1 REPTILIA: TESTUDINES: abaxillare

Catalogue of American Amphibians and combination. 927 Kinosternon cruentatum abaxillare: Artner 2003:xix. First use of combination. E. Reyes-Grajales and J. B. Iverson. 2020. Kinosternon scorpioides abaxilare: Acuña and Kinosternon abaxillare. Merchán 2003:31. Lapsus.

Kinosternon abaxillare Baur CONTENT. No subspecies are recognized. in Stejneger 1925 Central Mud DESCRIPTION. Kinosternon abaxillare is a medium to large Kinosternon, with males in Kinosternon abaxillare Baur in Stejneger some populations exceeding 156 mm in car- 1925:462. Type locality, “Tuxtla [Gutiér- apace length, and females reaching 160 mm. rez], Chiapas, .” Holotype, United The carapace is moderately to strongly tricar- States National Museum (USNM; now inate in all but the oldest, largest individuals. Smithonian Institution National Museum The shell is somewhat depressed (shell height of Natural History) 7518, the shell of an averages 35% of carapace length in males, adult male collected by Dr. C. H. Berendt, 38% in females). The first vertebral scute is apparently in 1863 or 1864 (see Smith and wider than long; vertebral scutes 1–4 have Smith 1980). The holotype is lost, but the distinct posterior notches at the midline in all junior author has examined the 11 para- but the oldest, largest individuals. In imma- types (USNM 7519-29; Cochran 1961). ture individuals, the second vertebral shield is Kinosternun abaxillare: Alvarez del Toro the longest, and in mature specimens (of both 1973:20. Lapsus. sexes) the third vertebral shield is the longest; Kinosternon scorpioides abaxillare: Berry in both size classes the fifth vertebral shield is 1979:3186-B (also Ernst and Barbour the shortest. The tenth marginal scute is high- 1989; Smith and Smith 1980). First use of er than the ninth. The shape of the carapace is

Figure 1. Male Kinosternon abaxillare from Villa Hidalgo, Chiapas, México. Photograph by Eduardo Reyes-Grajales. 2

Map. Distribution of Kinosternon abaxillare in Chiapas, México, and Huehuetenango, . The red dots indicate localities verified by the authors and the blue dots represent localities verified by collaborators (see Acknowledgments). The red triangles designate unverified records. approximately oval (more evident in imma- pletely closes the shell (width of posterior ture individuals and females than in males); plastral lobe averages 47% of maximum car- the margins are distinctly flared outward in apace length in males, 48% in females) with most populations. The carapace is highly adult males having a smaller plastron than variable in color, ranging from light brown to females. Maximum plastron length averages olive to black, with darker seams in all but the 100% of maximum carapace length in males, darkest individuals. and 99% in females (Reyes-Grajales 2019). The plastron has two kinetic hinges, an- The posterior plastral lobe bears only a tiny terior and posterior to the abdominal scutes, posterior notch, or more typically none at and is concave to flat in males but slightly all. Bridge length averages 28% of carapace convex or flat in females. The plastron com- length in males and 31% in females. The in- Account 927 3

Figure 2. Female Kinosternon abaxillare from Villa Hidalgo, Chiapas, México. Photograph by Eduardo Reyes-Grajales. terabdominal seam length averages 30% of skin of other soft parts is gray or brown, usu- carapace length in males and 33% in females. ally with many small, darker spots. There are Axillary scutes are usually absent, although three to four pairs of gular barbels, with the the axillary-abdominal scute seam may be in- anterior pair the largest. Elevated patches of complete in some individuals (<15% in a pop- horny scales (‘clasping organs’ or ‘vinculae’) ulation). For medial length of plastral scutes, are absent from the posterior thigh and leg the most common formula for all class sizes of both males and females. Males have lon- and sexes is interabdominal > interanal > gu- ger, more prehensile tails than do females; the lar > interhumeral > interfemoral > interpec- cloacal aperture is at or posterior to the car- toral. The interabdominal scute seam is long, apace margin in males but is anterior to the averaging 29% of maximum carapace length carapace margin in females. The tails of both (27–33%), and the interpectoral seam is short sexes have terminal spines, but the spines are (3%). The color of the plastron may be yellow, larger in males. orange, brown or black, usually with darker seams. The dorsal head shield is rhomboi- DIAGNOSIS. The body size is medium rel- dal, bell-shaped, or triangular. The maxillary ative to other Kinosternon (to only 170 mm sheaths are weakly to strongly hooked, more carapace length). In addition to its localized strongly hooked in older males than in fe- geographic distribution, the most important males or younger individuals. Head markings character distinguishing Kinosternon abax- consist of yellow, cream, or pale gray dots or illare from congeners is the lack of axillary reticulations on a gray or olive background. scutes; however, some individuals (< 10%) The jaw sheaths are cream to yellow (more have full or partial axillary scutes (Iverson evident in adult females) with darker vertical 2008; Reyes-Grajales 2019). streaks most conspicuous in older males. The 4

Figure 3. Details of the carapace (top row) and plastron (bottom row) of an adult female (left column) and an adult male (right column) of Kinosternon abaxillare from Villa Hidalgo, Chiapas, México. Photo- graphs by Eduardo Reyes-Grajales. PHYLOGENETIC RELATIONSHIPS. The (1960, 1973, 1982), Berry (1978, 1979), Ber- relationships among the of the ry and Iverson (2001), Bonin et al. (2006a, Kinosternon are still uncertain. A tree based 2006b), Legler and Vogt (2013), Reyes-Gra- on combined mtDNA + nuDNA presented by jales (2018a, 2018b), Smith and Smith (1980), Iverson et al. (2013) contained a clade with and Stejneger (1925). Specific descriptions Kinosternon abaxillare and Kinosternon oaxa- include comparisons with other Kinosternon cae as sister species, then most closely related (Kinosternon chimalhuaca, Kinosternon hirti- to a clade comprising Kinosternon scorpioi- pes, Kinosternon integrum, Kinosternon oaxa- des cruentatum and Kinosternon scorpioides cae, Kinosternon scorpioides and Kinosternon scorpioides. A subsequent phylogenetic anal- sonoriense; Reyes-Grajales 2019); shell shape ysis of the Kinosternidae (Spinks et al. and morphometrics (Iverson, 2008, 2010; 2014) did not include Kinosternon abaxillare, Reyes-Grajales et al. 2020; Sánchez-M. et al. but the sister species identified in the analysis 2000), and lack of cloacal bursae (Smith and by Iverson et al. (2013), Kinosternon oaxacae, James 1958). was most closely aligned with Kinosternon in- tegrum and Kinosternon sonoriense. ILLUSTRATIONS. Color photographs were published by Alvarez del Toro (1960, 1973, CONSERVATION STATUS. Not evaluated. 1982; dorsal views), Arai (1998; general pro- file), Legler and Vogt (2013; dorsal and ven- PUBLISHED DESCRIPTIONS. General de- tral views of male and female), Reyes-Graja- scriptions were provided by Alvarez del Toro les (2019; general, ventral, dorsal, head views Account 927 5

Figure 4. Dorsal (left column) and ventral (right column) views of adult, juvenile, and neonate (top to bottom) of Kinosternon abaxillare from Villaflores, Chiapas, México. Photographs by Eduardo Reyes-Gra- jales. of neonates, juveniles, males and females), and medium elevations across its range. The Turtle Working Group (2017; gen- general distribution was described by Turtle eral profile), and Vetter (2005; general profile, Taxonomy Working Group (2017). Addition- the plastron, the side of the head, and the al distributional information is available for bridge). Black-and-white photographs were México (Alvarez del Toro 1960, 1973, 1982; presented by Alderton (1988a, 1988b), Berry Bannikov 1969; Bonin et al. 1998, 2006a, (1978; dorsal, ventral, head and bridge area), 2006b; Bour 2004; Darevskiy and Orlov 1988; Freiberg (1972; general profile), and Pritchard Ferri 1999, 2002; Fotolulu 2018; Fritz and Ha- (1979a; general profile and plastron). vaš 2007; Iverson 1985, 1986, 1992; Johnson 1989; Nelson and Nickerson 1966; Rhodin et DISTRIBUTION. Kinosternon abaxillare is al. 2008; Smith and Smith 1980; Turtle Taxon- endemic to the Grijalva River basin in the omy Working Group 2007, 2009, 2010, 2011, Central Depression of Chiapas and north- 2012, 2014). west Guatemala (Reyes-Grajales et al. 2020; Sánchez-M. et al. 2000). Our personal data RECORD. None. and information presented by Rogner (1996) suggest that Kinosternon abaxillare occurs at PERTINENT LITERATURE. General re- altitudes of 300 to 860 m. This taxon occurs in views were published by Alvarez del Toro a variety of permanent, semipermanent, and (1960, 1973, 1982), Berry (1978, 1979), Ber- temporary aquatic habitats, primarily at low ry and Iverson (2001), Casas Andreu (1965, 6

2019; Reyes-Grajales et al. 2020, in review; Sánchez-M. et al. 2000), feeding (Reyes-Gra- jales 2018a, 2018b; Sánchez-M. et al. 2000), habitat (Alvarez del Toro 1960, 1973, 1982), longevity (Mertens 1970; Nöllert 1992; Slavens 1981, 1982, 1987, 1988; Slavens and Slavens 1991, 2000), morphology (Berlant and Stayton 2017; Casas Andreu 1965, 1967; Gijzen and Wermuth 1958a, 1958b; Iverson 1984; Kilias 1957; Smith and Smith 1980), Figure 5. Presence of axillary scutes in a juve- parasites (Reyes-Grajales 2018a, 2018b), nile of Kinosternon abaxillare from San Fernan- phylogeny (Bickham and Carr 1983; Iverson do, Chiapas, México. Photograph by Eduardo 1988; Iverson et al. 2013; Ramirez Guerra Reyes-Grajales. 2016; Spinks et al. 2014), population size and density (Iverson 1982; Reyes-Grajales 2018a, 1967), Gijzen and Wermuth (1958a, 1958b), 2018b, 2019; Reyes-Grajales et al. 2020; Sán- Pritchard (1967, 1979a, 1979b), Smith and chez-M. et al. 2000), predators (Reyes-Gra- Smith (1980). This taxon is listed in numer- jales, 2018a, 2018b), reproduction (Alvarez ous species checklists (David 1994; Ernst and del Toro 1973, 1982; Bull et al. 1974; Iver- Barbour 1989; Ferri 1999, 2002; Ferri and son 2008; Macip-Ríos et al. 2017; Moll 1979; Soccini 2015; Fritz and Havaš 2007; Iverson Vázquez Gómez et al. 2015), sexual dimor- 1985, 1986, 1991, 1992; Jordan 1972; King phism (Ceballos and Iverson 2014; Iverson and Burke 1989; Kirkpatrick 2006; Maldo- 2008; Reyes-Grajales 2019; Reyes-Grajales nado-Koerdell in Beltran 1953; Mertens and et al. 2020), size and growth (Iverson 2008; Wermuth 1955; Rhodin et al. 2008; Schilde Reyes-Grajales 2019), taxonomy (Anony- 2001; Smith and Smith 1976; Turtle Taxono- my Working Group 2007, 2009, 2010, 2011, 2012, 2014; Welch 1994; Wermuth and Mer- tens 1961, 1977). Other important references are activity (Reyes-Grajales 2019; Reyes-Gra- jales et al. 2020), anatomy (Parsons 1968; Smith and James 1958), blood and blood proteins (Sullivan and Riggs 1967a, 1967b, 1967c), captive breeding (Lucas and Biegler 1971; Pawley 1971; Slavens 1976), chromo- somes and karyotypes (Bickham and Carr 1983; Gilboa 1975; Moon 1972, 1974; Sites et al. 1979), climate change effects (Berrioza- bal-Islas 2018; Berriozabal-Islas et al. 2018), conservation status (Macip-Ríos et al. 2015; Rhodin et al. 2018), distribution (Amori and Luiselli 2019; Bannikov 1969; Bour 2004; Darevskiy and Orlov 1988; Johnson 1989; Figure 6. Dorsal view of hatchling (top) and Johnson et al. 2015; Köhler 2000; Smith and egg (bottom) of Kinosternon abaxillare from Villa Smith 1980; Smith and Taylor 1950a), ecology Hidalgo, Chiapas, México. Photograph by Eduar- (Iverson 1982; Reyes-Grajales 2018a, 2018b, do Reyes-Grajales. Account 927 7

Figure 7. Typical habitat of Kinosternon abaxillare in Villa Hidalgo, Chiapas, México. Photographs by Eduardo Reyes-Grajales. mous 2014; Berry 1978; Bickham and Carr 1982, 1987, 1988; Slavens and Slavens 1991, 1983; Bonin et al. 1998, 2006a, 2006b; Bour 2000; Smith and Smith 1980; Smith and Tay- 2004; Casas Andreu 1965, 1967; Cochran lor 1950b, 1966; Turtle Taxonomy Working 1961; CONABIO 2009; Darevskiy and Or- Group 2007, 2009, 2010, 2011, 2012, 2014), lov 1988; Forero-Medina and Castaño-Mo- and vernacular names (Darevskiy and Or- ra 2011; Fotolulu 2018; Frair 1972; Greene lov 1988; Fotolulu 2018; Franklin 2007; Lin- [1971]; IUCN/SSC and Freshwater er 1994; Obst 1986a, 1986b; Reyes-Grajales Turtle Specialist Group 1989; Iverson 1989; 2019; Wrobel 2004). Iverson et al. 2013; Liner 2007; Mueller 1998; Müller 1987; Otani and Kawazoe 2018; Rho- ETYMOLOGY. The specific nameabaxillare din et al. 2008; Rogner 1996; Rojas-Runjaic is a descriptive adjective derived from the et al. 2015; Rust 1938; Slavens 1976, 1981, Latin axillares, referring to the axilla (axillary

Figure 8. Variation in color and pigment in the carapace of Kinosternon abaxillare. Photographs by Eduardo Reyes-Grajales. 8 scute, in this case), and preceded by the pre- non scorpioides cruentatum: Biology and fix ab meaning “without;” hence, “without an taxonomic revision of the red-cheeked axillary scute.” mud turtle in Costa Rica. Reptilia 26:30- 34. ADDITIONAL VERNACULAR NAMES. Alderton, D. 1988a. & of the People in the Tuxtla Gutierrez region often call World. Facts on File, Inc., New York, New this turtle casquito, casquito moteada, casqui- York. 191 pp. to de cabeza pintada, tortuga, tortuga apesto- Alderton, D. 1988b. Turtles & Tortoises of the sa (unpublished observations by the authors), World. Blandford Press, London, United and casquito pardo (Liner 1994). This species Kingdom. 191 pp. has also been referred to as the Chiapas Mud Alvarez del Toro, M. 1960. Reptiles de Chi- Turtle (Obst 1986a, 1986b; Sokolov 1988; apas. Instituto Zoológico del Estado, Tux- Wrobel 2004), Чьяпасская Замыкающаяся tla Gutierrez, Chiapas, Mexico. 204 + [1] Черепаха (Darevskiy and Orlov 1988), Chi- pp. apas-Klappschildkröte (Müller 1987; Sokolov Alvarez del Toro, M. 1973 (“1972”). Los Rep- 1988); Central Chiapas Mud Turtle (Artner tiles de Chiapas. Segunda Edicion. Gobi- 2008; Berry and Iverson 2011; Franklin 2007; erno del Estado de Chiapas, Tuxtla Guti- Legler and Vogt 2013; Liner 1994; Muel- errez, Chiapas, Mexico. [3] + 178 + [1] pp. ler 1998; Turtle Taxonomy Working Group Alvarez del Toro, M. 1982. Los Reptiles de 2017), Zentral-Chiapas-Klappschildkröte Chiapas. Tercera Edición. Instituto de (Rogner 1996), Cinostserne [sic] de Chiapas Historia Natural, Tuxtla Gutierrez, Chi- (Wrobel 2004), Mexican Red-spotted Mud apas, Mexico. 248 pp [+ erratum sheet]. Turtle (Slavens 1981, 1982, 1987, 1988), and Amori, G. and L. Luiselli. 2019. Scale-depen- Baur’s Skorpion-Klappschildkröte (Fotolulu dent ‘functional’ assessment of the con- 2018). cept of endemism. Biological Journal of the Linnean Society 127:900–911. REMARKS. Kinosternon abaxillare was con- Anonymous. 2014. Recently proposed chang- sidered a subspecies of Kinosternon scorpioi- es of scientific names of turtles. Turtle des for 34 years; consequently, many general Survival, August 2014:62. literature references to Kinosternon scorpioi- Arai, [K.], [荒井, 敬子] (editor). 1998. Tur- des between the years of 1979–2013 do not tles Collection. Aqua Magazine Collec- pertain to Kinosternon abaxillare. tion Series 4. フェア・ウインド [Fair Wind Co., Ltd.], 佐倉市 Sakura City, Ja- ACKNOWLEDGMENTS. We thank An- pan. 116 + [28] pp. ders Rhodin, Rodrigo Macip-Ríos, Yessenia Artner, H. 2003. Nomenklatur aktuell. Die Sarmiento, Roberto Luna-Reyes, and Manuel rezenten Schildkrötenarten der Erde. E. Acevedo for locality information, and for 10(6):iv-xxxviii. providing information for literature cited. We Artner, H. 2008. The world’s extant turtle spe- thank Christian Rico for creating the map. cies, part 2. Emys 15(4):4-35. ERG thanks the Turtle Conservation Fund Bannikov, A. G. (editor) [Ванникова, А. and Chelonian Research Foundation for their Г., р е д.]. 1969. Жизнь Животных. support of fieldwork and laboratory research Земноводные, Пресмыкающиеся. Том (TCF-0760). 4. Часть 2 [Zhizn Zhivotnykh. Zemno- vodnye Presmykayushchiesya. Tom 4. LITERATURE CITED Chast’ 2] [ Life. Amphibians, Rep- Acuña, R. A. and M. Merchán. 2003. Kinoster- tiles. Volume 4, Part 2]. Издательство Account 927 9

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