REPTILIA: SQUAMATA: DIPSADIDAE Cubophis Vudii

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903.1 REPTILIA: SQUAMATA: DIPSADIDAE Cubophis vudii

Catalogue of American Amphibians and Reptiles.

Powell, R. 2014. Cubophis vudii.

Cubophis vudii (Cope) Bahamian Racer, Bahamian Brown Racer

Alsophis vudii Cope 1862:74. Type locality, “New Providence Id., Bahamas.” Syntypes, Academy of Natural Sciences of Philadelphia (ANSP) 5567, 5569–71, 5598–99, four adults (or subadults) and two juveniles (based on SVL Figure 1. An adult Cubophis vudii vudii from measurements of 53.3 cm, 43.5 cm, Cape Eleuthera, Eleuthera. Photograph by the 41.3 cm, 24.1 cm, 28.9 cm, and 40.5 author. cm, respectively; all measurements on preserved specimens), sexes unknown, sandy, taupe, gray, and reddish to reddish donated by Dr. H. C. Wood, Jr. on 7 May brown, dark brown, and black. Variable 1861 (examined by N. Gilmore at the dorsal patterns include uniform coloration, request of the author). faint lateral stripes, flecked or suffused with Dromicus angulifer: Boulenger 1894:120 darker color, irregular, diffuse banding, (part). combinations of flecking and banding, or Alsophis angulifer vudii: Stejneger 1905:337. a combination of faint lateral stripes and Alsophis rudii: Engelmann and Obst 1981:185. flecking. Tops of heads can be unpatterned or Lapsus. with a diffuse or sharply defined patterns that Alosophis vudii: Buckner 1993:19. Lapsus. might or might not include preorbital stripes Ocyophis vudii: Zaher et al. 2009:147. See but almost always with dark postorbital Phylogenetic Relationships. stripes that terminate on the temporals. In Cubophis vudii: Hedges et al. 2009:9. some populations, supralabials are bright orange, with the orange color extending onto CONTENT. Five subspecies are currently the neck. Ventral ground color varies from recognized: Cubophis vudii vudii, C. v. white, pale yellow, cream, and pinkish tan to aterrimus, C. v. picticeps, C. v. raineyi, and C. reddish brown, reddish, orange, pale orange, v. utowanae, but see Remarks. orange-gray, and brown, sometimes grading into black posteriorly. Venters range from DEFINITION. Cubophis vudii is a relatively unpatterned to variably suffused or spotted small “racer,” reaching a maximum known with tan (with dark borders), dark brown, or SVL of 84 cm (Franz and Dodd 1994). Dorsal black. Undersides of tails are pale yellow with scales are in 17 rows at midbody; ventrals various modifications based on the nature number 169–181 in males and 159–166 in of the ventral pattern. The color of the iris is females, subcaudals 112–120 in males and usually creamy tan above and reddish brown 101–120 in females. Supralabials usually below. number 8/8, infralabials usually 10/10, preoculars 1/1, postoculars 2/2, temporals DIAGNOSIS. Cubophis vudii is the only 1 + 2/1 + 2, and loreals 1/1 (Schwartz and “racer” in the Bahamas. Considering the Henderson 1991). variability in color, pattern, and ventral/ Dorsal ground color varies from tan, subcaudal scale counts, geographic origin 903.2

Map 1. Distribution of Cubophis vudii. Circles mark type localities and dots denote other records. Some symbols may represent multiple proximate locations. Guide to subspecies (numbered and demarcated by solid lines): 1. C. vudii vudii, 2. C. v. aterrimus, 3. C. v. picticeps, 4. C. v. raineyi, 5. C. v. utowanae (modified from Schwartz and Henderson 1991).

is the best way to distinguish this species Cope (1862), although Boulenger (1894) had from its congeners (C. cantherigerus, Cuban placed it in the genus Dromicus (which is Archipelago; C. caymanus, Grand Cayman now in the synonymy of several West Indian Island; C. fuscicauda, Cayman Brac; C. ruttyi, alsophiine genera). Zaher et al. (2009), based Little Cayman). on hemipenial morphology, resurrected the genus Ocyophis Cope for Hypsirhynchus ater PHYLOGENETIC RELATIONSHIPS. (type species) and five other species, including Cubophis vudii evolved as part of a C. vudii. Hedges et al. (2009) disagreed monophyletic West Indian radiation that because that grouping was not supported by probably originated from a single Mid an expanded molecular data set. They instead Cenozoic dispersal event from South America described the new genus Cubophis with five (Cadle 1985; Hedges 1996a, 1996b, 1996c; species (including C. vudii). Grazziotin et Vidal et al. 2000). Until recently, the generic al. (2012) recovered a polyphyletic Ocyophis arrangement within that radiation generally (sensu Zaher et al. 2009) and supported the followed Maglio (1970). He had included C. recognition of Cubophis. Pyron et al. (2013) vudii in the widely distributed genus Alsophis, found Cubophis monophyletic, not a sister to which it had been assigned originally by group to either Alsophis or Hypsirhynchus. 903.3

Figure 2. Syntypes of Alsophis vudii Cope 1862 (1863) from New Providence Island, Bahamas. All specimens accessioned at the Academy of Natural Sciences of Philadelphia (ANSP). Specimens in first column, top to bottom: ANSP 5567, 5569, 5570; specimens in second column, top to bottom: ANSP 5571, 5598, 5599. Images are not at same scale. Photographs by Ned Gilmore (ANSP).

PUBLISHED DESCRIPTIONS. In addition in supplementary information), Coborn to the original descriptions in Barbour and (1991), EOL (2014), Hedges (2013), Knapp Shreve (1935), Conant (1937), and Cope and Pagni (2011, eating an Andros Iguana, (1862), detailed descriptions are in Schwartz Cyclura cychlura cychlura; that photograph and Henderson (1991). was reproduced in Lemm and Alberts 2012), Mehrtens (1987), and Uetz and Hallermann ILLUSTRATIONS. Color photographs (2014). Additional color photographs are are in Bimini Biological Field Station on websites of the Bahamas National Trust (2004), Calsbeek and Cox (2010, eating (2012), Bahamas National Trust Education a Brown Anole, Anolis sagrei; available Office (2009), and The Center for Snake 903.4

Conservation (Young 2011). Black and white 1976), phylogenetics and systematics (Cadle photographs are found in Engelmann and 1984; Crother 1999b; Crother and Hillis Obst (1981, electrophoretic blood albumin 1995; Grazziotin et al. 2012; Hedges et al. separation) and Franz et al. (1996). Black and 2009; Maglio 1970; Vidal et al. 2000; Zaher et white illustrations include line drawings of al. 2009), predation (Mittermeier 2011), and individuals in Attrill et al. (1983) and Gape venom (Hayes 2008; Weinstein et al. 2011). and Sweeting (2004); Schwartz and Rossman The species also has been included in (1976) illustrated a left prefrontal bone. checklists, keys, general works, faunal accounts, indices, and popular works by Attrill DISTRIBUTION. Cubophis vudii is endemic et al. (1983), Barbour (1904, 1906, 1914, 1916, to and widely distributed in the Bahama 1930, 1935, 1937), Barbour and Loveridge Islands, where it occurs in essentially every (1929, 1946), Bright (2006), Buckner (1993), available habitat (Schwartz and Henderson Buckner et al. (2012), Buden (1981), But- 1991). Detailed island-by-island lists are in terfield et al. (1997), Cochran (1934, 1961), Buckner et al. (2012). The range was illustrated Cope (1894a, 1894b), Crother (1999a), Cun- previously in Schwartz and Henderson dall and Irish (2008), de Queiroz and Rodrí- (1991), Wieg (2009), and Hedges (2013). guez-Robles (2006), Dodd (2003), Dodd and Franz (1996), Frank and Ramus (1995), Franz FOSSIL RECORD. Late Pleistocene vertebrae and Buckner (1998), Franz et al. (1993), Frye from two individuals on New Providence (1991, 1994, 1995), Hass et al. (2001), Hayes Island were assigned to this species, with et al. (2004), Henderson and Binder (1980), the largest individual having an estimated Henderson and Powell (2004, 2009), Hender- snout-vent length of 880 mm (Pregill 1982). son and Sadjak (1986), Hutchins et al. (2003), Steadman et al. (2014) reported late Holocene James (2008), Kluge (1984), Knapp and Ow- fossils from Abaco Island. ens (2005), Knapp et al. (2011), Lawing et al. (2012), MacLean et al. (1977), Malnate (1971), PERTINENT LITERATURE. Relevant Matthews (1959), Mehrtens (1987), Meshaka citations are listed by topic: biogeography (2003), Miller (1968), Obst et al. (1984, 1988), (Maglio 1970; Schwartz 1968; Wieg 2009), Pavlidis (2002), Powell et al. (1996), Pregill diet (Barbour 1906; Bauer and Russell 1992; (1986), Pyron et al. (2011), Rabb and Hayden Calsbeek and Cox 2010, 2012; Calsbeek et al. (1957), Reinhardt and Lütken (1863), Rodrí- 2010; Cox and Calsbeek 2010a, 2010b; Franz guez-Robles and Greene (1996, 1999), Rosén and Dodd 1994; Henderson and Bourgeois (1911), Schmidt (1936), Schwaner and Des- 1993; Henderson and Crother 1989; Knapp sauer (1982), Schwartz and Henderson (1985, 2005; Knapp and Hines 2006a, 2006b; Knapp 1988, 1991), Schwartz and Thomas (1975), and Pagni 2011; Knapp et al. 2004, 2005, 2010; Sheehy (2006), Sokolov (1988), Stejneger Henderson and Sajdak 1986; Henderson et (1905), Tipton (2005), Uetz and Hallermann al. 1987; Meshaka 2001; Olson 2011; Olson (2014), Wallach (1998), Wallach et al. (2014), et al. 2012), distributional record (Crother Walley (1997), Werner (1929), and Wrobel and Slowinski 1987), eponym (Beolens et (2004). al. 2011; Shea 2012), growth (Franz and Dodd 1994), meristics and morphology REMARKS. Maglio (1970), based on skeletal (Burton 1939; Maglio 1970; Wieg 2009), morphology, suggested that C. v. utowanae natural history (overviews) (Henderson might be distinct at the species level, but Wieg and Powell 2009;Schwartz and Henderson (2009) disagreed. Wieg (2009:3), however, 1991), nomenclature (Follett 1958), overwater suggested that “the current subspecies dispersal (Knapp 2000), parasites (Franz designations oversimplify the variation 903.5 within the species and are not an adequate on Bimini Island; EOL 2014; Frank and Ra- reflection of variation present among the mus 1995; Hayes 2008; Wrobel 2004). island populations.” Barbour (1914) stated that Diadophis 1. Cubophis vudii vudii (Cope) rubescens Cope from New Providence Island, Bahamas (Cope 1885, 1900), belongs to this Alsophis vudii Cope 1862:74. See species species (= A. vudii). Boulenger (1884) had synonymy. previously referred this taxon to Dromicus Diadophis rubescens Cope 1885:403. Type angulifer, to which Stejneger (1905) referred locality, “New Providence, Bahama as “Alsophis angulifer (vudii).” The latter Islands.” Holotype, Academy of Natural (p. 337) also indicated that the single pore Sciences of Philadelphia (ANSP) 3472 on each scale “would preclude its being an (fide Barbour 1914), juvenile (sex not Alsophis, while on the other hand I can see determined), donated by Dr. H. C. no good reason why it may not be a form of Chapman (date of collection unknown) Leimadophis more or less closely allied to L. (examined by N. Gilmore at the request andreæ (Reinhardt and Luetken).” of the author). See Remarks. Halsophis vudii: Cope 1987(1888):439. COMMENT. This species is included in Leimadophis (?) rubescens: Stejneger Schedule 4 (“species indigenous to The 1905:337. See Remarks. Bahamas that are not included in Schedule Alsophis vudii vudii: Barbour and Shreve 1 and are believed to be endangered”) in 1935:363 (by inference). Chapter 250A (Wildlife Conservation and Ocyophis vudii vudii: Zaher et al. 2009:147 Trade Act) of the Statute Law of The Bahamas. (by inference). Cubophis vudii vudii: Hedges et al. 2009:9. ETYMOLOGY. The specific epithet vudii is a patronym honoring “my friend, Dr. H. C. DIAGNOSIS. This subspecies is characterized Wood, Jr., author of memoirs on Myriapoda by having 159–167 ventrals, 112–117 and extinct Cryptogamia” (Cope 1862:74). subcaudals, and a proportional tail length of The subspecific nameaterrimus is from the 0.301–0.319. Dorsal and ventral color and Latin ater (= black) and –imus (= having the patterns extremely variable, encompassing quality of), almost certainly pertaining to the most of the variations listed in the species dark color of these snakes; picticeps is from description, but usually not dark brown to the Latin (= painted head), presumably in black above or mostly black below. Tops reference to the markings on the heads of of heads and dorsum with sharply defined these snakes; raineyi is a patronym for Mr. patterns. The former usually includes light- Froelich Rainey of the Peabody Museum at bordered preorbital stripes and almost always Yale University, who served as the archeologist include dark postorbital stripes that terminate on the 1934 Utowana voyage; and utowanae on the temporals and do not extend onto the commemorates the research yacht Utowana. neck. Undersides of the tails are pale yellow and range from unpatterned to variously ADDITIONAL VERNACULAR NAMES. Vari- marked with darker pigment. ations including “Bahamian” or “Bahaman” in combination with “Racer” or “Brown Rac- 2. Cubophis vudii aterrimus (Barbour and er” are not listed. Other English-language Shreve) vernacular names used in the literature are Brown Runner (Franz and Dodd 1994) and Alsophis vudii aterrimus Barbour and Bimini Racer (not restricted to populations Shreve 1935:362. Type locality, “High 903.6

Rock, Grand Bahama Island, Bahamas.” vudii), but with a dark lateral stripe extending Holotype, Museum of Comparative onto the neck as in C. v. aterrimus. Zoology (MCZ) 37942, an adult female collected by the “Barbours” on 23 March 4. Cubophis vudii raineyi (Barbour and 1934 (not examined by author). Shreve) Ocyophis vudii aterrimus: Zaher et al. 2009:147 (by inference). Alsophis vudii raineyi Barbour and Shreve Cubophis vudii aterrimus: Hedges et al. 1935:363. Type locality, “Landrail Point, 2009:9. Crooked Island, Bahamas.” Holotype, Museum of Comparative Zoology (MCZ) DIAGNOSIS. This subspecies is characterized 37929, a subadult or small adult male by having 163 ventrals and ≥ 78 subcaudals (SVL = 592 mm) collected by T. Barbour (the tail in the holotype was incomplete). The in March 1934 (not examined by author). dorsum (except for the head) is black or very Ocyophis vudii raineyi: Zaher et al. 2009:147 dark brown, somewhat lighter anteriorly, and (by inference). spotted, especially anteriorly, with brown. Cubophis vudii raineyi: Hedges et al. 2009:9. The head is brown, with lighter spots on the internasals, prefrontals, and to a lesser extent DIAGNOSIS. This subspecies is characterized on the supraoculars, and often with a reddish by having 166–176 ventrals and 120 subcaudals area posteriorly. Dark pre- and postorbital (but parts of the tails of the paratypes are stripes are present, with the latter extending missing in many specimens, precluding an onto the neck. The venter is suffused with accurate assessment; the holotype has 120 dark red anteriorly and black posteriorly, subcaudals). Dorsal markings, including variously marked and spotted with white, those on the head, are indistinct. Heads which fades near the vent. Subcaudals are with little yellow, and little or no reddish black, sometimes spotted with white near the coloration is evident on the sides of the neck. tip of the tail. The preorbital stripe is narrower than in the other subspecies and is bordered below by a 3. Cubophis vudii picticeps (Conant) continuous whitish line comprising the lower edges of the preocular, loreal, and nasal (if Alsophis vudii picticeps Conant 1937:82. such a line is present in other subspecies, it is Type locality, “Bimini Islands, Bahamas.” much less clearly defined). Holotype, Museum of Comparative Zoology (MCZ) 43150, an adult female 5. Cubophis vudii utowanae (Barbour and collected by Y. H. Olsen in September Shreve) 1935 (not examined by author). Ocyophis vudii picticeps: Zaher et al. 2009:147 Alsophis vudii utowanæ Barbour and Shreve (by inference). 1935:365. Type locality, “Sheep Cay Cubophis vudii picticeps: Hedges et al. 2009:9. off northwest coast of Great Inagua Island, Bahamas.” Holotype, Museum of DIAGNOSIS. This subspecies is characterized Comparative Zoology (MCZ) 37941, an by having 120 subcaudals and a proportional adult male collected by J. C. Greenway, tail length of 0.327. Dorsal and ventral color Jr. in February 1934 (not examined by and patterns are exceedingly variable, but author). usually well defined and not dark brown to Ocyophis vudii utowanae: Zaher et al. black above or mostly black below (as in C. v. 2009:147 (by inference). 903.7

Cubophis vudii utowanae: Hedges et al. geography of the West Indies, with espe- 2009:9. cial reference to amphibians and reptiles. DIAGNOSIS. This subspecies is characterized Memoirs of the Museum of Comparative by having a high number of ventrals (181 Zoölogy at Harvard College 44:205–359, in the holotype) and subcaudals (129 in the 1 plate. holotype, which is greater than any of the Barbour, T. 1916. Additional notes on West other subspecies despite the holotype missing Indian reptiles and amphibians. Proceed- a small part of the tail). Dorsal markings, ings of the Biological Society of Washing- including those on the head, are indistinct, as ton 29:215–220. in C. v. rainey. However, unlike the latter, tops Barbour, T. 1930. A list of Antillean reptiles of heads are distinctly reddish posteriorly, and amphibians. Zoologica (Scientific with the coloration extending onto the neck, Contributions of the New York Zoologi- and the preorbital stripe is much wider and cal Society) 11:61–116. without a prominent whitish lower border. Barbour, T. 1935. A second list of Antillean reptiles and amphibians. Zoologica (Sci- ACKNOWLEDGMENTS. Aaron M. Bau- entific Contributions of the New York Zo- er, Robert W. Henderson, and John Virata ological Society) 19:77–141. helped locate elusive references. Ned Gilmore Barbour, T. 1937. Third list of Antillean rep- provided Figure 2 and information regarding tiles and amphibians. Bulletin of the Mu- the type specimens in the Academy of Natu- seum of Comparative Zoölogy at Harvard ral Sciences of Philadelphia (ANSP). College 82:77–166. Barbour, T. and A. Loveridge. 1929. Typical LITERATURE CITED reptiles and amphibians in the Museum Attrill, M., C. Edwards, and J. Williams. 1983. of Comparative Zoölogy. Bulletin of the Reptiles and Amphibians of the Bahamas. Museum of Comparative Zoölogy at Har- Bahamas National Trust, Nassau, The Ba- vard College 69:205–360. hamas. 21 pp. Barbour, T. and A. Loveridge. 1946. First sup- Bahamas National Trust Education plement to typical reptiles and amphibi- Office. 2009. Field Guide to the Pine ans. Bulletin of the Museum of Compara- Forests of the Bahamas. Available tive Zoölogy 96:59–214. at http://bahamaswildlife.org/pdfs/ Barbour, T. and B. Shreve. 1935. Concerning pineforestguide.pdf. Archived by some Bahamian reptiles, with notes on WebCite at http://www.webcitation. the fauna. Proceedings of the Boston So- org/6K3KRVRAW on 1 October 2013. ciety of Natural History 40:347–365. Bahamas National Trust. 2012. Snakes of the Bauer, A. M. and A. P. Russell. 1992. The evo- Bahamas. Available at http://www.bnt. lutionary significance of regional integ- bs/Snakes-of-The-Bahamas. Archived umentary loss in island geckos: A com- by WebCite at http://www.webcitation. plement to caudal autotomy. Ethology, org/6NVWKC36k on19 February 2014. Ecology & Evolution 4:343–358. Barbour, T. 1904. Batrachia and Reptilia from Beolens, B., M. Watkins, and M. Grayson. the Bahamas. Bulletin of the Museum of 2011. The Eponym Dictionary of Reptiles. Comparative Zoölogy at Harvard College The Johns Hopkins University Press, Bal- 46: 55–61. timore, Maryland. 296 pp. Barbour, T. 1906. Additional notes on Baha- Bimini Biological Field Station. 2004. Reptiles ma snakes. American Naturalist 40:229– of Bimini. Available at http://www6. 232. miami.edu/sharklab/aboutbimini_ Barbour, T. 1914. A contribution to the zoö- reptiles.html. Archived by WebCite at 903.8

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