1 REPTILIA: SQUAMATA: LEIOCEPHALIDAE Leiocephalus personatus

Catalogue of American Amphibians and in Mus. Compar. Zool.” (Museum of Reptiles 922 Comparative Zoology; MCZ), date unknown. Cannot be located (e.g., Powell, R. 2019. Leiocephalus personatus. Barbour 1914; Pregill 1992; Schwartz and Thomas 1975). Synonymy fide Boulenger Leiocephalus personatus Cope (1885), although attributed to Cochran Hispaniolan Masked Curly-tailed Lizard (1941, 2005) by Pregill (1992); see also Remarks. Liocephalus personatus Cope 1862:182. Type Leiocephalus personatus: Barbour 1914:302. locality, “Hayti (near Jeremie)” (= near Jérémie, Département de la Grand’Anse, CONTENT. Twelve subspecies are currently Haiti). Syntypes, Museum of Comparative recognized: Leiocephalus personatus actites, Zoology (MCZ) R–3615 (see Remarks), Leiocephalus personatus agraulus, Leiocephalus two adults (one male and one female) personatus budeni, Leiocephalus personatus “sent in a valuable collection made by elattoprosopon, Leiocephalus personatus mentalis, Dr. D. F. Weinland to Prof. Agassiz,” date Leiocephalus personatus personatus, Leiocephalus unknown (not examined by author). personatus poikilometes, Leiocephalus personatus Liocephalus trigeminatus Cope 1862:183. Type pyrrholaemus, Leiocephalus personatus scalaris, locality, “Hayti (near Jeremie)” (= near Leiocephalus personatus socoensis, Leiocephalus Jérémie, Département de la Grand’Anse, personatus tarachodes, and Leiocephalus personatus Haiti). Syntypes, “Dr. Weinland’s Coll. trujilloensis (see Remarks).

Figure 1. Adult male Leiocephalus personatus mentalis from Cabo San Rafael, La Altagracia Province, Dominican Republic. Photograph by Miguel A. Landestoy. 2

Map. Map of Hispaniola showing the distribution of Leiocephalus personatus; open circles indicate type localities, black dots mark other known records (some proximate localities are indicated by single dots), the red x marks a fossil locality, and question marks indicate records of specimens not assigned to subspecies. Numbers refer to subspecies and correspond to the numbers of the subspecific accounts: 1 – Leiocephalus personatus personatus; 2 – Leiocephalus personatus actites; 3 – Leiocephalus personatus agraulus; 4 – Leiocephalus personatus budeni; 5 – Leiocephalus personatus elattoprosopon; 6 – Leiocephalus personatus mentalis; 7 – Leiocephalus personatus poikilometes; 8 – Leiocephalus personatus pyrrholaemus; 9 – Leiocephalus personatus scalaris; 10 – Leiocephalus personatus socoensis; 11 – Leiocephalus personatus tarachodes; 12 – Leiocephalus personatus trujilloensis. Modified from Schwartz and Henderson (1991).

DESCRIPTION. Leiocephalus personatus scales include 6 (usually) supraoculars, 2–17 is a moderately sized member of the genus loreals, and 7–14 temporals; median head (maximum snout–vent length [SVL] in scales number 4–6 (mode 4), frontoparietals males is 86 mm, maximum SVL in females 3–5 (mode varies by subspecies), supraorbital is 63 mm; Schwartz 1967). An estimated semicircles usually complete, and parietals maximum SVL of 90 mm was mistakenly are almost always in contact. Males have a reported for Leiocephalus personatus by pair of enlarged postcloacal scales. Olson (1990); however, based on cited Sexual dichromatism is evident (Schwartz locality data, the species measured was and Henderson 1991). Throat color in males Leiocephalus melanochlorus. Estimated varies from solid black to immaculate yellow female SVL at maturity is 42.5 mm (Gifford with a contrasting black mental scale; males and Powell 2007). Lateral folds are absent in some subspecies might bear a few scattered (Schwartz and Henderson 1991). Dorsal and diffuse dusky spots but are never heavily, scales are imbricate, weakly tricuspid to regularly, and discretely spotted. Throats of denticulate, and keeled. Ventral scales are females have distinct black to dark gray spots. imbricate, weakly denticulate, and smooth. Males have no ventral pattern, whereas the Median dorsal crest scales are enlarged, ventral surfaces of females are heavily dotted greatly attenuate, strongly overlapping, much with black to dark gray spots. Males have a lower than median dorsal caudal scales, and dark brown to black facial mask, but no black number 41–64 in occiput–vent distance; one– neck or shoulder patches; the dorsum can half midbody scales number 18–28. Head be dark brown, reddish-brown dotted with 3 yellow, light brown, tan, tan flecked with gold, grayish-tan, sandy, practically black, tan to dark brown and often flecked or mottled with yellow or brick-red, and sometimes with tan, creamy, or sandy dorsolateral lines; transverse neck bars are present or absent, at times reduced to a series of black median dots on enlarged middorsal scales; flanks usually tan to brown with admixed red, green, and cream scales; the hindlimbs usually some shade of green; the forelimbs tan; the venter some shade of green; the tail brown above (often chevronate) and orange below. The facial mask of females often is reduced to a dark-edged temporal triangle; the dorsum is brown, with or without complete chevrons; flanks are tan with dark brown vertical Figure 2. Adult female Leiocephalus personatus bars; pale dorsolateral and lateral lines are mentalis from near Juanillo, La Altagracia Prov- prominent; the venter can be dirty yellow, ince, Dominican Republic. Photograph by Robert white, pale green, or opalescent; the limbs Powell. are brown; the underside of the tail is yellow- orange to bright orange or pink. Leiocephalus lunatus lacks a facial mask, male throats have discrete dots, female throats have DIAGNOSIS. Leiocephalus personatus faint gray dots, and a neck blotch is present can be distinguished from all Hispaniolan (Leiocephalus personatus has a facial mask, congeners using the following characteristics male throats solid black to patternless or (Henderson and Schwartz 1984; Henderson with few diffuse spots, female throats heavily et al. 1984; Köhler et al. 2016): Leiocephalus patterned with dark gray spots, and a neck melanochlorus, Leiocephalus schreibersii, and blotch is absent). Leiocephalus sixtoi are relatively large (male SVL of first two species to >100 mm and that PHYLOGENETIC RELATIONSHIPS. Leio- of the third to 88 mm) and have distinct lateral cephalus barahonensis, Leiocephalus lunatus, folds and lateral scales conspicuously smaller Leiocephalus personatus, Leiocephalus semil- than dorsal scales (Leiocephalus personatus ineatus, and Leiocephalus vinculum were hy- is smaller, lacks prominent lateral folds, pothesized to form the Leiocephalus persona- and lateral scales only slightly smaller than tus group (Schwartz 1967), but a study based dorsal scales); Leiocephalus pratensis lacks a on 39 morphological characters provided lit- regular row of middorsal scales (present in tle support for this hypothesis (Pregill 1992). Leiocephalus personatus); preauricular scales Two alternative tree topologies and a consen- of Leiocephalus barahonensis and Leiocephalus sus tree were presented by Pregill (1992), all semilineatus are much larger than adjacent of which showed Leiocephalus personatus in temporal scales (preauricular scales of a clade that included Leiocephalus lunatus of Leiocephalus personatus are not enlarged Hispaniola, Leiocephalus loxogrammus of the when compared to adjacent temporal scales); Bahamas, and Leiocephalus raviceps of Cuba, dorsal crest scales of Leiocephalus rhutidira all within a larger clade comprised mainly of and Leiocephalus vinculum are not attenuate or Hispaniolan species (Leiocephalus barahon- strongly overlapping (attenuate and strongly ensis, Leiocephalus lunatus, Leiocephalus per- overlapping in Leiocephalus personatus); and sonatus, Leiocephalus pratensis, Leiocephalus 4

Figure 3. Adult female Leiocephalus personatus pyrrholaemus from Frontón, Samaná Province, Domin- ican Republic. Photograph by Miguel A. Landestoy. rhutidira, Leiocephalus semilineatus) but also CONSERVATION STATUS. Because this including three Cuban species (Leiocephalus species is widely distributed, abundant, and cubensis, Leiocephalus raviceps, Leiocephalus not declining fast enough to qualify for a more stictigaster) in addition to a single Bahamian threatened category, Leiocephalus personatus species (Leiocephalus loxogrammus). is listed as being of Least Concern (LC) on Based on albumin immunological meth- the IUCN Red List of Threatened Species ods, lizards in the genus Leiocephalus repre- (Incháustegui and Landestoy 2016). sent a relatively recent radiation, with the old- est divergences within the genus occurring PUBLISHED DESCRIPTIONS. In addition less than 10 mya (Hass et al. 2001). Immu- to the original descriptions (Cochran 1932; nological distance (ID) values of Hispaniolan Cope 1862; Gali and Schwartz 1982; Gali et species (including Leiocephalus personatus) al. 1988; Mertens 1939a; Schwartz 1967, 1969, ranged from 0 to 4 (Leiocephalus personatus 1971), detailed descriptions were provided had an ID of 1 to Leiocephalus schreibersii, by Cochran (1941, 2005) and Schwartz and which was within the ± 2 ID-unit error range Henderson (1991). A list of morphological of the technique). (including skeletal), meristic, and pattern Leiocephalus personatus was the sister characters were published by Pregill (1992). species of Leiocephalus lunatus within a His- paniolan clade that also included Leiocephalus ILLUSTRATIONS. Color photographs of barahonensis, Leiocephalus schreibersii, and adult males were published by Bartlett and Leiocephalus semilineatus, but distinct from Bartlett (2009), Drewes (2012 [in captivity]), a Cuban clade that included Leiocephalus ca- Dollinger (2019), EOL (2019), Graff and rinatus, Leiocephalus macropus, Leiocephalus Bringsøe (1996), Hedges (2018), Henderson raviceps, and Leiocephalus stictogaster (Gif- (2001), Henkel and Schmidt (2007, 2010), ford 2008). A subsequent report demonstrat- Landestoy and Ortíz (2015), Matsumoto ed Leiocephalus personatus was most closely (1996), Powell et al. (1996), Purser (2008), related to Leiocephalus psammodromus, but Uchiyama (1995), and Wikipedia contributors only six species were included in that study (2019). Color photographs of males, females, (Pyron et al. 2013). and juveniles were published by Rogner (1997b). Color photographs of males and 5

published by Henkel and Schmidt (2007) is Leiocephalus personatus. Black-and-white photographs of Leio- cephalus personatus trujilloensis (including the holotype), Leiocephalus personatus mentalis, and Leiocephalus personatus pulcherrimus (=Leiocephalus personatus scalaris) were provided by Mertens (1939a). The same photographs of the habitat at the type locality of Leiocephalus personatus trujilloensis and of male and female Leiocephalus personatus mentalis Figure 4. Top of the head of an adult male were published by Mertens (1940a, 1940b). Leiocephalus personatus (AMNH 16155). From A photograph of an adult male Leiocephalus Schmidt (1921a). personatus scalaris from Cayo Monte Chico (Cayos Siete Hermanos) was presented females were provided by Ackermann (2006), by Burns et al. (1992). Photographs of an Dahms Tierleben (2018 [in captivity]), adult male and an adult female Leiocephalus Herrmann (1994 [in captivity]), Hoffman personatus mentalis were published by (2005), Landestoy and Reyes (2018), Rey Powell (2002); the photograph of the female (2017 [unlabeled Leiocephalus personatus also was published by Powell (2003). A mentalis]); color photographs of juveniles were photograph of an adult female Leiocephalus published by Herrmann (1994). An unlabeled personatus mentalis from Haiti was presented color photograph of Leiocephalus personatus by Klingelhöffer (1957), though this taxon is accompanied a description of Leiocephalus not known from that country. Photographs carinatus by Flank (1998). Photographs of of natural habitats were published by Gifford male and female Leiocephalus personatus et al. (2008). Photographs of an adult pair trujilloensis were provided by Sánchez Muñoz (undetermined subspecies) were presented (2019). Color photographs of adult male by Spycher (1984) and one photograph of an Leiocephalus personatus scalaris from Florida adult male (undetermined subspecies) was were published by Bartlett and Bartlett (2006 published by Reiprich (2009). [near the Miami International Airport], 2011 Black-and-white illustrations include a [adult male]) and Krysko et al. (2011; Key drawing of the top of the head of American Biscayne); an additional color photograph of Museum of Natural History (AMNH) 16155 an adult male Leiocephalus personatus scalaris provided by Schmidt (1921a), a drawing without locality information was published published by Kästle (1972b, 1986), and line by Bech and Kaden (1990). All images of drawings of dorsal head scalation and black- MCZ and Smithsonian National Museum of and-white illustrations of color patterns Natural History (USNM) type specimens of of Leiocephalus personatus personatus, Leiocephalus personatus were published by Leiocephalus personatus scalaris, and EOL (2019). Leiocephalus personatus mentalis published by The color photographs labeled Cochran (1941, 2005). The posterior end of “Leiocephalus personatus” by Obst et al. the dentary was illustrated by Pregill (1992). (1984, 1988) and Rogner (1992a, 1997a) are Drawings of the auricular opening and Leiocephalus schreibersii; one of two color preauricular scale, lateral and dorsal views of photographs labeled “Bunter Masakenleguan median dorsal crest scales, lateral views of the (Leiocephalus personatus)” [sic] by Dollinger head and neck of male Leiocephalus personatus (2019) is Leiocephalus schreibersii. The color personatus and Leiocephalus personatus photograph labeled Leiocephalus schreibersii mentalis, and ventral views of the throats 6

is absent only from extremely high elevations (elevational range: sea level to ~625 m), the extremely xeric Valle de Neiba and Cul- de-Sac Plain, and the Barahona Peninsula (Henderson and Powell 2009; Schwartz 1967; Schwartz and Henderson 1991). In general, the species occurs in mesic habitats; if in drier areas, it usually is restricted to the most densely shaded situations (Schwartz and Henderson 1991). These lizards readily exploit both natural and human-modified (agricultural and urban) Figure 5. Lateral views of the head and neck of habitats, although the species apparently an adult male Leiocephalus personatus personatus avoids exposed situations like beaches (Burns (MCZ 74636) from Roseaux, Département du et al. 1992; Henderson and Powell 2009). Sud, Haiti (top) and of an adult male Leiocephalus Introduced populations assigned to personatus mentalis (MCZ 75131) from Juanillo, Leiocephalus personatus scalaris have been La Romana Province, Dominican Republic. From documented in Miami-Dade County, Florida Schwartz (1967). Illustration reproduced with (Bartlett 1994; Bartlett and Bartlett 1999, permission granted by Editor of Tulane Studies in 2006; Butterfield et al. 1997; Kraus 2009; Zoology and Botany. Krysko et al. 2010, 2011, 2019; Lever 2003; Powell et al. 1998, 2012, 2019). Populations established during the 1970s apparently of male Leiocephalus personatus personatus, were extirpated in the 1980s, but became Leiocephalus personatus trujilloensis, Leiocephalus reestablished in the mid-1990s when imports personatus tarachodes, Leiocephalus personatus for the pet trade were resumed (Bartlett and actites, and Leiocephalus personatus budeni were Bartlett 1999). Although individual lizards presented by Schwartz (1967). Drawings of were documented (Krysko et al. 2010, 2011 the dorsum, lateral view of the head, teeth, [photograph included in Krysko et al. 2011]) and diagrammatic dorsal, ventral, and lateral on Key Biscayne in 2004, additional lizards (head only) patterns published by Olson were not found subsequently at that site; also, (1990) actually were images of Leiocephalus no lizards were found during surveys at the melanochlorus. Miami International Airport (Meshaka et al. 2004), supporting the contention whether DISTRIBUTION. Leiocephalus personatus “these current populations will actually has an extensive but largely disjunct become truly established is questionable” distribution across much of Hispaniola (Bartlett and Bartlett 1999:187) Although (including several satellite islands), ranging listed on some websites that focus on along the northern and southern shores of introduced species (e.g., Ferriter et al. 2008; the Tiburon Peninsula (including Grosse FWC 2018, 2019), the species was not listed Caye and Île-à-Vache) in southwestern Haiti, in recent publications (e.g., Meshaka 2011; northern Haiti eastward along the northern Powell et al. 2016) or listed as questionable or coast (including the Cayo Monte Chico [Cayos problematic (Powell et al. 2012, 2019; Krysko Siete Hermanos] and Isla Cabras) into the et al. 2019), and the Florida populations extreme southeastern Dominican Republic, that were at best tenuously established are along parts of the southern Dominican coast, presumed extirpated. and with extensions or isolated pockets The native range was illustrated previously toward the center of the island. The species by Cochran (1941, 2005), Dollinger (2019), 7

Hedges (2018), Schwartz (1967), and al. 1998, 2012, 2019; van Wilgen et al. 2009), Schwartz and Henderson (1991). captive husbandry (Anonymous 1991; Bech and Kaden 1990; Bindler and Duchene 2009; FOSSIL RECORD. Late Pleistocene fossils Dollinger 2019; Drewes 2007, 2012; Flower from the Cerro de San Francisco (Elías Piña 1929; Frommer 1984; Graff and Bringsøe Province), Dominican Republic, were as- 1996; Herrmann 1994; Hoffman 2005; Kästle signed to Leiocephalus personatus (Etheridge 1972b, 1986; Klingelhöffer 1957; Purser 2008; 1965). Rogner 1992a, 1995, 1997a, 2005; Spycher 1984), diseases in captivity (Bauwens et PERTINENT LITERATURE. Topics al. 2014; Gaigg 2008; Peters 1978 [quoted addressed in the literature include (note that in Petzold 1982, 1984, 2008], Rossier et al. references listing species of Leiocephalus 2016), endoparasites (Duszynski et al. 2008; formerly considered subspecies of Huntington and Cisper 1994), extinction Leiocephalus personatus [see Remarks] are risk (Incháustegui and Landestoy 2016), not cited): original descriptions (Cochran fossil record (Etheridge 1964, 1965, 1966c), 1932; Cope 1862; Gali and Schwartz 1982; Gali longevity in captivity (Slavens 1981, 1982, et al. 1988; Mertens 1939a; Schwartz 1967, 1987, 1988; Slavens and Slavens 1991; 1969, 1971), apparent failed introductions Snider and Bowler 1992), morphology, in Florida (Bartlett 1994; Bartlett and performance, anti-predator behavior Bartlett 1999, 2006, 2011; Butterfield et al. (Gifford et al. 2008; Schwartz 1979; Wassersug 1997; Ferriter et al. 2008; FWC 2018, 2019; et al. 2005a, 2005b), natural history (Gifford Kraus 2009; Krysko et al. 2010, 2011, 2019; and Powell 2007; Henderson and Powell Lever 2003; Meshaka et al. 2004; Powell et 2009; Henderson et al. 1987a, 1987b, 1988;

Figure 6. Ventral views of the throats of adult male Leiocephalus personatus (from left to right):Leio - cephalus personatus personatus (MCZ 74636) from Roseaux, Département du Sud, Haiti; Leiocephalus personatus trujilloensis (ASFS X7734; now KU 244449) from Santo Domingo, Distrito Nacional, Domin- ican Republic; Leiocephalus personatus tarachodes (MCZ 81087, holotype) from 6 km SE Nagua, María Trinidad Province, Dominican Republic; Leiocephalus personatus actites (MCZ 81088, holotype) from Sosúa, Puerto Plata Province, Dominican Republic; and Leiocephalus personatus budeni (MCZ 81089, holotype) from 12 km NE Jarabacoa, 2000 feet [= 610 m], La Vega Province, Dominican Republic. From Schwartz (1967). Illustration reproduced with permission granted by Editor of Tulane Studies in Zoology and Botany. 8

Figure 7. An adult male Leiocephalus personatus actites from Estero Hondo, Puerto Plata Province, Dominican Republic. Photograph by Miguel A. Landestoy.

Kästle 1971, 1972a, 1984; Mertens 1940b; Fotolulu (2018), Frank and Ramus (1995), Schwartz and Henderson 1991), pet trade Franz and Cordier (1986), Friedl et al. (1988), (Anonymous 1980; Araújo 2014; Bartlett and Garman (1887), Hedges (2018), Hedges Bartlett 1997; Huikeshoven 2010; Shiau et al. et al. (2019), Heino (2004), Henderson 2006), phylogeography (Gifford 2005, 2008), and Schwartz (1984), Henderson et al. systematics (Pregill 1992), and broad, mostly (1984), Henkel and Schmidt (2007, 2010), phylogenetic studies in which Leiocephalus Hernández (2013), Herter (1960), Hutchins personatus was used as an outgroup (Harvey et al. (2003), iNaturalist (2019), Jes (2006), and Gutberlet 2000; MacLeod et al. 2016; Kästle (1972b, 1986), Kluge (1984), Köhler et Matos et al. 2016; Mello 2014; Okajima and al. (2016), Landestoy and Ortíz (2015), Liner Kumazawa 2009, 2010; Olmo 1981, 1984; (1997, 1999), MacLean et al. (1977), Mertens Pyron et al. 2013; Schulte 2013; Schulte et al. (1938, 1939b, 1940a, 1946, 1967, 1969), 2003; Streicher et al. 2016; Wiens 1993; Yu et Midtgaard (2019), NCBI (2018), Nelson and al. 2017). Powell (2002), Noble and Bradley (1933), Leiocephalus personatus also has been Nolan (1913), Obst et al. (1984, 1988), Olson included in checklists, keys, general works, (1990, 1995 [although cited locality data articles focusing on other species, or indicate that he had examined Leiocephalus faunal accounts (some including brief melanochlorus instead of Leiocephalus descriptions): Alfonso (2016), Alfonso et al. personatus]), Petzold (1982, 1984, 2008), (2012), Anonymous (1991), Arment (2005), Petzold et al. (1970), Powell (1993, 1999a, Barbour (1914, 1930, 1935, 1937), Barbour 1999b, 2002, 2003, 2012), Powell and Gifford and Loveridge (1929, 1946), Bauer et al. (2010), Powell and Henderson (2008), Powell (2012), Bergmann (2008), Böhme (1983), et al. (1996, 1998, 1999, 2012, 2019), Purser Böker (1939), Boulenger (1885), Burns (2008), Reiprich (2009), Rey (2017), Rogner et al. (1992), Christman (1974), Cochran (1997b), Sánchez Muñoz (2019), Schmidt (1924, 1928, 1934a, 1934b, 1941, 1953, (1921a, 1921b), Schwartz (1965, 1968, 1980), 1961, 2005), Cope (1868, 1887), Crombie Schwartz and Henderson (1985, 1988), (1970), Dlugolecki et al. (2015), EOL (2019), Schwartz and Thomas (1975), Schwartz et al. Etheridge (1966a, 1966b, 2000), Felix (1988), (1978), SEA/DVS (1990, 1992), Seligmann 9 and Labra (2013), K. Smith (2006, 2009), M. was assigned the newer catalog number in Smith (1929), Uchiyama (1995), Uetz et al. 1985 (J. Rosado, in litt., 23 January 2017). (2017), Wikipedia contributors (2019), and The annotations by Schwartz and Henderson Wrobel (2004). (1991) and Uetz et al. (2019) that the syntypes Many additional websites (e.g., Dahms of Leiocephalus personatus had been lost Tierleben 2018; Rey 2017; Sánchez Muñoz were obviously in error and might have been 2019) provide information on Leiocephalus mistaken references to the unlocated syntypes personatus; most deal with husbandry, but of Leiocephalus trigeminatus. some provide photographs or descriptions At various times, a number of congeneric and many address some aspects of natural Hispaniolan taxa have been considered history. Unfortunately, some of those sites subspecies of Leiocephalus personatus (e.g., (e.g., Dollinger 2019; Exotic-Pets.co.uk 2018) Schwartz 1965) and are so listed in many mistakenly illustrate Leiocephalus schreibersii. publications (e.g., Cochran 1941, 1961, 2005; Etheridge 1966c; Mertens 1938, REMARKS. Cope (1862) described 1939a, 1939b): Leiocephalus altavelensis, Liocephalus personatus and Liocephalus Leiocephalus barahonensis, Leiocephalus ba- trigeminatus on consecutive pages, listing rahonensis aureus, Leiocephalus barahonensis the same geographic origin for both. Cope beatanus, Leiocephalus eremitus (Mertens (1868:122) provided a key to species of 1939a, by inference), Leiocephalus lunatus, Liocephalus with one option leading to both Leiocephalus lunatus arenicolor, Leiocephalus species; an appended annotation stated: “The lunatus louisae (also noted by Henderson last two are much alike in structural features, and Powell 2004), Leiocephalus semilineatus, but differ greatly in coloration; they do not and Leiocephalus vinculum. Leiocephalus ba- appear to be sexes of the same animal, as I have rahonensis, Leiocephalus lunatus, Leiocephalus seen both ♂ and ♀ of the latter.” However, as personatus, Leiocephalus semilineatus, and noted by Barbour (1914), Cope (1887:438) Leiocephalus vinculum constituted the stated: “TheL. trigeminatus is probably the hypothesized Leiocephalus personatus group immature stage of L. personatus, with which (Schwartz 1967; but see Phylogenetic Dr. Boulenger properly unites it.” Relationships). Two specimens (MCZ R-3615 and Cochran (1941, 2005) listed “Leiocepha- MCZ R-168525) are listed as syntypes of lus personatus Garman … (part)” in the syn- Leiocephalus personatus, the latter of which onymy of Leiocephalus personatus mentalis;

Figure 8. Adult male Leiocephalus personatus trujilloensis from Santo Domingo, Distrito Nacional, Do- minican Republic. Photograph by Miguel A. Landestoy (from Landestoy and Reyes 2018). 10

Figure 9. An adult male Leiocephalus personatus scalaris from Key Biscayne, Miami-Dade County, Flor- ida (11 October 2004). Photograph by Joseph P. Burgess. however, the cited locality (Puerto Plata) is ETYMOLOGY. Although not stated by Cope in the range of Leiocephalus personatus ac- (1862), the specific epithet evidently is de- tites Schwartz. Schwartz and Thomas (1975) rived from the Latin personatus (= masked; and Schwartz and Henderson (1988) indicat- Brown 1956), an obvious allusion to the dark ed that specimens from San Cristóbal (San facial mask of this species. The subspecific Cristóbal Province) seem not to be referable epithet actites is from the Greek aktites (= to Leiocephalus personatus trujilloensis, and shore dweller; Brown 1956), undoubtedly a also noted that a specimen from St.-Marc, reference to the fact that the distribution of Département de l’Artibonite, Haiti was “left the subspecies “is restricted to the coastline of unassigned subspecifically.”Leiocephalus a single province in northern República Do- personatus was included by Powell (1993) minicana” (Schwartz 1967:16). The subspecif- among species with a broad insular range and ic epithet agraulus is from the Greek agraulos numerous distributional records of uncertain (= dwelling in the field; Brown 1956), almost systematic status; he concluded that data were certainly a reference to the fact that “natives” insufficient to draw conclusions regarding indicated that “the lizards had been collected the possible specific status of taxa current- under rocks in a large open meadow, stud- ly identified as subspecies. Pregill (1992:49) ded with a few pines, and partly under cul- remarked: “Leiocephalus personatus is an es- tivation” (Schwartz 1967:23). The subspecif- pecially variable species, as demonstrated by ic epithet budeni is a patronym honoring D. the numerous populations given subspecific W. Buden, co-collector of the holotype. The designation, and in the character discordance subspecific epithet elattoprosopon is from noted above [in his account of the species]. A the Greek elatton (= less) and prosopon (= more detailed examination of this widespread face, mask), a reference to the reduced dark Hispaniolan taxon is warranted.” mask in this subspecies when compared to 11

Leiocephalus personatus personatus (Gali et tailed Lizard (Arment 2005; Bartlett and Bart- al. 1988). Although not stated by Cochran lett 1999, 2006; Lever 2003), Green-legged (1932:178) and although mentalis is from the Curlytail Lizard (FWC 2018), Jewelled Curly- Latin (= of the mind; Brown 1956), the sub- tailed Lizard (Bauwens et al. 2014), Masked specific epithet mentalis appears to be a ref- Curlytail (Obst et al. 1988), and various other erence to the unusual pattern of the chin and variations variously hyphenated or curtailed throat of males (“mental shield and lower lips (i.e., jeweled vs. jewelled; curlytail or curlytail of adult males sepia, remainder of throat pale lizard vs. curly-tailed lizard). china blue, immaculate or with a few minor In northern parts of the Dominican Re- pale-brown spots confined mostly to single public, the species is called “Mariguanita” scales”). The subspecific epithetpoikilometes (the meaning is uncertain) or “El Lagarto de “is Greek for ‘full of various wiles,’ an allu- Cola Rizada” (= curly-tailed lizard), where- sion to the difficulty encountered in securing as in the area around Baní, the name “Come these lizards” (Schwartz 1969:85). Although Maíz” (= corn-eater) is used (Rey 2017; M. not stated by Schwartz (1971), the subspecific A. Landestoy, personal communication), but epithet pyrrholaemus is from the Greek pyr- these names also are applied to other spe- rho (= flame-colored) andlaimos (= throat; cies of Leiocephalus. “Rana” (or, more rarely Brown 1956), certainly an allusion to the “Rano”), the Spanish word for frog, is widely deep orange throat of these lizards. Although used in the southern Dominican Republic for not stated by Cochran (1932), the subspe- any small lizard (M. A. Landestoy, personal cific epithet scalaris is from the Latin (= of a communication). In Haiti, lizards (including ladder; Brown 1956), possibly a reference to Leiocephalus personatus) usually are called the prominent nuchal crest scales of males. “Mabuya” or “Sandolit,” both of which are The subspecific epithet socoensis “is derived used for any small lizard (M. A. Landestoy, from the name of one of the localities (Boca personal communication). del Soco)” where this subspecies is known In German (e.g., Fotolulu 2018; Tierdoku to occur (Gali and Schwartz 1982:179). The contributors, 2012; Uetz et al., 2017), the spe- subspecific epithettarachodes presumably is cies is known as the Bunter Maskenleguan (= from the Greek taraktor (= disturber; Brown colorful masked iguana). This name is wide- 1956), but the meaning is uncertain and ly used in the European pet trade. German Schwartz (1967) did not provide an explana- common names for 11 of the 12 currently tion. Although not stated by Mertens (1939a), recognized subspecies were given by Fotolulu the subspecific epithettrujilloensis (originally (2018), with no subspecific epithet given for spelled “trujilloënsis”) certainly is a reference the nominate subspecies: Leiocephalus per- to the type locality, Cuidad Trujillo (= Santo sonatus actites, Nördlicher Maskenleguan; Domingo). Leiocephalus personatus agraulus, Cordille- ra-Maskenleguan; Leiocephalus personatus ADDITIONAL VERNACULAR NAMES. budeni, Buden’s Maskenleguan; Leiocephalus English common names include (note that personatus elattoprosopon, Tiburon-Masken- lists of references are not intended to be com- leguan; Leiocephalus personatus mentalis, plete): Haitian Curlytail Lizard (e.g., Frank Nordöstlicher Maskenleguan; Leiocephalus and Ramus 1995; FWC 2019; Incháustegui personatus poikilometes, Neiba-Maskenle- and Landestoy 2016; Uetz et al. 2019; Wrobel guan; Leiocephalus personatus pyrrholaemus, 2004), Hispaniolan Masked Curlytail (Hedg- Samanâ-Maskenleguan; Leiocephalus perso- es 2018, Hedges et al. 2019), West Indian natus scalaris, Siete Hermanos-Maskenle- Masked Lizard (Smith 1929), Green-legged guan; Leiocephalus personatus socoensis, Rio Curly-tail and Jewelled Curly-tail (Wikipe- Soco-Maskenleguan; Leiocephalus persona- dia contributors 2019), Green-legged Curly- tus tarachodes, Hato Mayor-Maskenleguan; 12 and Leiocephalus personatus trujilloensis, DIAGNOSIS. This subspecies is characterized Cristôbal-Maskenleguan. Note the use of a by males with a combination of pale green circumflex (most frequently used in French) throat, usually with a pair of transverse black instead of an acute accent for the Spanish lines at the level of the second and third chin place-names (i.e., Samaná and Cristóbal). shields; pale greenish ventral color; mask In Danish (Midtgaard 2019), the species present but somewhat faded in older adults is known as Maskeleguan (= Masked Iguana), with a pair of creamy to yellow-orange broad in French (Dollinger 2019) as L’iguane à vertical subocular bars which may in turn be queue coubée (= Tailed Iguana), and in Cyrillc partly or wholly confluent with a horizontal (Sokolov 1988) as обыкновенная масковая pale temporal bar and a postauricular bar; игуана (= Ordinary Masked Iguana). dorsal ground color from grayish-tan and sandy to practically black; dorsolateral longitudinal lines fairly prominent and 1. Leiocephalus personatus personatus Cope bordered medially by a broad ill-defined darker area of the dorsal zone; and nuchal Liocephalus personatus Cope 1862:182. See and scapular transverse bars absent in adults species synonymy. only at times indicated by a series of darker Liocephalus trigeminatus Cope 1862:183. See (dull brown, rather than dark brown or black) species synonymy. median crest scales (Schwartz 1967). Size is Leiocephalus semilineatus: Cochran 1928:54 large (maximum SVL in males is 86 mm, (“part; not of Dunn”). maximum SVL in females is 61 mm); mean Leiocephalus personatus personatus: Barbour number of loreals (4.3) moderate; median 1935:120. head shields modally 4; frontoparietals modally 4; and supraorbital semicircles more DIAGNOSIS. This subspecies is characterized often complete (58.9%) than not (Schwartz in males by the combination of a solid 1967). black throat confluent with a black loreal- temporal-lateronuchal area; three to five 3. Leiocephalus personatus agraulus Schwartz dark transverse dorsal bars on the neck and shoulders; and a broad dorsal zone bounded Leiocephalus personatus agraulus Schwartz by wide pale dorsolateral longitudinal lines 1967:21. Type locality, “1 mi. WSW (Schwartz 1967). Size is large (maximum Constanza, 4000 feet (1311 meters), La SVL in males is 79 mm, maximum SVL in Vega Province, República Dominicana.” females is 62 mm); mean number of loreals Holotype, MCZ R-81090, an adult male (5.6) high; mean number of temporals collected by a “native collector” on 4 July (11.1) high; median head shields modally 5; 1963 (not examined by author). frontoparietals modally 4 (Schwartz 1967). DIAGNOSIS. This subspecies is 2. Leiocephalus personatus actites Schwartz characterized in males by a combination of a tan to brown middorsal zone with dirty Leiocephalus personatus actites Schwartz tan dorsolateral lines; a dark blackish-brown 1967:14. Type locality, “Sosúa, Puerto head with white supraorbital stripes; sides of Plata Province, República Dominicana.” body pea-green flecked with orange; venter Holotype, MCZ R-81088, an adult male and upper surfaces of hindlimbs bright pea- collected by A. Schwartz and R. Thomas green; and throat immaculate but blackish on 15 October 1963 (not examined by and with some bright orange on the chin author). (Schwartz 1967). Size is moderate (maximum 13

SVL in males is 74 mm, maximum SVL in DIAGNOSIS. This subspecies is characterized females is 60 mm); mean number of loreals in males by a pale gray middorsal region (4.5) moderate; median head shields modally with pale buff lateral stripes; ventral color 6; and supraorbital semicircles more often pale green with very pale green (nearly complete (88.7%) than not (Schwartz 1967). white) spotting; mask present but restricted and usually dorsal to ear opening; chin 4. Leiocephalus personatus budeni Schwartz and anterior throat dark gray rather than black; and sides of body red with pale gray- Leiocephalus personatus budeni Schwartz green centers on scales (Gali et al. 1988). 1967:19. Type locality, “12 km NE Size is large (maximum SVL in males is 83 Jarabacoa, 2000 feet (656 meters), La mm, maximum SVL in females is 60 mm); Vega Province, República Dominicana.” mean number of loreals (5.7) high; median Holotype, MCZ R-81089, an adult male head shields modally 5; and supraorbital collected by D. W. Buden and a “native semicircles more often complete (95%) than collector” on 27 November 1964 (not not (Gali et al. 1988). examined by author). REMARK. Schwartz (1967) “questionably” DIAGNOSIS. This subspecies is characterized assigned a single female specimen from in males by a combination of a grayish-brown Aquin, Département du Sud, Haiti, to dorsal zone with prominent buffy dorsolateral Leiocephalus personatus personatus (Gali et stripes and one transverse nuchal and one al. 1988). transverse scapular bar; sides of body darker gray-brown with no green or brick-red scales; 6. Leiocephalus personatus mentalis lateral stripes faintly pinkish; venter white Cochran with a faint greenish tinge; throat greenish with prominent dark brownish-gray smudges; Leiocephalus personatus mentalis Cochran and usually a transverse bar at the level of 1932:178. Type locality, “Jovéro, [El the second chin shield (Schwartz 1967). Size Seibo Province,] Dominican Republic.” is small (maximum SVL in males is 66 mm, Holotype, USNM 65772, an adult male maximum SVL in females is 52 mm); mean collected by Dr. W. L. Abbott on 19 number of loreals (4.0) low; median head February 1923 (not examined by author). shields modally 4; frontoparietals modally 4; and supraorbital semicircles more often DIAGNOSIS. This subspecies is characterized complete (59.4%) than not (Schwartz 1967). in males by a combination of a bright yellow and completely immaculate throat; mental 5. Leiocephalus personatus elattoprosopon scale dark brown; face mask extremely bold Gali, Schwartz, and Suarez dark brown and prominent against brown to tan dorsal and head coloration; dorsolateral Leiocephalus personatus elattoprosopon Gali lines tan and fairly prominent to absent et al. 1988:19. Type locality, “ca. 1 km depending on the shade of dorsal coloration; inland, basal portion of the Morne Dubois and no nuchal or scapular transverse bars but ‘peninsula,’ Dépt. de [sic] Sud, Haiti.” median crest scales tipped with dark brown Holotype, National Museum of Natural to black on the neck and shoulders giving a History (USNM) 197370, an adult male median series of dark dots (Schwartz 1967). collected by “native collectors” on 20 July Size is small (maximum SVL in males is 72 mm, 1971 (not examined by author). maximum SVL in females is 58 mm);mean 14 number of loreals (4.0) low; median head and R. Thomas on 24 August 1969 (not shields modally 5; frontoparietals modally 5 examined by author). (Schwartz 1967). DIAGNOSIS. This subspecies is character- REMARK. This subspecies intergrades with ized in males by a combination of a deep or- Leiocephalus personatus tarachodes (Schwartz ange to greenish orange throat with labials 1971). and adjoining marginal scales deep golden orange, reddish orange, or dull orange; ven- 7. Leiocephalus personatus poikilometes ter pale green; mental scale with large (rather Schwartz than small) black to dark gray blotch; dark brown face mask bold and prominent against Leiocephalus personatus poikilometes dorsal gray coloration; dorsolateral lines dark Schwartz 1969:82. Type locality, “10 gray to absent; no nuchal or scapular trans- km SE El Jorillo, 2050 feet (625 m), San verse bars but median crest scales tipped with Juan Province, República Dominicana.” dark brown to black on neck and shoulders; Holotype, USNM 165935, an adult male mask extends behind ear opening rather collected by R. K. Bobilin, J. K. Lewis, than terminating before ear opening; a dark and A. Schwartz on 6 August 1968 (not gray blotch on the sub- and infralabials [fide examined by author). Schwartz 1971] and the next row or two of gular scales below the angle of the jaws; top DIAGNOSIS. This subspecies is character- of head at its most extreme solid black with ized in males by a combination of a tan to scattered pale yellow flecks (Schwartz 1971). grayish tan middorsal zone with cream dor- Size is moderate (maximum SVL in males is solateral lines; upper surface of head tan, suf- 76 mm, maximum SVL in females is 59 mm); fused with dark brown and cream and with mean number of loreals (3.8) low; median white supraorbital lines; sides of body bright head scales modally 4; frontoparietals modal- orange flecked with green; venter and up- ly 3; and supraorbital semicircles more often per surfaces of hindlimbs bright pea-green; complete (87.9%) than not (Schwartz 1971). and throat immaculate but with some black clouding on chin followed by orange to rusty 9. Leiocephalus personatus scalaris Cochran suffusion (Schwartz 1969). Size is moderate (maximum SVL in males is 73 mm, maxi- Leiocephalus personatus scalaris Cochran mum SVL in females is 58 mm); mean num- 1932:181. Type locality, “Cap-Haïtien, ber of loreals (3.7) low; median head shields [Département du Nord,] Haiti.” Holotype, modally 5; and supraorbital semicircles more USNM 74054, an adult male collected often complete (92.9%) than not (Schwartz by A. J. Poole on 3–6 March 1928 (not 1969). examined by author). Leiocephalus personatus pulcherrimus Mertens 8. Leiocephalus personatus pyrrholaemus 1939a:50. Type locality, “Moncion 452 m Schwartz H.” (= 2 km S Monción, elevation 450 meters, Santiago Rodríguez Province, Leiocephalus personatus pyrrholaemus República Dominicana; Schwartz 1967). Schwartz 1971:178. Type locality, “9 km E Holotype, Forschungsinstitut and Las Galeras, Samaná Province, República Naturmuseum Senckenberg (SMF) 25757, Dominicana.” Holotype, Carnegie an adult male collected by R. Mertens on Museum of Natural History (CM) 52287, 15 March 1939 (not examined by author). an adult male collected by J. R. Dennis Synonymy fide Schwartz (1967). 15

DIAGNOSIS. This subspecies is characterized green; and underside of tail bright orange and in males by a combination of green to yellow- undersides of hindlimbs dull pea green (Gali green venter and throat, the latter almost and Schwartz 1982). Size is moderate (maxi- immaculate or with only a few dusky smudges mum SVL in males is 75 mm, maximum SVL usually not aligned into any discernible in females is 59 mm); mean number of loreals transverse bars; dorsal ground color varying (4.1) moderate; median head shields modally from tan to dark brown, often flecked or 4, frontoparietals modally 5; and supraorbital mottled with yellow, cream or brick-red even semicircles more often complete (80%) than in small individuals; dorsolateral lines faint not (Gali and Schwartz 1982). in adults (both sexes); and mask prominent and outlined above by an orange canthal- supraocular-temporal line and below by 11. Leiocephalus personatus tarachodes two or three orange vertical subocular bars, Schwartz which at times are continuous with an orange bar across the temporal region; nuchal and Leiocephalus personatus tarachodes Schwartz scapular bars faint or usually absent, their 1967:11. Type locality, “6 km SE Nagua, positions indicated by dark brown crest scale María Trinidad Sánchez Province, dots (Schwartz 1967). Size is large (maximum República Dominicana.” Holotype, MCZ SVL in males is 82 mm, maximum SVL in R-81087, an adult male collected by A. females is 63 mm); mean number of loreals Schwartz and R. Thomas on 26 October (3.9) low; median head shields modally 4; 1963 (not examined by author). frontoparietals modally 5 (Schwartz 1967). DIAGNOSIS. This subspecies is characterized REMARK. This subspecies intergrades with in males by a combination of green or gray Leiocephalus personatus tarachodes (Schwartz throat with two rather bold transverse lines 1967). composed of black smudges at the level of the fourth and sixth chin shields; mask present 10. Leiocephalus personatus socoensis but not especially conspicuous because Schwartz of dark brown head coloration, bordered below by two or three bright yellow to cream Leiocephalus personatus socoensis Gali and subocular and temporal spots; bright green Schwartz 1982:177. Type locality, “25 km ventral color; and dorsal ground color tan E San Pedro de Macorís, Río Cumayasa, with transverse nuchal and scapular bars La Romana Province [now San Pedro rarely present and usually reduced to a series de Macorís Province], República of black median dots on the dorsal crest scales Dominicana.” Holotype, USNM 197371, (Schwartz 1967). Size is moderate (maximum an adult male collected by local collectors SVL in males is 75 mm, maximum SVL in on 26 June 1975 (not examined by author). females is 63 mm); mean number of loreals (3.8) low; median head shields modally 4; DIAGNOSIS. This subspecies is character- frontoparietals modally 4; and supraorbital ized in males by a combination of tan to brown semicircles more often incomplete (63.2%) dorsum with dorsolateral stripes duff buffy; than complete (Schwartz 1967). throats with black smudges anteriorly, chin black but throats of large males not spotted REMARK. This subspecies intergrades with on immaculate chin; subocular region with Leiocephalus personatus scalaris (Schwartz a white stripe, with or without a single black 1967) and with Leiocephalus personatus slash; ventral ground color green to yellowish mentalis (Schwartz 1971). 16

12. Leiocephalus personatus trujilloensis LITERATURE CITED Mertens Ackermann, T. 2006. Schreibers Glattkopfle- guan Leiocephalus schreibersii. Natur und Leiocephalus personatus trujilloënsis Tier Verlag GmbH, Münster, Germany. Mertens 1939a:45. Type locality, 61 pp. “Küste am Deutsch-Dominikanischen Alfonso, Y. U. 2016. Oviductal egg develop- Tropenforschungs-Institut in Ciudad ment in the curly-tailed lizard Leiocepha- Trujillo, etwa 4 km westlich der Ozama- lus carinatus aquarius. Revista Cubana de Mündung” (= coast at the German- Ciencias Biológicas 5(2):1–4. Dominican Research Institute in Santo Alfonso, Y. U., G. Fajardo, E. Suarez, and Domingo [Distrito Nacional, Dominican K. L. Krysko. 2012. Sexual size dimor- Republic] about 4 km west of the phism, ovipositioning, and hatching in mouth of the Ozama River). Holotype, Leiocephalus macropus asbolomus (Squa- Forschungsinstitut and Naturmuseum mata: Leiocephalidae) in Alexander von Senckenberg (SMF) 26213, an adult male Humboldt National Park in eastern Cuba. collected by R. Mertens on 15 April 1939 IRCF Reptiles & Amphibians 19:230–236. (not examined by author). Anonymous. 1980. Classified ads. Notes from NOAH 7(11):14–15. DIAGNOSIS. This subspecies is characterized Anonymous. 1991. Species of wild animals in males by a combination of greenish-orange bred in captivity during 1988/1989 and throat with black smudges, confluent on multiple generation captive births. In- sides of head and neck with a black mask; ternational Zoo Yearbook 30:311–489. transverse neck bars reduced or absent; and [Reptiles, pp. 326–342]. a broad dark brown to rich reddish-brown Araújo, B. M. C. de. 2014. Utilização de Répteis dorsal zone, in large adult males not bordered Como Animais de Esctimação: Impli- by pale dorsolateral lines, and dotted with cações Conservacionistas. Unpublished bright yellow (Schwartz 1967). Size is thesis, Universidade Estadual da Paraí- moderate (maximum SVL in males is 78 mm, ba Campus I, Campina Grande, Brazil. maximum SVL in females is 60 mm); mean Available at http://dspace.bc.uepb.edu.br/ number of loreals (3.7) low; median head jspui/bitstream/123456789/4189/1/PDF - shields modally 4; frontoparietals modally 5 Bruna Monielly Carvalho de Araújo.pdf. (Schwartz 1967). Archived by WebCite at http://www.web- citation.org/784UyOa1N on 2 May 2019. ACKNOWLEDGMENTS. José Rosado, Arment, C. 2005. Herper’s Life List. A Field Museum of Comparative Zoology, Harvard Checklist for the Native and Introduced University, graciously examined the MCZ Herpetofauna of the Continental United syntypes of Leiocephalus personatus and States and Canada. Coachwhip Publica- helped in an effort to resolve the mystery tions, Landisville, Pennsylvania. 199 pp. of the unlocated syntypes of Leiocephalus Barbour, T. 1914. A contribution to the zoö- trigeminatus (see Remarks). Darrel R. geography of the West Indies, with espe- Frost clarified the status of the unpublished cial reference to amphibians and reptiles. database (Etheridge 2000) maintained by the Memoirs of the Museum of Comparative AMNH. Miguel A. Landestoy and Joseph Zoölogy at Harvard College 44:205–359, P. Burgess provided photographs. John S. 1 plate. Parmerlee, Jr. produced the map. Aaron M. Barbour, T. 1930. A list of Antillean reptiles Bauer helped with references. and amphibians. Zoologica (Scientific 17

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