa, and those species heclassified in Gopherus. We confirm the absence of thischaracter in agassizii, berlandieri, emys, impressa, andpolyphemus, but also did not observe it in areolatus.
Polarity: An interdigitated contact between thesurangular and dentary is absent in all outgroups and the
majority of the ingroup. We consider its presence to be
derived.
(26) Height of the processus coronoideus; 0 = as high as
dentary, Fig. 51; 1 = rising significantly above the dentary, Fig. 52 (modified from Hirayama, 1985, 45).
The coronoid of most turtles is a small bone that
produces a minor knobby projection that rises only little

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Asiatic Herpetological Research

above the adjacent dentary, if at all. According to
McDowell (1964) and Hirayama (1985) the coronoid islarger and rises moderately above the dentary inborneensis and crassicollis, and in reevesii and subtrijuga the coronoid is very large and produces a robust
process that sits high above the dentary. We confirm
these observations, however, we were also able toobserve moderately developed coronoid processes inkachuga and tentoria. Among ‘emydids,’ we also
observed moderately developed coronoids in barbouri,
ernsti, flavimaculata, geographica, gibbonsi, kohnii,
and terrapin. Unlike Hirayama (1985), we only utilize
one derived character state, because it is difficult toobjectively measure and discretize the relative height ofthe coronoid among turtles.
Polarity: The coronoids of caretta, serpentina, andodoratus are small and do not rise above the dentary, but
the coronoid of spinifera is well developed and forms amoderate process. The lower jaw is not described for
Mongolemys. We consider well-developed coronoids tobe derived within testudinoids.
(27) Foramen dentofaciale majus; 0 = small, Fig. 53; 1= large and situated within a large lateral fossa, Fig. 54(Hirayama, 1985, 47).
The foramen dentofaciale majus of most testudinoids is a small opening that is situated on the lateralside of the mandible, just ventral and slightly anterior tothe coronoid. The foramen dentofaciale majus is greatly
enlarged in thurjii and ocellata and is situated at theanterior end of an expanded lateral fossa (Hirayama.
1985). We confirm the enlargement in those taxa, andfurther note its presence in petersi.
Polarity: The foramen dentofaciale majus is small inall outgroups and the vast majority of the ingroup. Itspresence is considered to be derived.
Triturating Surfaces(28) Participation of palatine in the triturating surfaceof the upper jaw; 0 = absent, Fig. 55; 1 = present, Figs.
56, 57 (Hirayama, 1985, 26; Gaffney and Meylan, 1988,
F2.1).
In some testudinoids, the palatine has a ventrolateral maxillary process that participates in the triturating
surface of the upper jaw. The degree of participation
varies among taxa, and within some species. A clear andextensive participation is present in barbouri, ernsti,
geographica, gibbonsi, petersi, rubriventris, scripta,
terrapin, texana, and versa. It is weakly developed inhamiltonii, ocellata, G. oculifera, ouachitensis, reevesii,
subtrijuga, and some individuals of baska.
Participation was used by Gaffney and Meylan(1988) to unite Terrapene spp., blandingii, guttata, G.
insculpta, marmorata, muhlenbergii, and orbicularis asVol. 10, p. 63

a clade within the ‘Emydidae.’ In our observations, however, this participation also is absent among other ‘emydids’ such as flavimaculata, kohnii, nigrinoda, picta, andreticularia (we were not able to evaluate adequately thepotential for polymorphism in these taxa). In addition, it
appears that the absence of a palatine participation represents the plesiomorphic state for testudinoids, thus
eliminating its value for diagnosing monophyletic
groups.
Polarity: The palatine does not participate in the triturating surface of spinifera, but does so in odoratus,
caretta, and serpentina. The palatine does not participate
in the triturating surface of Mongolemys and only occurs
in Testudinoids with highly derived secondary palates.
We consequently consider the participation of the palatine in the triturating surface to be derived within testudinoids.
(29) Participation of the vomer in the triturating surfaceof the upper jaw; 0 = absent, Figs. 56, 57, 58; 1 = present, Fig. 59 (Hirayama, 1985, 25).
The triturating surface is the grinding surface of thejaw. In most turtles, it is formed on the upper jaw predominantly by the maxilla and premaxilla. However, inturtles with extensive secondary palates the vomer may
also participate. In texana, the vomer may have a ventral
projection that barely separates the maxillae in the midline, but it does not participate in the triturating surface
proper because it sits in a dorsal concavity within thepalate (Fig. 56). Our scorings differ from those of
Hirayama (1985) for mouhotii (which Hirayama scoredas “intermediate apomorphic” and we score as absent
because it does not have a secondary palate) and subtrijuga (in our specimens the vomer does not descend to
the palatal surface, but this species may be polymorphic). This character is polymorphic in barbouri.
Polarity: Because the vomer does not participate in
the triturating surface of odoratus, spinifera, caretta,
serpentina, and Mongolemys, its participation is considered to be the derived condition for Testudinoidea.
(30) Presence and number of lingual ridges of the triturating surfaces of the upper and lower jaws; 0 = no lingual ridges present, Fig. 60; 1 = one lingual ridge present, Figs. 61-62; 2 = two lingual ridges present (modified from Hirayama, 1985, 29, 44; Gaffney and Meylan,
1988, F7.2, F9.1)
Most turtles lack lingual ridges on their triturating
surfaces (Fig. 60), but one or two such ridges are developed in a number of testudinoids (Figs. 61-62). Theseridges run parallel to the labial surface of the maxillaand dentary, and typically do not meet their counterpart
on the midline. They are not necessarily a continuous
structure (Fig. 61), and may be divided into several com-Vol. 10, p. 64

Asiatic Herpetological Researchponents. In some cases, an extensive ridge-like structure
can create the appearance of an additional ridge at theextreme lingual margin of the maxilla bordering theinternal nares; however, we consider these to be thethickened rim of the internal nares rather than an additional ridge. Among ‘batagurids’ and ‘emydids,’ one lingual ridge is present in alabamensis, borneoensis, decorata, kachuga, P. nelsoni, ocellata, petersi, rubriventris,
scripta, sinensis, tentoria, texana, and thurjii. Two lingual ridges are developed in baska. We found lingual
ridges in all tortoises we examined except erosa, belliana, and homeana.
Hirayama (1985) originally scored this character as
two separate characters, one for the mandible and one
for the maxilla. In our observations, the triturating surface of the lower jaw closely mimics that of the upper
jaw, creating an occlusal surface that closely reproduces
the function of cusps in mammalian cheek teeth. Bothcharacters were scored in parallel in Hirayama’s matrix,
and we see no reason to consider them independent.
Polarity: All outgroups lack lingual ridges on thetriturating surfaces. We consequently consider theirpresence to be derived.
(31) Well-developed serrations on labial or lingual
ridges of the triturating surfaces of the upper and lower
jaws; 0 = absent, Fig. 60; 1 = present, Fig. 61 (modified
from Hirayama, 1985, 21, 27, 41, 43, 46; Gaffney andMeylan, 1988, F9.2; Yasukawa et al., 2001, 11).
Well-developed serrations on the lingual and labial
ridges of the upper and lower jaws are developed in several ‘batagurids’ and ‘emydids’. A number of tortoises
and other ‘batagurids’ (e.g., carbonaria, pardalis, sulcata, annulata, and areolata) exhibit serrations on theirramphothecae, but unlike the bony, tooth-like serrationsof borneoensis, thurjii, petersi, or texana, these crenulations are weakly developed, leaving very little or no
trace of serrations on the underlying bone. In comparison to those taxa with well-developed serrations, it very
difficult to establish a consistent scoring system for taxa
with fine crenulations, because many specimens will not
exhibit any serrations, probably due to wear of the ramphothecae. Unlike Hirayama (1985) and Yasukawa et al.
(2001) we scored all taxa with such weak serrations as
absent.
Unfortunately, even in those taxa with well-developed serrations, the serrations are not always evenly
developed on all ridges. We consequently combined allof Hirayama’s (1985) characters relating to serrations
into one character. Because serrations commonly occur
on all available ridges, this treatment will also help toavoid unconsciously weighting the presence of serrations with up to five characters. In those taxa that havethem, the ridges themselves often have very different

2004

morphologies; this character needs to be criticallyreevaluated with adequate sample sizes for the relevant
taxa.
Polarity: All of our outgroup taxa and the majorityof the ingroup taxa lack strong serrations. We interpret
their presence to be derived.
(32) Median ridge or sulcus of the triturating surface of
the upper jaw; 0 = both structures absent, Fig. 60; 1 =median ridge present, Fig. 62; 2 = median sulcus present, Fig. 63 (modified from Hirayama, 1985, 30;
Crumly, 1985, 1994, 4; Gaffney and Meylan, 1988,
H3.1).
In a number of testudinoid turtles with partiallydeveloped secondary palates and lingual ridges, additional structures are formed along the midline of theupper jaw that typically correspond to reciprocal structures of the lower jaw. The upper jaw of petersi is characterized by a narrow sulcus (Fig. 63) and the mandible
exhibits a low median ridge. On the other hand, inbaska, borneoensis, thurjii, kachuga, agassizii,
berlandieri, and polyphemus, a ridge runs along the midline (Fig. 62), which typically corresponds to a sulcus in
the lower jaw. An incipient ridge also was reported inemys (Crumly, 1994), but we were not able to confirm
this on the specimen available to us.
Polarity: A median ridge is absent in all outgroups
and the vast majority of the ingroup. Its presence is considered to be derived.
(33) Posterior extension of the lower triturating surface
behind the symphysis of the dentary; 0 = absent, Fig.
64; 1 = present, Fig. 65 (Hirayama, 1985, 42; Gaffney
and Meylan, 1988, G5.2).
In several ‘batagurids,’ the triturating surface of thedentary forms a shelf along the midline that extends so
far posteriorly that the symphysis cannot be seen whenthe mandible is observed in dorsal view (McDowell,1964). Our scorings fully agree with those of Hirayama(1985) for the ‘Bataguridae,’ but we disagree with
Gaffney and Meylan (1988) who asserted that this character also occurs in some ‘Emydidae.’ Admittedly, several species currently placed in the genera Graptemys,
Pseudemys, and Trachemys have greatly expanded triturating surfaces of the dentary, but in all of the specimens
available to us, the symphysis is always visible in dorsal
view.
Polarity: An extended triturating surface of the dentary does not occur in any outgroup taxon. We consider
its presence to be derived.
Carapace(34) Carapace strongly tricarinate in adult; 0 = absent,
Figs. 66, 67; 1 = present, Fig. 68 (modified from

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Asiatic Herpetological Research

Hirayama, 1985, F; McCord et al., 1995, 10; Yasukawa
et al., 2001, 24).
Three distinct carapacial ridges are present in theadults of reevesii, hamiltonii, spengleri, subtrijuga, trijuga, and mouhotii. We cannot replicate Hirayama’s(1985) placement of this character as an autapomorphy
in hamiltonii. In our observations, the carinae in hamiltonii are not better developed than in some other taxa. In
fact, they are more weakly developed than those inmouhotii (Fig. 68). Because keels are present in theyoung and subadults of such taxa as crassicollis (Fig.
67), mutica, and sinensis, but disappear with age, andbecause we were not able to observe the juveniles ofmost species, we restricted this character to thosespecies that exhibit well-developed tricarinae as adults.
Three keels were reported to be present in the adults ofdentata (McCord et al., 1995), but we cannot confirm
this (tricarinae are not present on our younger specimens).
Polarity: Tricarinae are absent in our outgroupspecies (caretta, odoratus, serpentina, spinifera, gibba,
siebenrocki, subglobosa, and subrufa), but do appear
occasionally in some of their close relatives, such as
scorpioides, temminckii, and fimbriatus. We consider thepresence of tricarinae to be derived within
Testudinoidea.
(35) Significant serration of the posterior peripherals; 0= absent, Fig. 66; 1 = present, Fig. 68 (modified from
Hirayama, 1985, D; McCord et al., 1995, 11; Yasukawa
et al., 2001, 23).
We generally agree with previous observations
reported for this character (Hirayama, 1985). However,
because the carapace rim is at least slightly serrated inalmost all turtles, we rephrase the character definition to
include only significantly serrated posterior peripherals
as found, for example, in crassicollis, dentata, grandis,mouhotii, N. platynota, and spengleri. Among the ‘emydids’ and ‘testudinids,’ the peripherals of barbouri,
erosa, flavimaculata, geographica, homeana, kohnii,
nigrinoda, oculifer, G. oculifera, pseudogeographica,and versa also are serrated. It is important to note that
our scores are based on the peripheral bones; the amountof carapacial serration greatly depends on the presenceor absence of the marginal scutes in the specimens used,because the scutes greatly accentuate the amount of serration, if present. We find no conflict with the codings of
McCord et al. (1995) and Yasukawa et al. (2001).
Polarity: Serrated posterior peripherals are present
in caretta, but absent in odoratus, serpentina, and mostpleurodires. However, due to ingroup commonality andits absence in taxa placed within “Lindholmemydidae”
we conclude that its presence is derived.

Vol. 10, p. 65(36) Carapace of adult tectiform in cross-section with astrong posterior projection on the third vertebral scute;
0 = absent, Fig. 69; 1 = present, Fig. 70 (Hirayama,
1985, N).
According to Hirayama (1985), this character only
occurs in tecta and tentoria. For our sample, we wereable to confirm its presence in tecta and tentoria andalso observed it in barbouri (barbouri and other species
now classified in Graptemys may be sexually dimorphic
for this character). The descriptive term ‘tectiform’ issomewhat problematic, because any turtle shell can beconsidered ‘roofed’ (Fig. 69). We regard a carapace astectiform if its sides are more-or-less flat and meet along
the midline at a rather sharp angle (Fig. 70). Many testudinoids, and notably those ‘emydids’ currently classified within Graptemys, have a somewhat tectiform carapace as juveniles, but that morphology typically is lost in
the adults.
Polarity: Because all outgroups and the majority of
the ingroup do not have a tectiform carapace, we consider its presence to be derived.
(37) Shape and orientation of the second neural; 0 =second neural hexagonal, short sides positioned anteriorly, Fig. 71; 1 = second neural hexagonal, short sides
positioned posteriorly, Fig. 72; 2 = second neuraloctagonal, Fig. 73 (modified from Hirayama, 1985, G;
Yasukawa et al., 2001, 25).
(38) Shape and orientation of the third neural; 0 = third
neural hexagonal, short sides positioned anteriorly, Fig.
71; 1 = third neural hexagonal, short sides positioned
posteriorly, Fig. 72; 2 = third neural square, Fig. 74; 3= third neural octagonal, Fig. 75 (modified from
Hirayama, 1985, G; Yasukawa et al., 2001, 25).
Originally, Hirayama (1985) only discussed the orientation of the neurals in general, which is fully sufficient for his ‘batagurid’ ingroup, because almost all individuals exhibit his two suggested character states.
However, in most tortoises the second and/or third neurals are not hexagonal, but rather are square or octagonal, making it impossible to assign them to one of
Hirayama’s (1985) character states. We consequently
split Hirayama’s original character into two characters,
restricted their application to the second and third neural, and added additional character states.
Our observations generally agree with those of
Hirayama (1985) and Yasukawa et al. (2001) for‘batagurids,’ with the exception of annandalei in which
we found the short side of the second and third neurals
to be positioned anteriorly, and not posteriorly as wasindicated by Hirayama (1985).
Polarity: The short side of the second and third neural bones faces posteriorly in odoratus and spinifera but

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Asiatic Herpetological Research

2004

anteriorly in caretta. The shape of the second and third
neurals is extremely variable in serpentina. However, in
Lindholmemys and Mongolemys the short side of thesecond and third neurals is positioned anteriorly. We
consider that condition to be primitive for
Testudinoidea.

Meylan (1988) that this scute is absent in carbonaria,
chilensis, elegans, nigra, pardalis, and sulcata and additionally code homeana and erosa as polymorphic.
Polarity: The cervical scute is present in all cryptodiran outgroups that have scutes on their carapace. We
consider its absence to be derived.

(39) Medial contact of the seventh and/or eighth costal
bones; 0 = absent, Fig. 76; 1 = present, Fig. 77(Hirayama, 1985, V; Yasukawa et al., 2001, 26).
In some testudinoid turtles, the posterior costal
bones meet along the midline due to the reduction of theposterior neural elements. The original character definition provided by Hirayama (1985) and Yasukawa et al.
(2001) was worded to indicate a contact between theseventh and eighth costal bones among some‘batagurids.’ We are unable to reproduce their results if
the character definition is taken literally. For example, in
all our specimens of amboinensis and galbinifrons, theeighth costals meet on the midline, but the seventh
costals do not. However, if the character definition ismodified to include any contact of the seventh or eighth
costals, our results are concordant with those of
Hirayama (1985) and Yasukawa et al. (2001). In addition
to the Asian box turtles, we report a medial contact ofthe posterior costals in baska, carolina, coahuila, T. nelsoni, T. ornata and rubida.
Polarity: A medial contact of the seventh and/or
eighth costals is absent in serpentina, but present inspinifera, caretta, many kinosternids, and many pleurodires. Although a composite reconstruction of a putative “lindholmemydid” from the Early Cretaceous ofJapan was illustrated with the seventh costals in contact
at the midline (Hirayama et al., 2000, fig. 11), such acontact is absent in other specimens of Lindholmemys
and Mongolemys. Its presence within testudinoids is predominantly in the highly derived box turtles, and we
conclude that its presence is derived for Testudinoidea.

(41) Number of vertebral scutes; 0 = five, Fig. 80; 1 =six or more, Fig. 81 (Hirayama, 1985, P).
We confirm Hirayama’s (1985) observation that
there are at least six vertebral scutes in N. platynota.
Additional scutes occasionally occur in other species,
but are best considered abnormalities; they typicallylack the symmetrical associations with adjacent pleuralscutes seen in N. platynota.
Polarity: All testudinoids, except for N. platynota,
have five vertebral scutes. We consider the presence ofsix scutes to be derived.

(40) Cervical scute; 0 = present, Fig. 78; 1 = absent,
Fig. 79 (modified from Crumly, 1985, 1994, 34; Gaffney
and Meylan, 1988, H5.2, H10.1; Shaffer et al., 1997,
41).
The presence and shape of the cervical scute is usedcommonly to determine phylogenetic relationships
within tortoises. According to Crumly (1985) the cervical scute is very narrow or absent in all tortoises except
agassizii, berlandieri, emys, flavomarginatus, impressa,
and polyphemus. We generally agree with these observations, but when this character is applied to all testudinoids intraspecific variation is so great that the character
becomes essentially useless. We consequently limit ourscoring to the mere presence or absence of the cervicalscute. We confirm the observations of Gaffney and(42) Position of the anterior sulcus of the fourth vertebral scute; 0 = sulcus lies on the fifth neural, Fig. 82; 1= sulcus lies on fourth neural, or on the suture between
the fourth and fifth neural, Fig. 83; 2 = sulcus lies on thesixth neural, or on the suture between the fifth and sixth
neural, Fig. 84 (modified from Hirayama, 1985, L+M).
(43) Position of the posterior sulcus of the fourth vertebral scute; 0 = sulcus lies on the eighth neural, or on thehomologue of the eighth neural, if the seventh is reduced(e.g., in most tortoises), Fig. 85; 1 = sulcus lies on theseventh neural, or on the suture between the seventh andeight neural, Fig. 86; 2= eighth neural absent, sulcus
overlies costals that meet at the midline, Fig. 87 (modified from Hirayama, 1985, L+M).
The size of the fourth vertebral scute was addressedwith two characters by Hirayama (1985) but the total
range of morphological variability in testudinoids is notencompassed by his character definitions. In most testudinoids, the fourth vertebral scute covers the posterior
half of the fifth neural bone, the sixth and seventh neurals, and the anterior half of the eighth neural. A numberof variations are known, and simply counting the number of neural bones covered by this scute results in problems by creating a false perception of homology. For
instance, a fourth scute that partially overlies the fourthand seventh neurals and fully covers the fifth and sixth,
can strictly be said also to cover four neurals, but the elements involved are only partially homologous with thecommon condition. We attempt to resolve these issues
by establishing two new characters that preserve what
we think was Hirayama’s (1985) original intent, but that
permit a more accurate representation of the association
between the fourth vertebral scute and the underlying
neural bones.

2004

Asiatic Herpetological ResearchSome problems that are associated with scoring thischaracter include the prevalence of scute abnormalities
among testudinoids (e.g., Coker, 1905, 1910; Newman,
1906; Zangerl and Johnson, 1957). Specimens exhibiting such abnormalities were scored as unknown. The
notable exception to this is N. platynota, in which asixth, or even a seventh, vertebral scute is always present.
Polarity: Determining the polarity through outgrouprelationship is somewhat difficult, because almost every
outgroup exhibits a different condition, especially forthe posterior sulcus. Based on ingroup commonality, andthe presence of our zero state in both Lindholmemys andMongolemys, we consider the sulci of the fourth vertebral scute to be primitively situated on the fifth andeighth neural bones.
(44) Posterior margin of first vertebral scute significantly narrower than its anterior margin; 0 = absent, Fig.
88; 1 = present, Fig. 89 (modified from Hirayama, 1985,
C).
When originally proposed, this character wasapplied to a posterior constriction of all vertebral scutes(Hirayama, 1985). If strictly applied, this character isabsent in all taxa, because the fifth vertebral scute neveris constricted along its posterior edge relative to theanterior edge. If each scute is viewed by itself, it
becomes apparent that especially the fourth vertebral
scute tends to be narrowed posteriorly, as can beobserved in all species now classified in the genera
Graptemys, Heosemys, Trachemys, and Testudo among
others. According to Hirayama (1985), posterior narrowing is limited to crassicollis and borneensis and unites
them as a synapomorphy. We were able to replicate this
distribution only if the character definition was restricted to the first vertebral scute, in which the posterior margin is significantly narrower than its anterior margin in
those two species only. In making this change, however,this character becomes at least partly redundant withcharacters 45 and 47.
Polarity: Outgroup analysis reveals that posterior
narrowing of the first vertebral scute is present only inodoratus; it is absent in Lindholmemys andMongolemys. We regard the presence of a posterior narrowing of the first vertebral scute to be derived.
(45) Anterior half of the first vertebral scute much narrower than posterior half, especially in adults; 0 =absent, Fig. 90; 1 = present, Fig. 91 (modified from
Hirayama, 1985, R).
We confirm the clear presence of an anteriorly narrowed first vertebral scute in dentata and spinosa asreported by Hirayama (1985), and note that grandis ispolymorphic. Because the anterior sulcus of the first ver-Vol. 10, p. 67

tebral scute commonly is restricted to the nuchal bone inseveral other taxa, but the scute shows no anterior narrowing, we limit the original character definition to theshape of the first vertebral scute only. This peculiar morphology seems to be the result of growth that is limited
to the anterior edge and the posterior half of the lateraledge of the first vertebral scute. As a consequence, thischaracter is not apparent in juveniles, but becomesincreasingly accentuated in adults.
Polarity: Anterior narrowing of the first vertebral
scute is absent in all outgroups and within the large
majority of the ingroup. We consider its presence to be
derived.
(46) Significant contact of the tenth marginal scute withthe fifth vertebral scute; 0 = absent, Fig. 92; 1 = present, Fig. 93 (modified from Hirayama, 1985, K).
Contact of the tenth marginal scute with the fifth
vertebral scute was reported previously only in baska,
smithii, tecta, and tentoria (Hirayama, 1985). We areable to confirm the presence of a very well developed
contact in all but smithii (not seen), and we add spinosato the list of species in which this contact may occur (itis polymorphic for spinosa; contact is present in CAS
228368, but absent in the smaller CAS 228459, so ontogenetic differences may explain the polymorphism). We
also note slight contacts in some specimens of otherspecies (e.g., agassizii, borneoensis, carbonaria, homeana, orbicularis, pardalis, and polyphemus), but by
rewording Hirayama’s (1985) original character toinclude only significant contact, we are able to retain
what we believe was his original intent.
Polarity: Due to the absence of contact in all outgroups in which it is applicable and the predominancewithin the ingroup, we consider its presence to be
derived.
(47) Contact of the second marginal scute with the firstvertebral scute; 0 = absent, Fig. 94; 1 = present, Fig. 95(Hirayama, 1985, O; see also Tinkle, 1962, table 1,‘Seam A’).
According to Hirayama (1985), the first vertebral
scute usually (>90%) contacts the second marginal scutein japonica, leprosa, and caspica. He also noted that thescutes are sometimes in contact in N. platynota andbealei. For our sample, we are able to confirm this contact as a polymorphism for bealei, caspica, japonica,
and leprosa, but the contact is clearly absent in all our
specimens of N. platynota. We also note that these scutes
are sometimes in contact in picta, amboinensis, orbicularis, and terrapin. Together with all of the above, thesetaxa were scored as polymorphic. The only taxa to
exhibit a consistently well-developed contact are reticularia and blandingii.

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Polarity: A clear contact between the second marginal scute and the first vertebral scute does not exist in
caretta or serpentina, but both morphologies occur inkinosternids and pleurodires. The scutes are not in contact in Lindholmemys and Gravemys, but they are in contact in Mongolemys. The polarity for this character isambiguous.
(48) Contact of the sixth marginal scute with the third
pleural scute; 0 = absent, Fig. 96; 1 = present, Fig. 97(modified from Hirayama, 1985, B; see also Tinkle,
1962, table 3, ‘Seam C’).
The contact between the sixth marginal and third
pleural scutes is easily enough rendered as a simple‘presence or absence’ character, but this hides the rangeof possible morphological variation. The degree of contact can range from extensive to a condition where thetwo scutes just barely contact at their corners. Several
taxa exhibit a condition where these scutes either barely
touch or do not touch one another at their corners, butwhenever several specimens were available to us, they
typically turned out to be polymorphic. For this reason
Gaffney and Meylan (1988) called this character ‘dubious.’ We scored such borderline cases as polymorphic,even if not enough specimens were available to corroborate this.
We confirm Hirayama’s (1985) observations
regarding the absence of a contact between these scutes
in baska, ocellata, tecta, tentoria, and thurjii, and we
add petersi and spinosa to that list. Taxa that we score aspolymorphic include annandalei, borneensis, borneoensis, caspica, crassicollis, grandis, hamiltonii, japonica,
punctularia, reevesii, sinensis, subtrijuga, and trijuga.
Whereas all ‘emydids’ lack a contact, tortoises exhibit
both character states.
The presence of a contact between the sixth marginal scute and the third pleural scute was considered by
Hirayama (1985) to unite crown group ‘batagurids’ and‘testudinids’ as a synapomorphy. Given the patchy distribution of this character, and widespread polymorphism, it seems to be of little use.
Polarity: A contact between the sixth marginal scuteand the third pleural scute is absent in most outgroups.
The exception is caretta; this is not surprising because
caretta has five instead of four pleural scutes. We consider the presence of a contact to be derived.
(49) Twelfth marginal scute; 0 = two present, their common sulcus only partially subdivides the pygal bone,Fig. 98; 1 = two present, but their common sulcus fully
subdivides the pygal bone, Fig. 98; 2 = both twelfth
marginal scutes fused along the midline, Fig. 99 (modified from Mlynarski, 1976; Crumly, 1985, 1994, 35;
Gaffney and Meylan, 1988, H2.1).

2004

According to McDowell (1964, p. 240 and table 1
number 4) members of the ‘Emydinae’ can be distinguished from the ‘Batagurinae’ based on an incomplete
subdivision of the pygal bone by the median sulcus of
the posterior-most marginals. We found exceptions withpicta, N. platynota, pulcherrima, reevesii, reticularia,
and spengleri, which do not always clearly exhibit thepattern that would be predicted by McDowell’s (1964)
statement, but we note that the expression of this character will depend significantly on the shape of the pygalbone. In all tortoises except emys and impressa, thetwelfth marginal scutes are fused to from a single supracaudal scute that covers the posterior part of the carapace (Crumly, 1985). For this condition, we created athird character state.
Polarity: The twelfth marginal scutes are fully separated in all outgroups that have them, and their common sulcus fully subdivides the pygal bone. The pygalbone in “lindholmemydids” is polymorphic, with the 0state found in Lindholmemys and Mongolemys, and the
1 state in Gravemys. Either state 0 or state 1 is primitive
for Testudinoidea; the midline fusion of the twelfth marginals is a derived feature for ‘Testudinidae’.
Bridge(50) Sutured contact between plastron and carapace; 0= present, plastron and carapace are tightly connectedby an osseous bridge, Fig. 100; 1 = absent, plastron is
attached to carapace by connective tissue, Fig. 101(modified from Hirayama, 1985, Q; Shaffer et al., 1997,
58; Yasukawa et al. 2001, 21a).
(51) Presence and development of anterior buttresses; 0= anterior buttresses absent, Fig. 102; 1 = anterior buttresses present but small, and not in contact with the first
costal bones, Fig. 103; 2 = anterior buttresses long andthin and just barely in contact with the costal bones, if atall, Fig. 104; 3 = anterior buttresses well developed andin clear contact with the first costal bones, Fig. 105; 4 =anterior buttresses very large and in direct contact withthe first dorsal rib, Fig. 106 (modified from Hirayama,
1985, Q; Gaffney and Meylan, 1988, A14.2; Yasukawa
et al., 2001, 28).
(52) Presence and development of posterior buttresses;
0 = posterior buttresses absent, Fig. 107; 1 = posterior
buttresses present but small, and not in contact with thecostal bones, Fig. 108; 2 = posterior buttresses long andthin and just barely in contact with the costal bones, if atall, Fig. 109; 3 =posterior buttresses well developedand in clear contact with costal bones V and VI, Fig.
110; 4 = posterior buttresses well developed but only inclear contact with costal bone V, Fig. 111 (modified from
Hirayama, 1985, Q; Gaffney and Meylan, 1988, A14.2,
Shaffer et al., 1997, 55; Yasukawa et al., 2001, 29).

2004

Asiatic Herpetological Research(53) Medially-directed pivoting process for plastral
hinge developed on fifth peripheral bone; 0 = absent,
Fig. 112; 1 = present, Fig. 113.
(54) Complete or almost complete overlap of hyoplastron/hypoplastron suture by the pectoral/abdominal sulcus; 0 = absent, Fig. 114; 1 = present, Fig. 115 (modified Gaffney and Meylan, 1988, F3.2; Burke et al., 1996,
16; McCord et al., 1995, 13; Yasukawa et al., 2001,
21b).
In most testudinoid turtles, the plastron is attachedto the carapace via a fully ossified bridge and variably
developed plastral buttresses. In species with a kinetic
plastron, the bridge is typically absent and the plastron is
attached to the carapace via connective tissues (e.g.,
amboinensis, blandingii, carolina, dentata, galbinifrons,mouhotii, orbicularis, T. ornata). The original configuration of this character tied the presence of plastral kinesis to the reduction of the buttresses (Hirayama, 1985).
However, within testudinoid turtles the morphology ofthe buttresses varies significantly and independently
from plastral kinesis. We consequently split this character into three discrete characters concerned with themorphology of the bridge and the buttresses.
We also developed two new characters that pertain
to the morphology of the bridge region: the presence ofmedially-directed processes on the fifth peripherals thatact as pivots for the plastral bones during shell closure(Bramble, 1974) and a revised plastral kinesis character(Gaffney and Meylan, 1988; McCord et al., 1995) thatconsiders plastral-kinesis to be well developed only inthose taxa in which the pectoral/abdominal sulcus fully
overlaps the hyoplastron/hypoplastron suture, allowing
optimal movement between the two plastral lobes. Most
taxa with plastral kinesis also have well-developed pivoting processes on the fifth peripherals, but notable
exceptions are orbicularis and N. platynota. In
blandingii, the process is modified into an anteroposteriorly-elongated, flattened process that extends along
most or all of the length of the fifth peripheral (Fig. 113).
In some specimens of blandingii, a similar structure isdeveloped on the sixth peripheral as well.
Polarity: Reconstructing the basal condition forthese characters within testudinoids is difficult, because
all living cryptodiran outgroups do not have plastral buttresses and commonly lack osseous bridges. However,the bridge of Gravemys, Mongolemys, andLindholmemys is osseous, shows no signs of kinesis, and(at least in Lindholmemys) the anterior and posterior buttresses are well developed and touch the costal bones.
We consider that morphology to be primitive for
Testudinoids.

Vol. 10, p. 69(55) Contact between inguinal and femoral scutes; 0 =absent, Fig. 116; 1 = present, Fig. 117 (Crumly, 1985,
1994, 42; Gaffney and Meylan, 1988, H3.3, H15.2).
Within tortoises the complete or frequent absence ofa contact between the inguinal scute and the femoral
scute was used previously to hypothesize the monophyly of several smaller clades, for example {graeca + hermanni + horsfieldi + kleinmanni + marginata + tornieri}(Crumly, 1985) and {agassizii + berlandieri + flavomarginatus + polyphemus} (Gaffney and Meylan 1988). We
confirm the absence of a contact in representatives of thefirst group, but not in the second. Among the second
group (traditionally classified together in Gopherus) the
contact is strongly reduced, but still is present. Among‘batagurids’ and ‘emydids,’ a contact is absent in all taxa
with the noteworthy exception of hamiltonii.
Polarity: Determining the polarity for this characteris somewhat difficult because all living outgroups havean arrangement of plastral scutes that is rather different
from testudinoids. However, based on ingroup commonality and the absence of a contact in the“Lindholmemydidae,” we conclude that the presence ofa contact between the inguinal and femoral scutes
should be considered derived for Testudinoidea.
(56) Presence of musk glands; 0 = inguinal and axillary
gland present; 1 = axillary gland present only; 2 = musk
glands absent (modified from Crumly, 1985).
(57) Presence of anterior musk duct foramina; 0 = musk
glands and their foramina present, Fig. 118; 1 = musk
glands present, but foramina not developed; 2 = musk
glands and foramina absent (modified from Hirayama,
1985, A; Gaffney and Meylan, 1988, F1.2, F5.3; Burkeet al., 1996, 20).
(58) Presence of posterior musk duct foramina; 0 =musk glands and their foramina present, Fig. 119; 1 =musk glands present, but foramina not developed; 2 =musk glands and foramina absent (modified from
Hirayama, 1985, A; Gaffney and Meylan, 1988, F1.2,
F5.3; Burke et al., 1996, 20).
According to Hirayama (1985), the presence ofmusk duct foramina characterizes the paraphyletic
assemblage ‘Batagurinae’ (sensu Hirayama, 1985, notGaffney and Meylan, 1988). We believe the differenceof opinion between Hirayama (1985) and Gaffney andMeylan (1988) regarding this character is based on failure to make clear the distinction between the presence ofmusk glands and the presence of musk duct foramina.
Musk glands are developed in almost all extant turtles(Waagen, 1972), and we consequently agree with
Gaffney and Meylan (1988) that their presence should
be considered primitive for all cryptodiran turtles.

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However, even though most turtles have musk glands,
true musk duct foramina are developed only in somepleurodires (e.g., Chelodina, Emydura), some ‘emydids,’ and all ‘batagurids,’ making a monophyletic
Testudinoidea (sensu Hirayama, 1985) possible. Distinct
musk duct grooves are present on the anterior peripherals of Kinosternidae, and tiny foramina are sometimes
associated with these (Hutchison, 1991).
Because the presence of musk glands does not necessarily result in the development of musk duct foramina, we decided to score these two characters separately.
We relied on an unpublished thesis on the musk glandsof turtles (Waagen, 1972) to determine the presence ofmusk glands for most taxa. In scoring taxa not investigated by Waagen (1972) we only recorded presence ofmusk glands if musk duct foramina provided positive
evidence for their presence (e.g., baska, bealei, borneensis, galbinifrons, kachuga, mouhotii, petersi, pulcherrima, spengleri, and tentoria). Many tortoises, consequently, had to be scored as unknown, because they
were not analyzed by Waagen (1972) and do not exhibit
musk duct foramina (e.g., elongata, homeana, horsfieldi, and pardalis).
Our scoring of the musk duct foramina is derived
from a combination of osteological observation and data
provided by Waagen (1972). Taxa not reported to possess musk glands (Waagen, 1972) were checked formusk duct foramina, but none were found. For thosespecies that Waagen reported as having musk glands, wesought musk duct foramina on osteological specimens.
Several taxa with musk glands, but only lightly ossified
bridges, do not exhibit musk duct foramina (e.g.,
blandingii, dentata, flavomarginata, orbicularis, andpulcherrima) or show an asymmetry with foramina only
visible anteriorly (e.g., N. platynota). In taxa that possess them, the musk duct foramina are sometimes contained entirely within the peripherals (e.g., N. platynota), and sometimes between the peripheral and the plastral buttress (e.g., reevesii).
Polarity: Given the presence of musk glands in all
extant outgroups (Waagen, 1972), their absence should
be considered derived. Musk duct foramina are notdescribed for “lindholmemydids” but J. H. Hutchison
specifically searched for them in Mongolemys specimens housed at IVPP and found no trace of them.
Because musk duct foramina are developed in the vast
majority of the ingroup, we consider their absence to be
derived for testudinoids.
Plastron(59) Extensive overhanging lip of the epiplastra; 0 =absent, Fig. 120; 1 = present, Fig. 121 (Gaffney andMeylan 1988, H5.1, 9.2).
In most testudinoid turtles, the epiplastra are rather

2004

flat with a slight increase in thickness along the anterior
margin. In contrast, many tortoises have strongly thickened epiplastral margins that sometimes form an overhanging lip along the interior rim of the plastron.
According to Gaffney and Meylan (1988) the presenceof such an overhang of the epiplastra unites all tortoises
except those classified in the genera Manouria (Fig.
120) and Gopherus, with a reversal occurring in giant
insular forms. We confirm this general pattern, but wenote the absence of an overhang in tornieri. A interior
overhang is absent also in all extant ‘emydids’ and‘batagurids,’ but a small overhang is present in extinct
Ptychogaster and Echmatemys (Mlynarski, 1976, figs.
78, 81), taxa generally considered to belong to either the‘Emydidae’ or ‘Batagurinae.’
Polarity: An overhanging lip on the epiplastra isabsent in all outgroups and the majority of the ingroup.
We consider its presence to be derived.
(60) Intersection of the entoplastron by the humeropectoral sulcus; 0 = absent, Fig. 122; 1 = present, Fig. 123(Hirayama, 1985, X; Crumly, 1985; Gaffney andMeylan, 1988, F5.1; McCord et al., 1995, 15).
This character was used to help resolve relationships within ‘batagurids’ by Hirayama (1985) but wewere unable to replicate his results in our analysis. We
agree with Crumly (1985) that the sulcus crosses theentoplastron in at least one species classified in thegenus Indotestudo (i.e., elongata), but the sulcus is at theentoplastron/hyoplastron suture in our specimen of
forsteni. The condition in species now commonly classified in Testudo varies widely (e.g., the suture crosses theentoplastron in graeca and horsfieldi, but does not inhermanni or kleinmanni). We also agree with Gaffney
and Meylan (1988) on their distribution of this character
among the ‘Emydidae,’ however, our scoring for picta,
orbicularis, and blandingii is polymorphic, because thesulcus generally runs along the suture between the entoplastron and the hypoplastra, but may barely fall oneither side.
Polarity: The polarity is ambiguous if only extant
taxa are considered. The plastron of most outgroups is
too different from that of testudinoids to be of any use
for polarizing this character. For instance, the plastron ofspinifera lacks scutes, and that of serpentina, odoratus,and caretta is too heavily modified to enable a meaningful comparison. Both character states are commonly
found in ‘batagurids,’ ‘emydids,’ and testudinids, making an ingroup analysis futile. The humeropectoral sulcus is distinctly posterior to the epiplastron in Gravemys
and Mongolemys, so we consider an intersection of thissuture with the entoplastron to be derived.

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Asiatic Herpetological Research(61) Anal notch of the plastron; 0 = present, Fig. 124; 1= greatly reduced, Fig. 125; 2 = absent, Fig. 126 (modified from Hirayama, 1985, W; Yasukawa et al., 2001,
22).
The plastron of most testudinoid turtles has a significant anal notch. The absence of such an anal notch foramboinensis, galbinifrons, and flavomarginata wasreported by Hirayama (1985) and we confirm thoseobservations. An anal notch also is absent in belliana,
carolina, coahuila, erosa, homeana, T. nelsoni, and T.
ornata. To accommodate the presence of a reduced anal
notch we modify Hirayama’s (1985) character by creating a third character state. A reduced anal notch is foundin blandingii, orbicularis, N. platynota, and reticularia.
In at least one species (mouhotii), a distinct anal notch ispresent in larger individuals, but small specimens have a
reduced notch (scored as polymorphic in our matrix);
this suggests that development of an anal notch may besubject to ontogenetic variation in at least some testudinoids.
Polarity: In caretta and serpentina an anal notch is
not present, however, their plastra are narrow andtapered posteriorly. The fleshy plastron of spinifera issmooth along its posterior margin, but this cannot be
observed in osteological preparations. A notch is weakly developed in at least some Lindholmemys, but isabsent in Mongolemys. A notch is present in kinosternids, pleurodires, most of the ingroup, and in Gravemys.
We conclude that its absence should be consideredderived for our ingroup.
(62) Anal scutes fused; 0 = absent, Fig. 127; 1 = present, Fig. 127 (Hirayama, 1985, Z).
The anal scutes of adult galbinifrons, and flavomarginata are at least slightly fused, especially along theirposterior medial border. We fully agree with Hirayama’s(1985) treatment for this character. Anal scute fusion canbe identified easily in macerated specimens (Fig. 127),
because the anal scutes will not separate from one another, as will all other scutes.
Polarity: Anal scute fusion is absent in the vast
majority of turtles, and is considered to be the primitive
condition.
(63) Plastral scutes with vibrant, radiating color pattern; 0 = absent, Fig. 128; 1 = present, Fig. 129(Hirayama, 1985, S; McCord et al., 1995, 16 Yasukawa
et al., 2001, 32).
Vibrant, radiating color patterns of the plastral
scutes of dentata, grandis, and spinosa were noted by
Hirayama (1985) and McCord et al. (1995). We add
tcheponensis to this list, as well as the ‘testudinids’ geometricus and P. oculifera. In our specimens, the patternof dentata and tcheponensis is not as vibrant as in gran-Vol. 10, p. 71

dis and spinosa.
Polarity: Vibrant, radiated color patterns are missing in all outgroups and the majority of the ingroup.
Their presence is derived.
Postcranium and Soft Tissue(64) Development of a suprascapula; 0 = absent; 1 =present, Fig. 130 (Gaffney and Meylan, 1988, F3.1;
Burke et al., 1996, 11).
(65) Development of an episcapula; 0 = absent; 1 =present, Fig. 130 (Gaffney and Meylan, 1988, F4.1;
Burke et al., 1996, 11).
The presence of both a suprascapula and an episcapula apparently is limited to blandingii and the species
currently classified in Terrapene. A suprascapula is alsopresent in orbicularis. Both structures are involved in
the locking mechanism of the anterior plastral lobe during shell closure (Bramble, 1974). These structures are
difficult to verify in most osteological preparations,because they may dissociate from the scapula and be difficult to recognize, and because they may ossify only inolder individuals. The specimen we dissected to illustrate these features (TNHC 62532, a T. ornata with carapace length of 103 mm) has a completely cartilaginous
episcapula, and a predominantly cartilaginous suprascapula (Fig. 130). It is therefore much easier to confirm
their presence than verify their absence. We followed
Bramble’s (1974) account of these structures and scoredour matrix accordingly, as probably did Gaffney andMeylan (1988) and Burke et al. (1996).
Polarity: Suprascapulae and episcapulae are absent
in all outgroups and the majority of the ingroup. Their
presence is considered to be derived.
(66) Shape of coracoid blade; 0 = long and narrow, Fig.
131; 1 = short and very wide, Fig. 131 (Crumly, 1985,
1994; Gaffney and Meylan, 1988, H1.7).
The coracoid blade of all ‘emydids’ and‘batagurids’ is an elongate bone with a narrow, shortshaft and a long, wedge-shaped coracoid blade that isabout two times wider than the base. In tortoises, thisbone is still wedge-shaped, but relatively much shorter
and with a blade that is considerably wider, typicallyfour times the width of the base (Crumly, 1985, 1994;
Gaffney and Meylan, 1988). We agree with previously
published observations.
Polarity: The coracoid blade of caretta, serpentina,
and odoratus is long and narrow and that of spinifera islong, but not wedge-shaped. We consider a long and narrow coracoid blade to be primitive for Testudinoidea.
(67) Number of manual claws; 0 = five, Fig. 132; 1 =four, Fig. 133 (modified from Hirayama, 1985, J).

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Most testudinoid turtles have five manual claws
with the exception of baska and horsfieldi, both ofwhich have only four. We did not verify this character
independently for all species, due to an overall lack ofarticulated skeletons and our limited access to pickled
specimens. However, because the number of claws of
the forelimbs is an easily determinable, discrete numberthat is regularly noted and described in the literature, wescored all remaining taxa from the comprehensive information provided by Ernst and Barbour (1989).
Polarity: Five manual claws are present in serpentina, odoratus, and almost all pleurodires (excluding
species currently classified in Chelodina andHydromedusa); three are present in spinifera, and two incaretta. The condition in “lindholmemydids” isunknown. Based on ingroup commonality, we consider
five claws to be primitive for the ingroup.
(68) Number of phalanges of manus and pes; 0 = digital formula of 2-3-3-3-3 or 2-3-3-3-2, Fig. 132; 1 = digital formula with less than 2-3-3-3-2, Fig. 133 (Crumly,
1985; Gaffney and Meylan, 1988, H1.1).
The digital formula of most testudinoid turtles is 23-3-3-3 or 2-3-3-3-2. Among tortoises, the manus andpes are greatly shortened and the digital formula is typically reduced to 2-2-2-2-2 or less (Auffenberg,
1974:135-136; Crumly, 1985). Due to the dissociatednature of most of the material we viewed, we were notable to verify the digital formulae of most of the turtle
taxa we included. However, when articulated hands andfeet were present, we never found anything to contradict
the statements made above. We scored all tortoises
based on information provided by Auffenberg (1974)
and Crumly (1985).
Polarity: All outgroups and the majority of the
ingroup do not have a reduced digital formula. We consequently consider the reduced formula to be derived.
(69) Webbing between digits; 0 = present, well developed, Fig. 134; 1 = absent, or at least strongly reduced,
Fig. 135 (Hirayama, 1985, b).
Due to their semi-aquatic nature, most testudinoids
have well-developed webbing between the digits of theirhands and feet. In more terrestrial species, however, thewebbing often is reduced. Unfortunately, there seems to
be a gradient in the development of webbing, fromextremely well developed (e.g., baska, reticularia) tomoderately developed (e.g., dentata, guttata) to virtually non-existent (e.g., spengleri). We nevertheless wereable to reproduce Hirayama’s (1985) distribution for the‘batagurids’ with the exception of grandis and spinosa,
which have reduced webbing (grandis is the only ‘borderline’ taxon we found, but its webbing is reduced relative to those taxa we scored as having well-developed

2004

webbing). Among ‘emydids,’ we note that the webbing
is heavily reduced in carolina, T. nelsoni, and T. ornata.
All tortoises lack webbing.
Polarity: All outgroups and the majority of ingrouptaxa have webbed hands and feet. We consider theabsence of webbing to be derived.
(70) Sexual size dimorphism; 0 = absent; 1 = present,
female much larger than male (Gaffney and Meylan,
1988, F5.2; Burke et al., 1996, 37).
In almost every species of turtle, there is some
expression of sexual size dimorphism (Berry and Shine,
1980; Gibbons and Lovich, 1990). The difference in size
between the sexes can be expressed as a ratio and typically shows considerable variation depending on the
population (Gibbons and Lovich, 1990). We initiallytried to score this character with three character states, as
done by Burke et al. (1996), differentiating between
species with larger males, larger females, and equally
sized sexes, but we abandoned that, because exact data
are missing for most ‘batagurid’ taxa. We consequently
only score taxa as being sexually dimorphic if females
are at least 1.4 times larger than the males. Our scoresare derived from Gibbons and Lovich (1990) and Ernst
and Barbour (1989).
Polarity: Sexual size dimorphism is prevalent inmost outgroups. In spinifera the female is much larger,
in odoratus and serpentina the male tends to be slightly
smaller, in caretta the sexes are of similar size. The outgroup polarity is thus ambiguous, but in the majority of
the ingroup pronounced sexual dimorphism is absent.
Problematic Characters
We encountered difficulties in evaluating a number ofpreviously used characters, and we provide some summary statements for those in this section. Most of thesecharacters were not pursued thoroughly in our study
because we were not able to understand the originaldescriptions, were unable recover discrete character
states, or because at an early point in our investigation of
the character we detected significant variation in expression of character states within taxa. In the latter casegreater sample sizes or new methodological techniques(e.g., Wiens, 1995; Smith and Gutberlet, 2001) will be
required to tease out a phylogenetic signal.
(A) Frontal contribution to the supratemporal rim(Hirayama, 1985, 4).
The anterior extent of the upper temporal emargination is difficult to define in many taxa, and is impossible
to determine in those with a fully emarginated temporal
region (e.g., T. ornata). The result is a high degree of
ambiguity and a general lack of discrete character states.

2004

Asiatic Herpetological Research(B) Contact between postorbital and quadrate(Hirayama, 1985, 10).
In the vast majority of ingroup taxa, there is no contact between the postorbital and the quadrate. Such acontact was observed only in japonica and punctularia
by Hirayama (1985). In specimens of japonica available
to us, we were not able to confirm this contact. CAS
228348 is a skeleton from a diseased specimen of punctularia. On the right side of the skull there is a possible(but only slight) contact. It is possible that the contact is
actually between the postorbital and the quadratojugal(Fig. 136). We also found a minimal contact in one specimen of annulata. Given these diverging observations
and the minute contact that is present in our material, we
regard (for now) any contact within the ingroup as anabnormality.
(C) Absence of the “posterior process of the postorbital” (Hirayama, 1985, 8).
We cannot determine unambiguously what
Hirayama (1985) meant by this character. In our assessment of testudinoids, both a posterolateral and a posteromedial process of the postorbital can occur. In
Hirayama’s (1985) analysis, only grandis and spinosalack a “posterior process of the postorbital.” These
species are also the only ‘batagurid’ taxa to fully lack atemporal arch. We suspect that this character may somehow be referring to a lack of a bony temporal arch.
(D) Processus inferior parietalis “medially approximating each other, cranial cavity anteroventrally narrowing” (Hirayama, 1985, 5; McCord et al., 1995, 7;
Yasukawa et al., 2001, 2).
We acknowledge the validity of this character as
was originally worded by McDowell (1964). However,
we find it difficult to determine how strongly the constriction of the brain case must be before it can be considered present. We were unable to develop unambiguous discrete character states for this feature.
(E) Subdivision of the foramen nervi trigemini (Crumly,
1982; Hirayama, 1985, 6).
This character was used originally by Crumly(1982) to infer phylogenetic relationships within‘Testudinidae’. For his ingroup, Crumly (1982)
observed a great amount of polymorphism, with no single species either completely lacking or always exhibiting a subdivision of the foramen. He also noted asymmetry for this character between the left and right side ofsome individuals. We confirm the common presence ofa subdivided trigeminal foramen in representatives of‘Testudinidae,’ and the occasional presence in individuals of ‘Emydidae’ and ‘Bataguridae’ (e.g., areolata, dentata, flavimaculata, N. platynota, and rubida). A signifi-

Vol. 10, p. 73

cant amount of variation can be observed in two specimens of borneensis available to us, that exhibit left/rightasymmetry and the full spectrum from a fully intact (Fig.
137), to partially subdivided (Fig. 138), to fully subdivided (Fig. 139) trigeminal foramen. Given that mosttaxa are represented by three or fewer skulls in ourstudy, it is apparent that we are not able to fully document the amount of variation exhibited by testudinoids.
(F) Contact between postorbital and squamosal(Hirayama, 1985, 9).
Gaffney et al. (1991) noted that absence of this contact is associated with the upper temporal emarginationand considered it informative at the level of their analysis. Within our ingroup, all turtles have substantial upper
temporal emarginations, resulting in the contact being
just barely present, or just barely absent, or polymorphic(e.g., picta, petersi, texana, crassicollis). See comments
above under character 9.
(G) Median premaxillary notch (Hirayama, 1985, 19;
Yasukawa et al., 2001, 9)(H) Large cusps near the suture of the premaxillae andmaxillae (Hirayama, 1985, 28).
Initially, we were faced with the problem of defining these two characters independently from one another, because the presence of two tightly spaced, opposing
cusp-like structures along any margin will automaticallyresult in the development of a median notch. An additional problem relating to these characters is the question of whether these features should be observed on theramphotheca or the maxilla.
Large, tooth-like cusps are clearly present in a number of taxa (e.g., thurjii) but so is the full spectrum ofsmaller cusps, making it impossible to clearly define
discrete character states. Furthermore, if all species were
evaluated for medial notches that existed even if thecusps were removed, all taxa in our sample would show
a medial notch. We were unable to develop a consistentmethod for scoring this character for all testudinoid
species.
(I) “Antero-medial portion of the upper triturating surface formed by premaxillae and maxillae” (Hirayama,
1985, 23; Yasukawa et al., 2001, 13).
We are neither able to replicate the full meaning of
this character nor formulate truly discrete character
states. A connection with the development of the secondary palate is evident, but the morphology of this region
seems to be sufficiently covered by a number of othercharacters.

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Asiatic Herpetological Research(J) Participation of the vomer in the foramen praepalatinum (Crumly, 1982, 10; Hirayama, 1985, 32; Yasukawa
et al., 2001, 15).
Within testudinoid turtles, the foramen praepalatinum perforates the nasal cavity at the border between
the premaxilla and the vomer. When the foramen is positioned slightly more anteriorly, it is fully surrounded by
the premaxilla, when it is minutely farther posterior it issurrounded by the vomer. Considering the impact ofsuch minute changes, it is not surprising that our scoring
for this character generally seems to be in conflict withthat of Hirayama (1985) and Crumly (1982). This character appears to be subject to great intraspecific variation.
(K) Foramen palatinum posterius enclosed within thebrain cavity (Hirayama, 1985, 34).
According to Hirayama (1985), in reevesii (only)
the foramen palatinum posterius is enclosed within theregion of the brain cavity due to a flared descending
process of the parietal. We cannot confirm this observation for any testudinoid turtles (including three specimens of reevesii).
(L) Participating bones in the processus trochlearis
oticum (Hirayama 1985, 37; Gaffney and Meylan, 1988,
Gaffney et al. 1991, 6; McCord et al., 1995, 8; Shaffer etal., 1997, 74, 258 Yasukawa et al., 2001, 18).
The relative participation of the prootic, parietal,
and quadrate to the processus trochlearis oticum wasused previously by a number of authors to infer phylogenetic relationships within turtles. Our observations
confirm the great variety of morphologies that can beobserved in this region. However, the amount of
intraspecific variation is considerable and the full spectrum of possible morphologies seems to be filled, making it difficult to discern discrete character states. Future
research in the area may result in more clearly defined
discrete character states.
(M) Length of the crista supraoccipitalis (Hirayama,
1985, 40).
A long crista supraoccipitalis was observed by
Hirayama (1985) for borneensis. The character states heused are defined by relative length of the crista supraoccipitalis to the “condylo-basal length.” Unfortunately,
we could not replicate this because it is not clear exactly how the length of the crista was measured.
Furthermore, a true morphological gap seems to be
missing between the admittedly very long crista ofborneensis and other ‘batagurids’ with an elongatedcrista. This character is problematic, because it is poorly defined and lacks discrete character states.

2004(N) Bony sutures and sulci lost in old adults (Hirayama,
1985, I).
According to Hirayama (1985), loss of sutures andsulci occurs in baska, borneoensis, and borneensis only.
We are able to confirm this, but we do not have individuals of all other species that are sufficiently old enoughto positively confirm that they also do not exhibit thisfeature at old age. In subsequent treatments, loss of
sutures and loss of sulci should be treated as separatecharacters.
(O) Ossification of cornu branchiale II (Hirayama,
1985, 48; Yasukawa et al., 2001, 20).
This character was used previously to unite tortoises with a number of ‘batagurid’ taxa (Hirayama, 1985).
The hyoid apparatus of turtles is often disarticulated inskeletal preparations, making is difficult to positively
confirm if an ossified cornu branchiale is present orabsent. However, for those taxa for which we were able
to observe the hyoid apparatus, we were not able to confirm Hirayama’s (1985) observation of a reduced cornu
branchiale II in some ‘batagurids’ (e.g., galbinifrons,
spengleri). Instead, these taxa exhibit a cornu branchiale
II that is not significantly different from most other‘batagurids.’(P) Double articulation between the fifth and sixth cervical vertebrae (Hirayama, 1985; Gaffney and Meylan,
1988, F1.5).
Most articular surfaces of the cervical column are
rather homogenous within all testudinoid turtles(Williams, 1950). A double articulation between the fifthand sixth cervical previously was considered to be aunique character that unites the ‘Emydinae’ (McDowell,1964). This character also was used by Hirayama (1985)
and with reservations by Gaffney and Meylan (1988).
Our observations generally confirm the presence of amore or less clear double articulation in most ‘emydids,’
however, this features is also present in a number of‘batagurids’ confirming that this character is highly variable within the ingroup (Williams, 1950; Gaffney andMeylan, 1988). Unfortunately, discrete character states
are lacking; we were able to observe the full morphological spectrum from a clear singular articulation to a clear
double articulation.
(Q) Scapular prong with lateral concavity (Hirayama,
1985, E).
Hirayama (1985) reported this character as an
autapomorphy for subtrijuga only. However, we cannot
identify this morphology in any of our specimens of subtrijuga.

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Asiatic Herpetological Research(R) Large facet of the ilium for the testoscapularis andtestoiliacus (Hirayama, 1985, T; Yasukawa et al., 2001,
34)(S) Extensive development of both testoscapularis andtestoiliacus (Hirayama, 1985, U).
An extensive development of the testoscapularisand testoiliacus muscles together with an associatedlarge scar on the ilium was reported by Bramble (1974)
for Asian and North American box turtles. Whereas wehave no reason to doubt his assessment of the development of these muscles for box turtles, we were not able
to score this character for most of the remaining taxa.
The shape of the ilium was explored and illustrated forsome ‘emydids’ and ‘batagurids’ by Yasukawa et al.
(2001:122-123).
(T) Ossification of the epipubis (Gaffney and Meylan,
1988, F5.5).
The identification (and confirmation of presence orabsence) of an ossified epipubis (Fig. 141) is somewhat
difficult for most species, because it seems to ossify
rather late in ontogeny, and can fall off during preparation. Our tentative observations confirm the presence ofan ossified epipubis in numerous adult ‘emydids’ and‘batagurids,’ typically terrestrial forms (e.g., T. ornata,
G. insculpta, N. platynota, yuwonoi). An improved sample of adult specimens of all taxa, however, is necessary
to reveal the true distribution of this character.
(U) Diploid Number of Chromosomes (Hirayama, 1985,
Ø; Shaffer et al., 1997, 43).
The diploid number of chromosomes was used byCarr and Bickham (1986) to hypothesize a sister grouprelationship between the ‘Emydinae’ and subtrijuga, followed by borneensis and crassicollis and finally the restof the ‘Batagurinae.’ Whereas most ‘batagurids’ allegedly have 52 chromosomes, subtrijuga, borneensis, crassicollis, and ‘emydids’ are supposed to have 50 chromosomes. We view these results with caution, because abrief review of the relevant literature reveals great differences in chromosomal counts for a variety of taxa.
For instance, according to the work of Killebrew (1977)
and Bickham (1981), amboinensis has 52 chromosomes,
however, Gorman (1973) reported only 50. Similar conflicts can also be found for dentata (Bickham, 1981;
DeSmet, 1978; Gorman, 1973; Stock, 1972), subtrijuga(Bickham, 1981; Killebrew, 1977), trijuga (DeSmet,
1978; Carr and Bickham, 1986), sinensis (Bickham,
1981; Killebrew, 1977; Stock, 1972), crassicollis(Killebrew, 1977; Stock, 1972; Bickham and Baker,1976), and some of the species currently placed in
Graptemys (Killebrew, 1977; McKown 1972),
Trachemys (DeSmet, 1978; Killebrew, 1977; Stock,Vol. 10, p. 75

1972) and Mauremys (Killebrew, 1977; Stock, 1972).
This conflict in primary data is probably best understoodwhen considering the nature of testudinoid chromosomes: whereas 14 pairs of chromosomes have a considerable size, all of the remaining ones are extremely
small. Given these circumstances, it seems reasonable tohypothesize that one pair of chromosomes may beunrecognized during analysis.
(V) Plica media spade-shaped (Gaffney and Meylan,
1988, F7.1).
The penile soft anatomy of turtles was comprehensively reviewed by Zug (1966) and one of his characters,
the shape of the plica media, was used by Gaffney andMeylan (1988) to unify species placed in Chrysemys,
Deirochelys, Trachemys, and Pseudemys as a monophyletic group. In his detailed description of the plica
media, Zug (1966) referred to the shape of this structure
as being “spade-shaped” in those taxa, but made similar
claims for other taxa too. Furthermore, based on theillustrations that were provided by Zug (1966) for othertaxa, the plica media of species placed in Graptemys,
Malaclemys, Rhinoclemmys, and Platysternon appear“spade-shaped” also, even though Zug (1966) did not
explicitly use those descriptive words. This anatomicalsystem should be carefully reevaluated for all testudinoids, with special attention given to definition of discrete characters.
(W) Ossifications within the fenestra postotica.
In some taxa, portions of the fenestra postotica areclosed or obscured by ossifications (noted, but without
exemplars, by Gaffney, 1972). In our largest specimenof grandis (CAS 228443) a short, spike-like ossified
process extends posterodorsally from the dorsal edge ofthe quadrate process of the pterygoid, and crosses thefenestra postotica. It is situated ventral to the stapes (columella auris), medial to the incisura columella auris, andlateral to the fenestra ovalis (Fig. 141). In some specimens, the dorsal tip of a similar structure approaches ormeets a posterodorsally-inclined process that extends
from the dorsal surface of the pterygoid, near the suturewith the prootic. In our largest specimen of N. platynota(CAS 228342) the two processes meet to enclose thestapedial shaft in a ring of bone situated at approximately the midpoint between the fenestra ovalis and the
medial opening of the incisura columella auris. Ourother, smaller, specimen of N. platynota shows no development of these processes. It seems likely that there isan ontogenetic component in the expression of this feature. It does not appear to have any systematic significance, but in any case it is not widespread within
Testudinoidea.

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Conclusions
Our observations can be used to draw some tentative conclusions regarding the current level of understanding about morphological variation within testudinoid turtles. In addition, we provide some cautionary
statements about the quality of morphological data now
in use for assessing systematic relationships among
these turtles. It is clear from a perusal of the relevant literature, and from our data, that there is reasonably
strong morphological support for a monophyletic‘Testudinidae.’ Support for monophyly of ‘emydines’
and ‘batagurines’ is not as impressive. The paraphyly of‘Batagurinae’ (with respect to ‘Testudinidae’) was
explicitly proposed by Hirayama (1985) and has been
generally accepted since that time, although some
strides have been made towards resolving relationships
among some ‘batagurine’ taxa. The monophyly of‘Emydinae’ seems to have been implicitly assumed bymany workers, but remains to be established in the context of a rigorous phylogenetic analysis of all relevant
taxa. The interpretation of several morphological features shared between some ‘emydines’ and some‘batagurines’ as either convergence or synapomorphy
remains an important and interesting challenge. For
example, there are intriguing morphological similarities
between subtrijuga and some species classified in thegenus Graptemys (e.g., contact of the jugal and descending process of the parietal; contact of the quadratojugal
with the articular facet of the quadrate; contact between
the quadratojugal and the maxilla; ventral process of thepterygoid approaching the articular surface of thequadrate). These similarities may be due to functional
convergence as a result of a molluscivorous diet, butthey raise questions about the propriety of utilizing subtrijuga as an outgroup for systematic studies of ‘emydines’ (e.g., Burke et al., 1996). Additional similarities
are reported for chromosome numbers in subtrijuga and‘emydines’ (see ‘Problematic Character’ U, above).
A seriously deficient understanding of morphological variation is one of the greatest inadequacies of current perspectives on morphological data in turtles generally, and testudinoids specifically. Few published studies
have been conducted to evaluate the range and causes ofsexual, ontogenetic, intra- and inter-population variationin morphological characters within testudinoids. Our
preliminary considerations of ontogenetic variation,
combined with reports of sexual variation (e.g., Berryand Shine 1980; Gibbons and Lovich, 1990; Stephens,
1998) and new studies exploring the complex interactions of morphological evolution with behavioral characteristics and environmental conditions in turtles (e.g.,
Lindeman, 2000; Herrel et al., 2002; Joyce and Gauthier,2003; Claude et al., 2003) emphasize the importance of
pursuing these questions further. Our decision not to

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produce a phylogenetic hypothesis in this paper was
based primarily on two considerations. The first is therelatively small sample size we used for many taxa inthis study (although it is comparable to sample sizes
from other, previously published, studies), and the factthat several taxa are not represented in our work. The
second consideration is our sense that the current understanding of morphological variation in testudinoid turtles is insufficiently mature to permit reliable phylogenetic hypotheses based solely on morphological data.
The most expedient way to address the need for greater
documentation of variation within testudinoid species is
to utilize existing museum collections to the greatest
extent possible, and secondarily to develop responsible
collecting programs that are designed with this need inmind.
Our results also indicate that morphological data
matrices currently in the literature should not be taken atface value. We had particular difficulties replicatingsome of the scoring in the Hirayama (1985) matrix. Thatseminal analysis (completed prior to the widespread useof computer-assisted analytical methods in systematics)
laid the foundation for nearly all subsequent work on‘batagurine’ morphology (including our own), and its
importance in shaping our current conceptualization of‘batagurine’ phylogeny cannot be overstated. The workof pathfinders in all fields of inquiry is often subjectedto the greatest scrutiny by the next generation ofresearchers. Our statements and contradictory observations in this paper in no way denigrate Hirayama’s work;
instead, we view our efforts as minor attempts to correct
the few inconsistencies in his analysis, and to contributeour observations to the body of knowledge that he began
to synthesize 20 years ago.
The accurate interpretation and understanding of
the morphological descriptions of previous authors wereamong the great challenges we faced when we began ourstudies of testudinoid skeletal morphology. Much of ourconfusion could have been averted if adequate illustrations accompanied all published character descriptions,but such documentation often is an expensive undertaking. Our photographs of character states discussed inthis paper are intended to facilitate communication
among turtle enthusiasts, and to provide a baseline for
future comparisons and discussions about testudinoid
morphology. We hope that adequate illustration of all
newly proposed characters will become standard practice among turtle systematists. It seems likely that ourinterpretations of characters will differ in some respects
from those of our colleagues, and we anticipate that ourdecisions regarding ‘problematic characters’ (discussed
above), and our choices with respect to ‘lumping’ or‘splitting’ previously published character states, will
generate much spirited discussion in the years ahead.

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As practicing vertebrate paleontologists, we also
hope that our efforts here will stimulate additional investigation and publication of the extensive fossil record of
testudinoid turtles. Morphology is, of course, of paramount importance for the interpretation of fossils. Our
visits to many museums in the last several years
revealed an abundance of unpublished testudinoid fossilmaterial. Although a phylogenetic analysis was not agoal of this research, we adopt the convenient and now
familiar means of summarizing morphological data by
providing a character matrix that summarizes some of
our observations. Although character data can be used toassist paleontologists in diagnosing fossil testudinoid
specimens, we also encourage paleontologists to utilize
our study as a starting point for basic morphologic
descriptions. Description of differential diagnostic characters generally is an adequate minimum for the erectionof a new taxon. However, such a diagnosis is, in itself,
not particularly helpful to systematists trying to score amatrix and place a fossil into a broader phylogenetic
context. Descriptions of new fossil specimens (and taxa)
will be most useful if they include discussions of character state data for all preserved anatomical regions.
This current summary of morphological characters thattraditionally are used in systematic treatments of testudinoids can be used as a preliminary guide to the anatomical regions and features that would be most useful when
included as part of a description of new fossil material.

Acknowledgements
We are greatly indebted to a number of institutions
and their staff for providing generous access to skeletal
and wet specimens: Jens Vindum (CAS), Stephen
Rogers and John Wiens (CM), Alan Resetar andMaureen Kearney (FMNH), John Simmons and Linda
Trueb (KU), Jim McGuire (LMNH), José Rosado(MCZ), Travis LaDuc and David Cannatella (TMM),
Harold Dundee (TUMNH), and Gregory CaldwellWatkins and Jacques Gauthier (YPM). We extend ourthanks to Rick Van Dyke for the gift of specimens to
support this research. Gabe Bever, Julien Claude, Jason
Downs, Jacques Gauthier, Jason Head, Pat Holroyd,Howard Hutchison, Lyndon K. Murray, Jim Parham, andKrister Smith (in alphabetical order) provided usefulcomments and criticisms regarding this project. All
images containing YPM specimens were reproducedwith courtesy of the Peabody Museum of Natural
History, Yale University, New Haven, CT. Special
thanks go to Ted Papenfuss and Jim Parham for providing us with the unique opportunity to publish the resultsof this study, and to Julien Claude and Howard
Hutchison for their detailed and insightful reviews. The
Brett Sterns Award for Chelonian Research (CAS), theVisiting Scholar Fund (FMNH), and Yale University

Vol. 10, p. 101

Graduate Fellowships provided funding to WGJ.
Additional financial support was provided by the
Geology Foundation of The University of Texas atAustin.

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Appendix 1
List of specimens used.
agassizii, CAS 33867, FMNH 216746, FMNH 250843;
alabamensis, CM 95968, CM 95991; amboinensis, CAS
153872, CAS 228345, CAS 228369, CAS 228412, CAS
228419, FMNH 224009; annandalei, FMNH 258876,
FMNH 258879, FMNH Moll3036; annulata, CAS
SUR7425, CM 87903, YPM 15410; arachnoides, MCZ
54050; areolatus, MCZ 42214; areolata, CM 47957,
CM 87904; barbouri, CAS SUR12063, TUMNH 15400,
TUMNH 15428, TUMNH 15429, TUMNH 16899;
baska, FMNH 224095, FMNH 224097, FMNH 224213,
FMNH 224124, FMNH 224226; bealei, CM 118554,
FMNH 255270, FMNH 226542; belliana, CAS 228394,
CAS 228404; berlandieri, TNHC 2813; blandingii,
CAS 12837, CAS 228346, CAS 228373, CAS 228434,
CAS 228448, YPM 14677, YPM 10382; borneensis,
FMNH 224001, FMNH 224003, FMNH 224004,
FMNH 224005, FMHH 224140; borneoensis, FMNH
224107, FMNH 224122, FMNH 224129, FMNH
251499, MCZ 42198; carbonaria, CAS 228397, CAS
228411, CAS 228427, CAS 228430; caretta, CAS 8383,
FMNH 31021; carolina, CAS 228375, CAS 228378;
caspica, CAS 141118, CM 118517, FMNH 19714,
FMNH 74505, FMNH 98764; chilensis, CM 112252;
coahuila, KU 46929, KU 92623, MCZ 120335; crassicollis, CAS 228335, CAS 228451, FMNH 11091, MCZ
7821, MCZ 134451; decorata, CM 118590; dentata,
CAS 134332, KU 47170, MCZ 29567, CAS 228333,
CAS 228362, CAS 228414; elegans, CAS 228396; elongata, CAS 228408, FMNH 183740, FMNH 257382;
emys, CAS 228344, FMNH 63749, FMNH 224034; ernsti, TUMNH 13460, TUMNH 13462, TUMNH 16899;
erosa, YPM 12653; flavimaculata, TUMNH 15375,
TUMNH 15404, TUMNH 15747, TUMNH 15787;
flavomarginata, CAS 18040, CAS 228356, CAS
228357, CAS 228359, FMNH 216515; flavomarginatus,
MCZ 164926; forsteni, CAS 228433; galbinifrons, CAS
228358, CAS 228360, CAS 228367, CAS 228376, YPM
14074, YPM 14080; geographica, CAS 12809,
TUMNH 15391, TUMNH 15392, TUMNH 15410,
TUMNH 15430; geometricus, MCZ 32184; gibba, CAS
228392; graeca, CAS 217732, CAS 228435, LMNH
54855, LMNH 54861; grandis, CAS 228341, CAS
228443, FMNH 224038, YPM 15431; guttata, CAS
8696, CAS 228372, CAS 228386, FMNH 211573;
hamiltonii, MCZ 120333, YPM 10399; hermanni, CAS
228400, CAS 228401, CAS 228402; homeana, CAS
228409, CAS 228423, CAS 228428, CAS 228429, CAS
228455, FMNH 19794; horsfieldi, CAS 120707, CAS
228398, CAS 228399, CAS 228421, CAS 228425, YPM
16450; impressa, CAS 228416; G. insculpta, CAS
228406, CAS 228407, CAS 228413; japonica, YPM

Vol. 10, p. 105

15482, YPM 15486; kachuga, FMNH 224128, FMNH
224152; kleinmanni, CAS 228422, CAS 228426, CAS
228431; kohnii, TUMNH 10237, TUMNH 12121,
TUMNH 14544, TUMNH 15678; leprosa, CM 137031;
marmorata, CAS no number, CAS 188533, CAS
220052, FMNH 22076; mouhotii, CAS 228365, CAS
228374, CAS 228420, CAS 228444; muhlenbergii,
LMNH 55352, MCZ 52248; mutica, LMNH 54883; P.
nelsoni, CM 67311; nigra, CAS 8125, CAS 8289;
nigrinoda, TUMNH 15147, TUMNH 15317, TUMNH
15408, TUMNH 15750; ocellata, CAS 228336; G.
oculifera, TUMNH 26, TUMNH 3359, TUMNH 7548,
TUMNH 12402, TUMNH 16928; P. oculifera, CAS
165598, CAS 220645, CAS 220646; odoratus, YPM13622, CAS 228351, CAS 228352, CAS 228353; orbicularis, CAS 173223, CAS 228347, CAS 228415, CAS
228447, LMNH 10347, YPM 15479; T. ornata, CAS
228381, CAS 228382, CAS 228383, CAS 228384,
TNHC 62532; ouachitensis, CM 61656, CM 84696;
pardalis, CAS 148630, CAS 228349, CAS 228410,
CAS 228418, CAS 228432, CAS 228449; petersi, CAS
8608, CM 124246, CM 124247; picta, CAS 13889, CAS
228379, CAS 228380, CAS 228385; N. platynota, CAS
228342, CAS 228450, CM 118586, FMNH 224216,
YPM 14678; polyphemus, CAS 14090, LMNH 43344,
LMNH 43354, LMNH 59320; pulcherrima, CAS
11754, CAS 228355, CAS 228366, CAS 228377; pulchra, LMNH 48092; punctularia, YPM 465, CAS
228348; reevesii, CAS 31437, CAS 228361, CAS
228364; reticularia, CAS 228338, CAS 228388, LMNH
14565, LMNH 14569, LMNH 49145, LMNH 54856,
LMNH 78580; rubida, CM 87907, CM 87908;
rubriventris, CM 34409, CM 45188; scripta, CAS
SUR8642, CAS 228436, CAS 228442, LMNH 15684;
serpentina, CAS 228452, YPM 6369, CAS 228457;
siebenrocki, CAS 228393; signatus, CAS 228405, MCZ
42217, MCZ 42218; sinensis, CAS 18031, CAS
228339; spengleri, CAS 21008, CAS 228331, CAS
228332, CAS 228343, CAS 228371, YPM 11653, YPM
14445; spinifera, CAS 65705, CAS 228350, CAS
228354, YPM 656; spinosa, CAS 228368, CAS 228459;
subglobosa, CAS 228334; subtrijuga, CAS 16996, CAS
228445, CAS 228446, CAS 228453, CAS 228454; subrufa, CAS 228389, CAS 228390, CAS 228391; tcheponensis, CAS 228363, CAS 228370; tecta, CM 89923;
tentoria, FMNH Moll3026, FMNH Moll3028, FMNH
Moll3032, FMNH 259430; tentorius, MCZ 41944,
MCZ 46604; terrapin, CAS 43640, CAS 228340, CAS
228387, YPM 2983; texana, CAS 30965, CAS 228437,
CAS 228438, CAS 228439, CAS 228440, CAS 228441,
CAS 228456, CAS 228458; thurjii, FMNH 224135,
FMNH 224153, MCZ 62523, MCZ 62524, YPM 14072,
YPM 14073, YPM 14074; tornieri, CAS 139704, CAS
228395, CAS 228403, CAS 228417, CAS 228424; tri-Vol. 10, p. 106

Asiatic Herpetological Researchfasciata, CAS 228337, MCZ 5218; trijuga, CAS 12463,
CM 89921, CM 124227, YPM 15453; versa, TUMNH
4484, TUMNH 10510, TUMNH 16192; yuwonoi, YPM
12626.
Appendix 2
Generic synonymies for all currently recognized testudinoid species and all outgroup species used herein.
Synonymies are based only on common usage during
the last 50 years.
Ingroupadiutrix
agassizii
alabamensis
amboinensis
angulata
annamensis
annandalii
annulataarachnoides
areolata
Rhinoclemmys
areolatus
aurocapitata
barbouribaska
bealeibelliana
berlandieri
blandingii
borneensis
borneoensis
boulengeri
caglei
callirostriscarbonaria
carolina
caspicachilensis
coahuilaconcinnacrassicollis
decorata
decussatadentatadenticulata
depressa
dhongokadiadematadorbigni

Chrysemys, Pseudemys, Trachemys
Gopherus, Xerobates
Chrysemys, Pseudemys
Cuora
Chersina, Goniochersus, Neotestudo,
TestudoAnnamemys, Mauremys
Hieremys
Callopsis, Geoemyda, Nicoria,
Rhinoclemmys
Pyxis
Callopsis, Geoemyda, Nicoria,
Homopus
Cuora
Graptemys, Malaclemys
Batagur
Clemmys, Sacalia
KinixysGopherus
Emydoidea, Emys, Neoemys
Orlitia
Callagur
Homopus
Graptemys
Chrysemys, Pseudemys, Trachemys
Chelonoidis, Geochelone
Terrapene
Clemmys, Mauremys
Chelonoidis, Geochelone
Terrapene
Chrysemys, Pseudemys
Siebenrockiella
Chrysemys, Pseudemys, Trachemys
Chrysemys, Pseudemys, Trachemys
Cyclemys, Geoemyda
Chelonoidis, Geochelone
Geoemyda, Heosemys
Kachuga
Geoemyda, Rhinoclemmys
Chrysemys, Pseudemys, Trachemys

dussumieri

2004

Aldabrachelys, Dipsochelys,
Geochelone, Megalochelys
elegans
Geochelone
elongata
Geochelone, Indotestudo
emolli
Chrysemys, Pseudemys, Trachemys
emys
Geochelone, Manouria
ernsti
Graptemys
erosa
Kinixysfemoralis
Homopusflavimaculata
Graptemys, Malaclemys
flavomarginata Cistoclemmys, Cuora, Geoemyda
flavomarginatus Gopherus
floridana
Chrysemys, Pseudemys
forsteni
Geochelone, Indotestudo, Manouria
funerea
Callopsis, Geoemyda, Rhinoclemmys
gaigeaeChrysemys, Pseudemys, Trachemys
galbinifrons
Cistoclemmys, Cuora
geographica
Graptemys, Malaclemys
geometricus
Psammobates
gibbonsi
Graptemys
graecaTestudograndisGeoemyda, Heosemys
guttata
Clemmyshamiltonii
Geoclemys
hermanni
Testudohomeana
Kinixyshorsfieldi
Agrionemys, Testudo
impressa
Geochelone, Manouria
G. insculpta
Calemys, Clemmys, Glyptemys
japonica
Clemmys, Mauremys
kachuga
Kachugakleinmanni
Pseudotestudo, Testudo
kohnii
Graptemys
leprosa
Clemmys, Mauremys
leytensis
Geoemyda, Heosemys
lobatsiana
Kinixysmarginata
Testudomarmorata
Clemmys, Emysmelanosterna
Geoemyda, Rhinoclemmys
mouhotii
Cuora, Cyclemys, Geoemyda, Pyxidea
muhlenbergii
Calemys, Clemmys, Glyptemys
muticaClemmys, Mauremys
nasutaCallopsis, Geoemyda, Rhinoclemmys
natalensis
Kinixysnebulosa
Chrysemys, Pseudemys, Trachemys
P. nelsoni
Chrysemys, Pseudemys
T. nelsoni
Terrapene
nigra
Chelonoidis, Geochelone
nigricans
Chinemys
nigrinoda
Graptemys, Malaclemys
ocellata
Morenia
G. oculifera
Graptemys, Malaclemys
P. oculifera
Psammobates
oldhamii
Cyclemys2004

Asiatic Herpetological Research

orbicularisEmys
T. ornata
Terrapene
P. ornata
Chrysemys, Pseudemys, Trachemys
ouachitensis
Graptemys
pani
Cuora
pardalisGeochelone
petersiMorenia
picta
Chrysemys
planicauda
Acinixys, Pyxis
G. platynota
Geochelone
N. platynota
Notochelys
polyphemus
Gopherus, Xerobates
pseudogeographica
Graptemys, Malaclemys
pulcherrima
Callopsis, Geoemyda, Rhinoclemmys
pulchraGraptemys, Malaclemys
pulchristriata
Cyclemys
punctularia
Geoemyda, Rhinoclemmys
quadriocellata Clemmys, Sacalia
radiata
Asterochelys, Geochelone
reevesii
Chinemys
reticularia
Deirochelys
rubidaCallopsis, Geoemyda, Nicoria,
Rhinoclemmys
rubriventris
Chrysemys, Pseudemys
scripta
Chrysemys, Pseudemys, Trachemys
signatus
Homopussilvatica
Geoemyda, Heosemys
sinensis
Ocadiasmithii
Kachuga, Pangshuraspekii
Kinixysspengleri
Geoemyda
spinosa
Geoemyda, Heosemys
stejnegeri
Chrysemys, Pseudemys, Trachemys
subtrijuga
Malayemys
sulcata
Geochelone
sylhetensis
Kachuga, Pangshurataylori
Chrysemys, Pseudemys, Trachemys
tcheponensis
Cyclemys, Geoemyda
tecta
Kachuga, Pangshura
tentoria
Kachuga, Pangshura
tentorius
Psammobates
terrapen
Chrysemys, Pseudemys, Trachemys
terrapin
Malaclemys
texanaChrysemys, Pseudemys
thurjii
HardellatornieriMalacochersus
travancorica
Geochelone, Indotestudo
tricarinata
Geoemyda, Melanochelys
trifasciata
Cistoclemmys, Cuora
trijuga
Geoemyda, Melanochelys
trivittata
Kachugavenusta
Chrysemys, Pseudemys, Trachemys
versa
Graptemys, Malaclemys
werneri
Testudoyaquiayniphora
yunnanensis
yuwonoizhouiVol. 10, p. 107
Chrysemys, Pseudemys, Trachemys
Asterochelys, Geochelone
Cuora
Geoemyda, Leucocephalon,
Notochelys
Cuora

Outgroup
caretta
Caretta
dumerilianus
Peltocephalus, Podocnemis
C. insculpta
Carettochelys
fimbriatus
Chelusgibba
Phrynops, Mesoclemmys
madagascariensis
Erymnochelys, Podocnemis
megacephalum Platysternon
odoratus
Kinosternon, Sternotherus,
Sternothaerus
scorpioides
Kinosternon
serpentina
Chelydrasiebenrocki
Chelodina, Macrochelodina
spiniferaApalone, Trionyx
subglobosa
Emydura, Tropicochelymys
subrufa
Pelomedusa
temminckii
Macroclemys, Macrochelys
Appendix 3
Distribution of character states for 70 characters among46 species of testudinoid turtles; a = 0/1; b = 0/2; c = 0/3;
d = 0/1/2; e = 1/2; f = 1/3; g = 2/3.
agassizii
11a10 00000
01000 03200
00000 10110
amboinensis
00a00 00010
00000 01110
20000 00000
annandalei
0010? 00020
00100 01100
00000 0000?
annulata00a00 00010
00000 03100
00000 00010
barbouri00001 11000
00001 10000
00000 00001
baska
00001 00000
11100 00010

000?0
00000

11010
00120

00110
22001

00001
22200

00001
00200

00000
0a011

00010
11110

00000
00001

00011
00000

10001
00a10

00010
34000

00000
00001

0010a
00000

00000
000?0

00010
11000

00000
0??01

01010
00000

10000
00000

10000
34000

101a0
22200

00010
00000

10001
10110

00000
43000

00a02
00000

Vol. 10, p. 108
00000 01000
bealei
00000 00010
00000 00000
00000 00000
blandingii
00000 00000
00000 0c000
10011 00000
borneensis
00100 00000
00000 00000
00000 00000
borneoensis
00001 00000
11100 00000
00000 00000
carbonaria
11010 00000
00000 01201
00000 10110
caspica
00000 00010
00000 00000
00000 00000
crassicollis
00a00 00000
00001 00000
00000 00000
elongata
11010 00000
00000 03200
00000 10110
dentata
00000 00010
00001 01100
00100 00000
emys
11010 00000
00000 01000
00000 10110
flavomarginata
00010 00020
00000 01110
21000 00010
graeca11010 00000
00000 03200
00000 10110
grandis0010a 00020
00001 01100

Asiatic Herpetological Research

00000
0??00

10000
0a010

00010
34000

00000
0000?

000?0
00000

00000
01001

10000
11110

00000
1120a

00010
00010

00000
00a10

00010
33000

10000
00000

00010
00100

10001
00a10

00010
43000

00001
00000

000?a
00000

11000
00120

00111
22001

00001
22210

0000a
00000

00000
0aa10

00010
34000

00000
00001

00000
00010

10000
00a10

00010
33000

10000
00001

000?a
00000

11000
00020

00111
22001

00001
???11

00000
00001

00000
00011

00000
11000

00000
01101

000?a
02000

11000
00aa0

00110
22001

00001
???00

0000a
00100

00000
00011

00010
11110

00000
?1101

000?1
00000

11000
00a20

10111
22000

00001
22211

00000
0000a

00000
00a10

000a0
34000

00000
00001

00100 00010
guttata
00010 00000
00000 00000
00000 00000
hamiltonii
01101 00001
00111 00000
00000 00000
homeana
11010 00000
00001 0110a
20000 10110
horsfieldi
11010 00000
00000 03200
00000 11110
kachuga
00101 00000
11100 00000
00000 0000?
mouhotii
00010 00010
00011 01100
a0000 00010
P. oculifera
11010 00000
00001 01200
00000 10110
orbicularis
00000 00000
00000 00000
10010 00000
T. ornata
00010 00010
00000 00010
20011 00010
pardalis
11010 00000
00000 0f201
00000 10110
petersi
01100 00000
12100 02000
00000 00001
picta
00001 10000
00000 00000
00000 0000a
N. platynota
01101 00010
00001 00000

2004

00000
00000

00000
00000

10000
34000

00000
10201

00000
00000

10000
00a10

00010
33001

00110
0000a

000?1
00000

11000
00020

00111
22001

00000
???1a

000?0
00000

11000
00020

a0111
22000

00001
???11

00010
01000

10001
10110

00010
43000

10001
00000

00000
00000

11100
00011

00010
11110

00000
00001

000??
00000

11000
00120

00111
22001

00001
???10

00000
00000

00000
0a001

a0000
11010

00000
1120a

00000
00200

00000
00001

10000
00110

00000
ed201

000?a
00000

11000
00120

00111
22001

00001
???10

00010
00100

10001
00110

00010
34000

01111
00000

00010
00000

00000
0a0a0

10000
34000

00000
2220a

00000
10100

00000
00001

00a10
03010

00000
00101

2004
10000 00000
polyphemus
11010 00000
01000 0a200
00000 10110
pulcherrima
00a00 00010
00000 01100
00000 00000
reevesii
00101 10000
00110 00000
00000 00000
reticularia
00000 10000
00000 00000
10000 00001
scripta
00001 10000
10000 00000
00000 0000a
sinensis00001 00000
00000 00000
00000 00000
spengleri
000a0 00000
00011 01100
00000 00010
subtrijuga
00100 01100
00110 00g00
00000 00000
tentoria
00001 00000
10100 10000
00000 0000?
terrapin
00001 10000
00000 00000
00000 00001
texana00001 10000
10000 00000
00000 0000a
thurjii
00001 00000
11100 00000
00000 00001
tornieri11010 00000
00000 0??00

Asiatic Herpetological Research
00000
000?0
00000

11000
00120

a0110
22001

00001
22200

00001
00000

00000
000a0

a0010
14000

00000
01?01

00000
00000

10000
00aa0

00010
33000

10110
00001

000?0
00000

00000
010a0

10000
34000

00000
22200

00010
00000

10000
00000

10000
34000

00101
22200

00010
00000

10001
00aa0

10010
33000

00001
00001

00100
00000

10100
00000

00010
34000

00000
00001

11000
00000

10000
00a10

01010
33000

10100
00000

00010
01000

10001
10110

00010
43000

10001
00000

00010
00000

10000
0a000

a0000
34000

10100
ed200

0a010
00000

10000
00000

a0000
34000

00101
22200

00010
00100

10001
00110

00000
43000

01011
00000

000?1
00000

11000
00020

00111
22000

00001
2220a

10110

Vol. 10, p. 109