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Home , Tropidophiidae, Tropidophis, Tropidophis celiae, Tropidophis haetianus, Tropidophis hendersoni, Tropidophis melanurus

907.1 REPTILIA: : hendersoni

Catalogue of American Amphibians and . and three females) provided by Díaz et al. (2014). The body is robust and the head is Torres, J., R. Powell, and O. H. Garrido. 2017. slightly wider than the neck. Parietal scales in . contact, partially separated, or separated by one scale; 9–11 supralabials (4–5 in contact Tropidophis hendersoni Hedges and with the eye), 10–11 infralabials, 1 preocular, Garrido and 3 postoculars. Dorsal scales are smooth Cuban Khaki Trope (weakly keeled in the middorsal region above the vent) and in 23–27 rows anteriorly, 25–27 : Schwartz and Marsh rows at midbody, and 19–20 rows posteriorly. 1960:73 (part). See Remarks. Middorsal scales are not enlarged except a Tropidophis haetianus haetianus: Schwartz few scales at the posterior end of the body 1975:310 (part). See Remarks. and on the tail. Ventrals number 190–204, Tropidophis hendersoni Hedges and Garrido subcaudals 30–34. Adult males have well- 2002:158. Type locality, “Guarda la Vaca developed pelvic spurs that retract into small [= Guardalavaca], Holguín Province, grooves and slightly longer tails than females , 21°07'35''N, 77°49'55''W, probably (42.5–43.2 mm in males vs. 34.7–37.7 mm in at sea level.” Holotype, Museum of females). Comparative Zoology (MCZ) 47896, an Dorsal color in life (Díaz et al. 2014) is tan adult female, collected by “Clench and to grayish-tan and the venter is paler, cream- Alayo” on 14 August 1945 (examined by colored to slightly orange. Head markings OHG). See also Comments. include brown postocular stripes extending over but not beyond the posterior supralabials, CONTENT. No subspecies are recognized. conspicuous subocular dark bars, additional less obvious dark bars on supralabials and DESCRIPTION. Tropidophis hendersoni is infralabials, a roughly triangular or diamond- a relatively small tropidophiid; the largest shaped figure on the posterior dorsal surface individual is an adult male with a snout- of the head, and a barely evident brown band vent length (SVL) of 332 mm and the largest across the snout and nares. The dorsal pattern female has a SVL of 327 mm (Díaz et al. consists of 48–57 pairs of often partially fused 2014). The following characters are based and not always paired large spots along either on the original description of a single adult side of the dorsal midline, smaller, irregularly female by Hedges and Garrido (2002) and aligned, and less contrasting spots on the data from six additional adults (three males flanks, resulting in 8–10 ‘rows’ of spots around

Figure 1. Adult male Tropidophis hendersoni (left and center) and adult female (right). Photographs by Raimundo López-Silvero Martínez. 907.2

Map. The distribution (in red) ofTropidophis hendersoni; the circle marks the type locality (Hedges and Garrido 2002) and the dot marks the other known locality (Díaz et al. 2014). The distribution is an estimate based on the extent of apparently suitable habitat.

midbody. The tip of the tail has a yellowish Tropidophis haetianus (Stull 1928; Schwartz cast in some individuals. The venter is and Henderson 1988) until Hedges (2002) immaculate or with sparse and widely spaced elevated them to and assigned them spots on the borders of some ventral scales. to the Tropidophis jamaicensis species group. Tropidophis hendersoni differs from all of DIAGNOSIS. Tropidophis hendersoni is easily those taxa in coloration, pattern, scalation, distinguished from its sympatric congeners, and size. For example, the Jamaican taxa have and Tropidophis fewer ventrals and more anterior scale rows, wrighti (Rodríguez Schettino et al. 2013). and Tropidophis haetianus haetianus is larger Tropidophis melanurus is larger (SVL in males than Tropidophis hendersoni and has a lateral to 770 mm, females to 957 mm; Hedges 2002) head stripe, which is absent in Tropidophis and has a predominantly striped pattern with hendersoni (Díaz et al. 2014; Hedges and no more than four rows of spots (if spots Garrido 2002). are present; Hedges and Garrido 2002). Hedges (2002) also assigned Tropidophis has a gracile habitus and celiae, from northwestern Cuba, to a different a very different pattern that consists of black (Tropidophis melanurus) species group, but spots in only four rows. it is superficially similar in appearance to In general appearance, Tropidophis Tropidophis hendersoni. The head is wider hendersoni is similar to Tropidophis in the latter (1.45 head/neck width ratio vs. haetianus from and three 1.31 in ), the pale nuchal Jamaican species (Tropidophis jamaicensis, band (formed from fused occipital spots) is Tropidophis stejnegeri, and Tropidophis obvious in Tropidophis celiae but less evident stullae) that were considered subspecies of in Tropidophis hendersoni. The latter also 907.3

Figure 2. The holotype of Tropidophis hendersoni (MCZ 47896), an adult female collected by “Clench and Alayo” on 14 August 1945 at Guardalavaca, Holguín Province, Cuba. Photograph by S. Blair Hedges (from Hedges 2017). has a salmon-colored venter, whereas that of the holotype was published by Hedges and Tropidophis celiae is cream, and the contrast Garrido (2002); the same photo was reprinted between the dorsal spots and background by Rodríguez Schettino and Garrido (2012). color is less obvious in Tropidophis hendersoni (Díaz et al. 2014; Hedges 2002; Hedges and DISTRIBUTION. Tropidophis hendersoni Garrido 2002; Hedges et al. 1999; Torres et al. is known only from the type locality and a 2013, Torres López et al. 2016). second location about 68 km to the northwest in and around the town of Gibara. The PHYLOGENETIC RELATIONSHIPS. region is a sub-coastal karstic plateau (Díaz Hedges and Garrido (2002) tentatively placed et al. 2014; Hedges and Garrido 2002) with this species in the Tropidophis maculatus abundant caves, many with sinkholes that species group, but Hedges (2002) placed it in allow sunshine to enter and shrubs and vines the species group. to grow in some chambers, including those where were collected (Díaz et al. PUBLISHED DESCRIPTIONS. Detailed 2014). The range was illustrated previously descriptions were provided by Hedges and by Hedges (2017), Rodríguez Schettino and Garrido (2002). Additional descriptions Garrido (2012), Rodríguez Schettino et al. were provided by Hedges (2002) and Díaz et (2013), Uetz et al. (2016), but only Hedges al. (2014), with the latter including data on (2017) included both localities. additional specimens. FOSSIL RECORD. No fossils are known. ILLUSTRATIONS. Color photographs were published by Díaz et al. (2014) and Hedges PERTINENT LITERATURE. The most (2017). A black-and-white photograph of comprehensive sources for the species are 907.4

the original description (Hedges and Garrido by Schwartz and Marsh (1960) and Schwartz 2002) and a note covering both ecology and Garrido (1975). The specimen was ten- (including predation on Eleutherodactylus cf. tatively assigned to Tropidophis haetianus Eleutherodactylus thomasi) and morphology haetianus by Schwartz and Marsh (1960), (Díaz et al. 2014). Tropidophis hendersoni was with this assignment confirmed by Schwartz included in checklists, general works, articles (1975). focusing on other species, or faunal accounts Tropidophis hendersoni was listed as Criti- by Anonymous (2004), Domínguez Díaz cally Endangered (CR) on the IUCN Red List and Moreno García (2003), Estrada (2012), by Powell et al. (2010) and on the Cuban Red González Alonso et al. (2012), Hedges (2017), List by both Rodríguez Schettino and Garrido Henderson and Powell (2007, 2009), Hutchins (2012) and Rodríguez Schettino et al. (2015). et al. (2003), Rodríguez Schettino and Rivalta González (2003), Rodríguez Schettino et ETYMOLOGY. The specific epithet al. (2010, 2013, 2015), Uetz et al. (2016), hendersoni is a patronym honoring Robert W. and Wallach et al. (2014). See Remarks for Henderson “for his many valued contributions information on the conservation status of the to West Indian ” (Hedges and species. Garrido 2002; Uetz et al. 2016).

REMARKS. Initially, the type specimen of ADDITIONAL VERNACULAR NAMES. Tropidophis hendersoni was assigned to Tropi- Henderson’s Trope (Beolens et al. 2011); dophis haetianus (then thought to include Majasito (Rodríguez Schettino and Garrido snakes from Hispanola, , and Cuba) 2012), but note that “Majasito” is the general

Figure 3. An adult male Tropidophis hendersoni (Museo Nacional de Historia Natural de Cuba [MNHNCu] 5055) ingesting a frog (Eleutherodactylus cf. Eleutherodactylus thomasi) in Cueva del Santo, Gibara, Holguín Province. Photographs by Luis M. Díaz (from Díaz et al. 2014). 907.5

Figure 5. Habitat of Tropidophis hendersoni in Gibara, Holguín Province, Cuba. Entrance of Cueva de Los Panaderos (above) and an area under a sinkhole with vegetation, leaf litter, and rocks at Cueva del Santo (below). Photographs by Antonio Cádiz (from Díaz et al. 2014).

hendersoni was first obtained by Dr. Albert Schwartz during his first expedition to Cuba in December 1954. The specimen was de- posited in the small private collection of the malacologist Miguel L. Jaume. When the Nat- ural History Museum of Cuba was founded Figure 4. Adult male Tropidophis hender- and Jaume named its director, he donated his soni (MNHNCu 5056; top) and subadult male entire collection to the museum. Schwartz (MNHNCu 5057; middle and bottom). Photo- had previously examined the specimen at graphs by Luis M. Díaz (from Díaz et al. 2014). Jaume’s home and received it as a loan. How- ever, Schwartz never returned the specimen name applied to all tropes in Cuba. Many to Cuba and after his death, it was deposit- websites (e.g., Midtgaard 2017; Wikipedia ed with part of his herpetological collection 2017) use the name “Cuban Khaki Dwarf at the Museum of Comparative Zoology at Boa"; in general, "Dwarf Boa" is used for Harvard University. Orlando Garrido, during snakes in the Tropidophis (e.g., Uetz et one of his trips to Harvard, was asked by Dr. al. 2016). Ernest E. Williams, then curator of the col- lection, to reexamine the Cuban specimens COMMENTS. The holotype of Tropidophis deposited in the collection. Garrido redis- 907.6 covered the specimen, which he and S. Blair 98–109 in Anfibios y Reptiles de Cuba Hedges subsequently described as a species (L. Rodríguez Schettino, ed.). UPC Print, new to science. Vaasa, Finland. Hoser (2013), based entirely on evidence Estrada, A. R. 2012. The Cuban Archipelago. presented in other studies, reassessed the Pp. 113–125 in Island Lists of West Indian family Tropidophiidae, in the course of which Amphibians and Reptiles (R. Powell and he assigned Tropidophis hendersoni to a new R. W. Henderson, eds.). Bulletin of the genus and a monotypic subgenus, thus creating Florida Museum of Natural History 51. the combination Wellsboa (Tonysilvaboa) González Alonso, H., L. Rodríguez Schetti- hendersoni. Although technically adhering no, A. Rodríguez, C. A. Mancina, and I. to rules set out by the International Code of Ramos García (eds.). 2012. Libro Rojo de Zoological Nomenclature (ICZN), the journal los Vertebrados de Cuba. Editorial Aca- in which these innovations appeared is not demia, La Habana, Cuba. 304 pp. peer-reviewed and the author has a history Hedges, S. B. 2002. Morphological variation of exploiting the work of others to create new and the definition of species in the names that do little more than complicate genus Tropidophis (Serpentes, Tropi- the of those groups he chooses dophiidae). Bulletin of the Natural His- to address (Kaiser 2014; Kaiser et al. 2013). tory Museum [London], Zoology Series Consequently, we choose not to recognize his 68:83–90. taxonomic names. Hedges, S. B. 2017. Caribherp. West Indi- an amphibians and reptiles [Amphibi- ACKNOWLEDGMENTS. We thank Tomás ans and reptiles of Caribbean Islands]: M. Rodríguez Cabrera for help with the Cuba. Temple University, Philadelphia, map, and Antonio Cádiz, Luis M. Díaz, Pennsylvania. Available at http://www. S. Blair Hedges, and Raimundo López- caribherp.org/index.php?il=Cuba&so=- Silvero Martínez for permission to use their class,%20ord,%20subord,%20family,%20 photographs. species&vw=y&dd=n&mob=y. Archived by WebCite at http://www.webcitation. LITERATURE CITED org/6pkdc38st on 15 April 2017. Anonymous. 2004. Liste der Neubeschrei- Hedges, S. B. and O. H. Garrido. 2002. A new bungen von Reptilien des Jahres 2002, snake of the genus Tropidophis (Tropi- mit Nachträgen des Jahres 2001. Elaphe dophiidae) from eastern Cuba. Journal of 12:85–90. Herpetology 36:157–161. Beolens, B., M. Watkins, and M. Grayson. Hedges, S. B., A. R. Estrada, and L. M. Díaz. 2011. The Eponym Dictionary of Reptiles. 1999. A new snake (Tropidophis) from The Johns Hopkins University Press, Bal- western Cuba. Copeia 1999:376–381. timore, Maryland. xiii + 296 pp. Henderson, R. W. and R. Powell. 2007. The Díaz, L. M., A. Cádiz, S. Villar, and F. Bermu- biology of boas and pythons: A retrospec- dez. 2014. Notes on the ecology and mor- tive look to the future. Pp. 2–22 in Biology phology of the Cuban khaki trope, Tropi- of the Boas and Pythons (R. W. Hender- dophis hendersoni Hedges and Garrido son and R. Powell, eds.). Eagle Mountain (Squamata: Tropidophiidae), with a new Publishing, LC, Eagle Mountain, Utah. locality record. IRCF Reptiles & Amphib- Henderson, R. W. and R. Powell. 2009. Nat- ians 21:116–119. ural History of West Indian Reptiles and Domínguez Díaz, M. and L. V. Moreno Amphibians. University Press of Florida, García. 2003. Serpientes del suelo. Pp. Gainesville, Florida. xxiv + 496 pp. 907.7

Hoser, R. T. 2013. A reassessment of the Rodríguez Schettino, L. and V. Rivalta Tropidophiidae, including the creation of González. 2003. Lista de especies. Pp. two new tribes and the division of Tropi- 162–165 in Anfibios y Reptiles de Cuba dophis Bibron, 1840 into six genera, and a (L. Rodríguez Schettino, ed.). UPC Print, revisiting of the Ungaliophiinae to create Vaasa, Finland. two subspecies within pana- Rodríguez Schettino, L.,V. Rivalta González, mensis Schmidt, 1933. Australasian Jour- and E. Pérez Rodríguez. 2010. Distribu- nal of Herpetology 17:22–34. ción regional y altitudinal de los reptiles Hutchins, M., J. B. Murphy, and N. Schlager de Cuba. Poeyana 498:11–20. (eds.). 2003. Grzimek’s Life Ency- Rodríguez Schettino, L., C. A. Mancina, and clopedia. Second Edition. Volume 7, Rep- V. Rivalta González. 2013. Reptiles of tiles. The Gale Group, Inc., Farmington Cuba: Checklist and Geographic Distri- Hills, Michigan. xvi + 593 pp. butions. Smithsonian Herpetological In- Kaiser, H. 2014. Best practices in herpetolog- formation Service 144. 96 pp. ical taxonomy: Errata and addenda. Her- Rodríguez-S., L., V. Rivalta-G., A. petological Review 45:257–268. González-R., and R. Fernández de Ar- Kaiser, H., B. I. Crother, C. M. R. Kelly, L. Lu- cila-F. 2015. Presencia del género Tropi- iselli, M. O'Shea, H. Ota, P. Passos, W. D. dophis (Serpentes: Tropidophiidae) en Schleip, and W. Wüster. 2013. Best prac- el Sistema Nacional de Áreas Protegidas tices: In the 21st century, taxonomic deci- de Cuba. Revista Colombiana de Ciencia sions in herpetology are acceptable only Animal 7:19–24. when supported by a body of evidence Schwartz, A. 1975. Variation in the Antillean and published via peer-review. Herpeto- boid snake Tropidophis haetianus Cope. logical Review 44:8–23. Journal of Herpetology 9:303–311. Midtgaard, R. 2017. English common names Schwartz, A. and O. H. Garrido. 1975. A re- of reptiles. Family Tropidophiidae (Dwarf consideration of some Cuban Tropidophis Boas). Available at http://www.nature- (Serpentes, ). Proceedings of the swindow.dk/UKnames_Tropidophiidae. Biological Society of Washington 88:77– html. Archived by WebCite at http:// 90. www.webcitation.org/6q1lgaSih on 26 Schwartz, A. and R. W. Henderson. 1988. April 2017. West Indian amphibians and reptiles: A Powell, R., G. C. Mayer, and S. B. Hedges. check-list. Milwaukee Public Museum 2010. Tropidophis hendersoni. The IUCN Contributions in Biology and Geology Red List of Threatened Species 2010: e. 74:1–264. T178429A7545322. Available at http:// Schwartz, A. and R. J. Marsh. 1960. A review dx.doi.org/10.2305/IUCN.UK.2010-4. of the pardalis-maculatus complex of the RLTS.T178429A7545322.en. Archived boid genus Tropidophis of the West In- by WebCite at http://www.webcitation. dies. Bulletin of the Museum of Compar- org/6gCr1ft5q on 22 March 2016. ative Zoology 123:49–84 + Figures 1–10. Rodríguez Schettino, L. and O. H. Garrido. Stull, O. G. 1928. A revision of the genus 2012. Hoja de datos del taxón. Tropidophis Tropidophis. Occasional Papers of the hendersoni Hedges y Garrido, 2002. P. 176 Museum of Zoology, University of Michi- in Libro Rojo de los Vertebrados de Cuba gan 195:1–149, Plates 1–3. (H. González Alonso, L. Rodríguez Schet- Torres, J., O. J. Torres, and R. Marrero. 2013. tino, A. Rodríguez, C. A. Mancina, and I. Nueva localidad para la boa enana Tropi- Ramos García, eds.). Editorial Academia, dophis celiae Hedges, Estrada y Díaz, 1999 La Habana, Cuba. (Serpentes, Tropidophiidae) en Cuba. 907.8

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