Curly-tailed lizard

Template:Use dmy dates Template:Short description Template:Automatic taxobox

Leiocephalidae, also known as the curlytail lizards or curly-tailed lizards, is a family of iguanian lizards restricted to the West Indies. One of the defining features of these lizards is that their tail often curls over. They were previously regarded as members of the subfamily Leiocephalinae within the family Tropiduridae. There are presently 30 known species, all in the genus Leiocephalus.

TaxonomyEdit

Phylogenetic evidence supports Leiocephalidae being the most basal extant member of the clade Pleurodonta, with it diverging from the rest of the suborder as early as the Late Cretaceous, about 91 million years ago.<ref>Template:Cite journal</ref> As with many other higher-order taxa endemic to the Caribbean, it likely colonized the Antilles from South America during the Cenozoic; however, its deep divergence time from other lizards supports a much more complex and less straightforward history in the West Indies compared to other modern taxa.<ref name=":0">Template:Cite journal</ref>

Phylogenetic analysis on the genus supports some members of the now-extinct Lesser Antillean Leiocephalus radiation being the most basal of the recent Leiocephalus, with the last-surviving members of this group, L. herminieri and L. roquetus, sharing traits not present in other curlytail lizards from the Greater Antilles and other areas, such as the absence of enlarged snout scales. The second most basal of the recent curlytail lizards is another recently extinct species, L. eremitus from Navassa, followed by all other members of the genus from the Bahamas and Greater Antilles. Another extinct species from the Lesser Antilles known only from fossil remains, L. cuneus of Antigua and Barbuda, is thought to be more closely related to more derived Leiocephalus from the Bahamas and Greater Antilles, such as L. carinatus, L. greenwayi and L. punctatus, than to the other, more basal Lesser Antillean and Navassa species.<ref name=":0" />

DistributionEdit

Curlytail lizards are native to the West Indies, with the extant (living) species in the Bahamas, Turks and Caicos, Cayman Islands, Cuba, Hispaniola (Haiti and the Dominican Republic) and nearby small islands.<ref name="worldcat.org">Template:Cite book</ref><ref name=CaribHerp>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Additionally, Leiocephalus carinatus and Leiocephalus schreibersii have been introduced to Florida.<ref name=Conant1998>Template:Cite book</ref>

Former distributionEdit

Curlytail lizards formerly had a much wider native range, being distributed south to Jamaica and east to Puerto Rico and several of the Lesser Antilles.<ref name="reptile-database.reptarium.cz">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> They went extinct in most of this range during the Quaternary extinction or Holocene extinction, with some members of the Lesser Antillean radiation surviving to recent times, perhaps remaining widespread until after European colonization. The last surviving members of the Lesser Antillean radiation, L. herminieri of Guadeloupe and L. roquetus of Martinique, went extinct during the early-mid 19th century.<ref name=":0" />

General anatomyEdit

The curly-tailed lizards vary in size depending on species, but typically are approximately Template:Cvt in snout-to-vent length. These lizards have no femoral pores, pterygoid teeth, or palatine teeth. Additionally, these lizards are observed to have overlapping scales.<ref name="worldcat.org"/>

BehaviorEdit

The curly-tailed lizards mostly forage on arthropods such as insects, but also commonly take flowers and fruits.<ref name="worldcat.org"/><ref name=Kircher2014>Template:Cite journal</ref> Large individuals will eat small vertebrates, including anoles.<ref name=Kircher2014/><ref>Template:Cite journal</ref>

As suggested by their name, most species of this family often lift their tail and curl it. This is done both when a potential predator is present and when not present, although in some curly-tailed lizard species it increases when a predator is present. It shows the fitness of the lizard to a would-be predator and—in the case of an attack—draws attention to the tail, which increases the lizard's chance of escaping.<ref name=Cooper2007>Template:Cite journal</ref><ref name=Kircher2017>Template:Cite journal</ref> Although it has been suggested that it also functions as a territorial display,<ref name=Conant1998/> studies have been unable to find support for this, as the tail curling does not vary when another member of the same species is present.<ref name=Kircher2017/>

Conservation status and extinctionsEdit

The conservation status of the species in this family varies greatly. Several species, for example Leiocephalus carinatus, are common and widespread. Others are rare and highly threatened, especially those restricted to a single small island or a single location on a larger island, like the critically endangered Leiocephalus (barahonensis) altavelensis from Alto Velo Island and critically endangered Leiocephalus onaneyi from Guantánamo Province in Cuba.<ref name=CaribHerp/> Primary threats to their survival are habitat loss (for example, expanding agriculture, charcoal production and grazing goats) and introduced predators (for example, small Indian mongoose).<ref>Template:Cite iucn</ref><ref name=EDGEendomychus>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name=EDGErhutidira>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Several species of Leiocephalus are already extinct, including all of the Jamaican, Puerto Rican and Lesser Antillean members of the genus.<ref name="reptile-database.reptarium.cz"/><ref>Template:Cite journal</ref> Some of these are only known from fossil or subfossil remains and became extinct in the Pleistocene or pre-Columbian era, but others such as two Lesser Antillean species and one from Navassa survived until comparatively recently, during the 19th century. Leiocephalus is the only known squamate genus to be entirely wiped out from the Lesser Antilles following European colonization; other reptilian genera that have also seen significant extirpations in the Lesser Antilles, such as Boa or Diploglossus, still retain relict populations on at least some islands, such as Dominica and Montserrat. This mass disappearance of Leiocephalus from the Lesser Antilles may be due to their inhabiting dry forests in littoral areas that were heavily exploited and deforested by early colonists. Few confirmed Leiocephalus fossil remains from after the early Holocene are known from the Lesser Antilles, which has raised doubts about their being only recently extirpated from this area; however, Leiocephalus fossil bones are small and closely resemble those of other lizard species, which may explain the lack of detection of Leiocephalus fossil bones from these areas aside from by the most highly trained palaeo-herpetologists.<ref name=":0" />

In modern times, three species, Leiocephalus endomychus, Leiocephalus pratensis and Leiocephalus rhutidira, have not been seen since the 1960s and 1970s and are recognized as critically endangered, possibly extinct, by the IUCN. They are among the "most wanted" EDGE species.<ref name="EDGEendomychus" /><ref name="EDGErhutidira" /><ref name="EDGEpratensis">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

Newly discovered speciesEdit

Lizards of this family are diurnal and mostly inhabit fairly open habitats in a generally well-studied part of the world. Consequently, the majority of the species and subspecies already were scientifically described several decades ago. In 2016, the first new curly-tailed lizard since the early 1980s was described. The species was found in the coastal dunes of Bahía de las Calderas in the southwestern Dominican Republic. This species differs from the rest within Leiocephalidae in that its bony parietal table is U-shaped versus V-shaped, the males have 3–4 enlarged post-postcloacal scales versus 2, and there are specific sexual dimorphism trails.<ref>Template:Cite journal</ref>

Species and subspeciesEdit

The following species and subspecies, listed alphabetically by scientific name, are recognized as being valid by the Reptile Database.<ref name=ReptileDatabase>Template:NRDB genus</ref>

Extant and recently extinct speciesEdit

Image	Scientific name	Common Name	Subspecies	Distribution
	Leiocephalus barahonensis Template:Small	orange-bellied curlytail	L. b. altavelensis Template:Small – Alto Velo curly-tailed lizard, Alto Velo curlytail (likely better regarded as a separate species)<ref>Template:Cite iucn</ref><ref>{{#invoke:citation/CS1\|citation	CitationClass=web }}</ref> L. b. aureus Template:Small L. b. barahonensis Template:Small L. b. beatanus Template:Small L. b. oxygaster Template:Small	Hispaniola
File:Leiocephalus carinatus armouri.jpg	Leiocephalus carinatus Template:Small	saw-scaled curlytail, northern curly-tailed lizard	L. c. carinatus Gray, 1827 L. c. aquarius Schwartz & Ogren, 1956 L. c. armouri Barbour & Shreve, 1935 L. c. cayensis Schwartz, 1959 L. c. coryi K.P. Schmidt, 1936 L. c. granti Rabb, 1957 L. c. hodsdoni K.P. Schmidt, 1936 L. c. labrossytus Schwartz, 1959, South Central Cuba, Playa Larga L. c. microcyon Schwartz, 1959 L. c. mogotensis Schwartz, 1959 L. c. virescens Stejneger, 1901 L. c. zayasi Schwartz, 1959	Bahama Islands, the Cayman Islands and Cuba
File:Cuban Brown Curlytail (Leiocephalus cubensis) (8591588491).jpg	Leiocephalus cubensis Template:Small	Cuban brown curlytail, Cuban curlytail lizard	L. c. cubensis Template:Small L. c. gigas Template:Small L. c. minor Template:Small L. c. pambasileus Template:Small L. c. paraphrus Template:Small	Cuba.
	Leiocephalus endomychus Template:Small	Hinche curlytail, Central Haitian curlytail (possibly extinct, last seen in 1976)		Haiti
	†Leiocephalus eremitus Template:Small	Navassa curlytail lizard (extinct, 19th century)		Navassa Island
	Leiocephalus greenwayi Template:Small	East Plana curlytail, Plana Cay curlytail lizard		Bahama Islands
	†Leiocephalus herminieri Template:Small	Martinique curlytail lizard (extinct, 19th century)		Trinidad and Tobago
	Leiocephalus inaguae Template:Small	Inagua curlytail lizard		Bahamas
	Leiocephalus loxogrammus Template:Small	San Salvador curlytail, Rum Cay curlytail lizard	L. l. loxogrammus Template:Small L. l. parnelli Template:Small	Bahamas
File:Leiocephalus lunatus.jpg	Leiocephalus lunatus Template:Small	Hispaniolan maskless curlytail, Santo Domingo curlytail lizard	L. l. arenicolor Template:Small L. l. lewisi Template:Small L. l. louisae Template:Small L. l. lunatus Template:Small L. l. melaenoscelis Template:Small L. l. thomasi Template:Small	Dominican Republic.
File:Cuban Side-blotched Curlytail (Leiocephalus macropus) (8592688740).jpg	Leiocephalus macropus Template:Small	Cuban side-blotched curlytail, Monte Verde curlytail lizard	L. m. aegialus Template:Small L. m. asbolomus Template:Small L. m. felinoi Template:Small L. m. hoplites Template:Small L. m. hyacinthurus Template:Small L. m. immaculatus Template:Small L. m. koopmani Template:Small L. m. lenticulatus Template:Small L. m. macropus Template:Small L. m. phylax Template:Small L. m. torrei Template:Small	Cuba.
File:Tiburon Curlytail imported from iNaturalist photo 112020623 on 21 April 2022.jpg	Leiocephalus melanochlorus Template:Small	Tiburon curlytail, Jérémie curlytailed lizard	L. m. hypsistus Template:Small L. m. melanochlorus Template:Small	Haiti.
	Leiocephalus onaneyi Template:Small	Guantanamo striped curlytail, Guantanamo striped curly-tailed lizard, Sierra curlytail lizard		Cuba.
File:Leiocephalus-personatus-maskenleguan.jpg	Leiocephalus personatus Template:Small	Hispaniolan masked curlytail, Haitian curlytail lizard		Hispaniola.
	Leiocephalus pratensis Template:Small	Haitian striped curlytail, Atalaye curlytail lizard (possibly extinct, last seen in 1966)	L. p. chimarus Template:Small L. p. pratensis Template:Small	Haiti.
File:Leiocephalus psammodromus 189402506.jpg	Leiocephalus psammodromus Template:Small	Turks and Caicos curlytail, Bastion Cay curlytail lizard	L. p. aphretor Template:Small L. p. apocrinus Template:Small L. p. cacodoxus Template:Small L. p. hyphantus Template:Small L. p. mounax Template:Small L. p. psammodromus Template:Small	Turks and Caicos Islands
	Leiocephalus punctatus Template:Small	Crooked Acklins curlytail, spotted curlytail lizard		Bahamas.
	Leiocephalus raviceps Template:Small	pallid curlytail, mountain curlytail lizard	L. r. delavarai Template:Small L. r. jaumei Template:Small L. r. kilinikowski Template:Small L. r. raviceps Template:Small L. r. uzzelli Template:Small	Cuba.
	Leiocephalus rhutidira Template:Small	Haitian black-throated curlytail, Lapierre curlytail lizard (possibly extinct, last seen in 1978)		Haiti.
	†Leiocephalus roquetus Bochaton, Charles, and Lenoble, 2021	La Désirade curlytail lizard, curlytail roquet (extinct, late 19th century)		Guadeloupe.
File:Leiocephalus schreibersii Prague Zoo.jpg	Leiocephalus schreibersii Template:Small	red-sided curlytail, red-sided curly-tailed lizard	L. s. nesomorus Template:Small L. s. schreibersii Template:Small	Hispaniola
	Leiocephalus semilineatus Template:Small	Hispaniolan pale-bellied curlytail, Thomazeau curlytail lizard, Pale-bellied Hispaniolan curlytail		Hispaniola.
File:Leiocephalus sixtoi P1010419.jpg	Leiocephalus sixtoi Template:Small	Hispaniolan dune curlytail		Hispaniola.
File:Cabo Corrientes Curlytail lizard Leiocephalus stictigaster - Flickr - gailhampshire.jpg	Leiocephalus stictigaster Template:Small	Cuban striped curlytail, Cabo Corrientes curlytail lizard	L. s. astictus Template:Small L. s. celeustes Template:Small L. s. exotheotus Template:Small L. s. gibarensis Template:Small L. s. lipomator Template:Small L. s. lucianus Template:Small L. s. naranjoi Template:Small L. s. ophiplacodes Template:Small L. s. parasphex Template:Small L. s. septentrionalis Template:Small L. s. sierrae Template:Small L. s. stictigaster Template:Small	Cuba.
File:Cayman curly-tailed lizard (Leiocephalus varius).jpg	Leiocephalus varius Template:Small	Cayman curlytail, Cayman curly-tailed lizard		Cayman Islands.
	Leiocephalus vinculum Template:Small	Gonave curlytail, Cochran's curlytail lizard		Haiti.

Fossil and subfossil speciesEdit

†Leiocephalus anonymous Template:Small – Hispaniola
†Leiocephalus apertosulcus Template:Small – Hispaniola
†Leiocephalus cuneus Template:Small – Leeward Islands curlytail (extinct, Late Quaternary of Antigua and Barbuda and potentially Anguilla, La Désirade, and Marie-Galante, but might have survived to recent times)
†Leiocephalus etheridgei Template:Small – Morovis curlytail (extinct, Late Pleistocene of Puerto Rico)
†Leiocephalus jamaicensis Template:Small – Jamaican curlytail (extinct, Late Pleistocene or Holocene of Jamaica)
†Leiocephalus partitus Template:Small – Guánica curlytail (extinct, Pleistocene or Holocene of Puerto Rico)

Nota bene: A binomial authority or trinomial authority in parentheses indicates that the species or subspecies was originally described in a genus other than Leiocephalus.

ReferencesEdit

Template:Reflist