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Leiocephalus psammodromus BARBOUR, 1920

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Higher TaxaLeiocephalidae, Iguania, Sauria, Squamata (lizards)
SubspeciesLeiocephalus psammodromus psammodromus BARBOUR 1920
Leiocephalus psammodromus aphretor SCHWARTZ 1967
Leiocephalus psammodromus apocrinus SCHWARTZ 1967
Leiocephalus psammodromus cacodoxus SCHWARTZ 1967
Leiocephalus psammodromus hyphantus SCHWARTZ 1967
Leiocephalus psammodromus mounax SCHWARTZ 1967 
Common NamesE: Turks & Caicos Curlytail, Bastion Cay Curlytail Lizard 
SynonymLiocephalus (sic) arenarius BARBOUR 1916: 217
Leiocephalus psammodromus BARBOUR 1920 (nom. subst.)
Leiocephalus psammodromus — SCHWARTZ & HENDERSON 1991: 443
Leiocephalus psammodromus — SMITH 1994: 120

Leiocephalus psammodromus psammodromus BARBOUR 1920
Leiocephalus psammodromus psammodromus — PREGILL 1992: 50
Leiocephalus psammodromus psammodromus — HEDGES et al. 2019

Leiocephalus psammodromus aphretor SCHWARTZ 1967
Leiocephalus arenarius aphretor SCHWARTZ 1967: 163
Leiocephalus psammodromus aphretor — PREGILL 1992: 50
Leiocephalus psammodromus aphretor — HEDGES et al. 2019

Leiocephalus psammodromus apocrinus SCHWARTZ 1967
Leiocephalus arenarius apocrinus SCHWARTZ 1967: 165
Leiocephalus psammodromus apocrinus — PREGILL 1992: 50
Leiocephalus psammodromus apocrinus — HEDGES et al. 2019

Leiocephalus psammodromus cacodoxus SCHWARTZ 1967
Leiocephalus arenarius cacodoxus SCHWARTZ 1967: 176
Leiocephalus psammodromus cacodoxus — PREGILL 1992: 50
Leiocephalus psammodromus cacodoxus — HEDGES et al. 2019

Leiocephalus psammodromus hyphantus SCHWARTZ 1967
Leiocephalus arenarius hyphantus SCHWARTZ 1967: 172
Leiocephalus psammodromus hyphantus — KLUGE 1984
Leiocephalus psammodromus hyphantus — PREGILL 1992: 50
Leiocephalus psammodromus hyphantus — HEDGES et al. 2019

Leiocephalus psammodromus mounax SCHWARTZ 1967
Leiocephalus arenarius mounax SCHWARTZ 1967: 169
Leiocephalus psammodromus mounax — PREGILL 1992: 50
Leiocephalus psammodromus mounax — HEDGES et al. 2019 
DistributionTurks I, Caicos I, scatteren on associated Cays

psammodromus: Turks Is.: "Bastion Cay," Big Sand Cay; Type locality: 'Bastion Cay," Turks Islands (unlocatable).

aphretor: Turks Is.: Long Cay, SE of Grand Turk I.

apocrinus: Caicos Is.: Big Ambergris and Little Ambergris cays.

cacodoxus: Caicos Is.: Ft. George Cay, Providenciales I., Sugar Loaf I.

hyphantus: Caicos Is.: Pine Cay, Water Cay, Stubb Cay.

mounax: Caicos Is.: Long Cay off Cockburn Harbour, South Caicos I.

INCERTAE SEDIS: The species is also known from (Caicos Is.) West Caicos I., Dellis Cay, Little Water Cay, Parrot Cay, North Caicos I., Middle Caicos I., East Caicos I., and (Turks Is.) Gibbs Cay, East Cay, Pear Cay; all are unassigned subspecifically.  
TypesHolotype: MCZ 11948.
Holotype: CM 40602 [aphretor]
Holotype: CM 40601 [apocrinus]
Holotype: MCZ 54185 [cacodoxus]
Holotype: UMMZ 126624 [hyphantus]
Holotype: CM 40603 [mounax] 
DiagnosisDESCRIPTION: Size moderate (SVL in males to 105 mm, in females to 83 mm);
strong sexual dichromatism, males with a pair of enlarged postanals; a distinct lateral fold, with smaller scales in fold; dorsals imbricate, keeled; ventrals imbricate, smooth, weakly denticulate; median dorsal crest scales enlarged, not strongly overlapping, lower than median dorsal caudal scales, 56-81 in occiput-vent distance; one-half midbody scales 28-56; supraoculars 5-8 (strongly modally 7); loreals 4-14; temporals 10-18; supraorbital semicircles more often complete; parietals more often in contact; no median azygous scale between posterior pair of median head scales; median head scales 3-12 (mode variable by population); prefrontal row 3-7 scales, usually complete; frontoparietal row 2-7 scales, complete or incomplete. Dorsum (males) pale yellowish tan or yellow; 2 nuchal, and 1 dark brown to velvety black, transverse axillary bars, followed by 2 postaxillary bars, these bars sometimes limited (by subspecies) to anteriormost 3, confluent or not, or at times entire dorsum overlaid with black stippling or marbling, rendering transverse bars inconspicuous; at times about 5 more vague, transverse bars before hindlimbs; interspaces between bars finely stippled or speckled with brown or marbled with black; each dorsal bar with concomitant dark lateral blotch or spot (= 3 dark spots on neck and above axilla, 2 postaxillary), these dark spots variously fused, in 1 subspecies giving a single (but obviously quinquepartite) blotch; limbs stippled with dark brown or dark grayish brown with paler spotting, or medium brownish, dotted black (forelimbs) and black and yellow (hindlimbs); upper surface of head yellowish tan with much paler and darker marbling, or brown very heavily flecked with yellow; throat pattern 2-3 brown or black Vs, apices pointed anteriorly, or their fragments (by subspecies), and with (at times) a brownish, transverse bar across chin, followed by a few scattered, brownish chest flecks or dots; throat ground color yellow, much clouded at times with grayish purple, or mottled gray and yellow; venter pale yellow or grayish yellow, immaculate; tail unicolor yellowish tan to brown above, with 1-2 dark dumbbells proximally, nonchevronate or chevronate only distally; dorsum (females) yellowish tan, dull tan, drab brownish, or dull gray, with about 10-15 brownish, blackish, or grayish brown, transverse bars (often constricted medially to form dumbbells) outlined with tan edges or tan to cream flecks; sides with or without a series of brownish squares corresponding to transverse dorsal bars; limbs tan, flecked with paler pigment; lateral fold not conspicuously marked with pale ovals, but occasionally with a cream line along fold edge; upper surface of tail basally chevronate, but banded with darker brown to tip; lower sides variously marbled, stippled, or flecked and with 4-7 short ventrolateral bars, composed of pearly scales, restricted to sides of venter; throat with 2-3 pairs of gray, diagonal lines or their fragments, followed by longitudinal dashes and gray spotting on chest, or reticulate; limbs brown with (forelimbs) brown or black flecking and marbling or (hindlimbs) diffuse, dark brown and cream marbling (Schwartz & Henderson 1991: 443).

DIAGNOSIS (cacodoxus): A subspecies af L. arenarius characterized by a cambinatian of moderately dark dorsum with four or five lateral black blotches fused into a single black lateral band, five (two nuchal, one supra-axillary, two postaxillary) dorsal black bars which do not cannect laterally with the black lateral blotch, remainder of dorsum without prominent bars or bar remnants and moderately stippled with dark, throat pattern consisting of a complete set of gray diagonal lines, often with a third pair of diagonals on the posterior portion of the throat, semicircles usually not complete, and parietals modally in contact (Schwartz 1967: 176)

COMPARISONS (cacodoxus, needs OCR corrections): By virtue of its throat pattern, L. a. cacodoxus is readily distinguishable from its closest neighbor, L. a. hyphantus. In males, the latter has a complex reticulate throat pattern on a dark ground, the former a clearly delineated gray pattern of diagonals on a clear pale ground. Doubtless in life there are pigmental differences in this region as well. The solid black anterolateral blotch in cacodoxus contrasts with the obsolescent blotches in hyphantus. Of the other subspecies, in throat pattern cacodoxus most closely resembles mounax, but the extra set of diagonals and the lateral solid blotch will differentiate the two with ease. These two subspecies occupy islands at almost the two extremes of the Caicos arc.
The Ft. George Cay series has been discussed above. The three males and one female have the throat patterns so clearly like the topotypes of cacodoxus, rather than the heavily patterned throats of hyphantus, that lizards on this basis are clearly referrable to the former subspecies. The lateral blotches are discrete and do not form a single blotch, and on this basis the lizards are closer to hyphantus. I assign them to cacodoxus, but possibly they might be better regarded as intergrades between these two races on the basis of combination of characters. In favor of this intergradient interpretation is the fact that the semicircles are modally complete whereas the parietals are modally not in contact-precisely the condition in hyphantus in contrast to that in cacodoxus.
As far as head scalation is concerned, only cacodoxus and aphl'etol' have the semicircles usually incomplete, and only cacodoxus has the parietals modally in contact. The modal head scale formula of cacodoxus (3-4 or 6-5) is confusing and doubtless due to the small series involved. Of interest is the fact that in having a modal condition of three prefrontals and parietal contaot, cacodoxus (inhabiting islands near the extreme western end of the Caicos Bank) resembles L. inaguae. L. a. cacodoxus is the only subspecies of L. al'enal'ius which has these two conditions modally (Schwartz 1967: 178)

DIAGNOSIS (hyphantus): A subspecies of L. arenarius characterized by a combination of pale dorsum with one (axillary) to three (usually one axillary, two nuchal) discrete lateral black blotches, three (two nuchal, one supra-axillary) black velvety transverse dorsal bars followed by about six to ten less prominent but still distinguishable dorsal black bars or pairs of sports to the sacrum, throat pattern consisting of a black reticulum, and supraorbital semicircles usually in contact (Schwartz 1967: 172)

COMPARISONS (hyphantus): Because of its reticulate throat pattern in both sexes and the virtually completely patterned dorsum in males, L. a. hyphantus is readily distinguishable from all previously named subspecies. As noted above, some male hyphantus resemble some arenarius in lack of dorsal black obliterative stippling and marbling. These peculiarly marked male hyphantus can be easily distinguished from arenarius by the reticulate throat pattern. In scales, hyphantus differs from aphretor in having the semicircles modally complete. The head scale formula of 4-5-4 is the same as that of aphretor, but differs from the formulae of other subspecies (Schwartz 1967: 175). 
CommentFor illustrations see Schwartz, 1967.

Nomenclature: Liocephalus (sic) arenarius BARBOUR 1916 is preoccupied by Steironotus [Ophryoesoides = Leiocephalus] arenarius TSCHUDI 1845.

Subspecies: Genetic data suggests that subspecies names do not reflect actual units of variation and need revision (Reynolds et al. 2012). 
EtymologyThe name hyphantus is derived from the Greek for "woven," an allusion to the net-like or reticulate throat pattern. 
  • Barbour, T. 1916. Additional notes on West Indian reptiles and amphibians. Proc. Biol. Soc. Washington 29: 215-220. - get paper here
  • Hedges SB, Powell R, Henderson RW, Hanson S, and Murphy JC 2019. Definition of the Caribbean Islands biogeographic region, with checklist and recommendations for standardized common names of amphibians and reptiles. Caribbean Herpetology 67: 1–53
  • Pregill,G.K. 1992. Systematics of the West Indian Lizard Genus Leiocephalus (Squamata: Iguania: Tropiduridae). Misc. Publ. Univ. Kans. Mus. Nat. Hist. 84: 1-69 - get paper here
  • Reynolds, R. Graham; Matthew L. Niemiller; Benjamin M. Fitzpatrick 2012. Genetic analysis of an endemic archipelagic lizard reveals sympatric cryptic lineages and taxonomic discordance. Conservation Genetics 13:953–963 - get paper here
  • Schwartz, A. 1967. The Leiocephalus (Lacertilia, Iguanidae) of the southem Bahama Islands. Annals of the Carnegie Museum 39 (12): 153-84.
  • Schwartz, A. & Henderson, R.W. 1991. Amphibians and Reptiles of the West Indies. University of Florida Press, Gainesville, 720 pp.
  • Smith, G. R. 1994. Observations on activity in two lizards (Anolis scriptus and Leiocephalus psammodromus). Bull. Maryland Herp. Soc. 30 (3): 120-125 - get paper here
  • Smith, Geoffrey R. 1992. Sexual dimorphism in the curly-tailed lizard, Leiocephalus psammodromus. Carib. J. Sci. 28 (1-2): 99-101
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