Leiocephalus

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Lepidosauromorpha
Superordo: Lepidosauria
Ordo: Squamata
Cladus: Unidentata Episquamata Toxicofera
Subordo: Iguania
Infraordo: Pleurodonta

Familia: Leiocephalidae
Genus: Leiocephalus
Species (29 + †3) †L. anonymous – †L. apertosulcus – L. barahonensis – L. carinatus – L. cubensis – L. cuneus – L. endomychus – L. eremitus – L. etheridgei – L. greenwayi – L. herminieri – L. inaguae – L. jamaicensis – L. loxogrammus – L. lunatus – L. macropus – L. melanochlorus – L. onaneyi – L. partitus – L. personatus – L. pratensis – L. psammodromus – L. punctatus – L. raviceps – L. rhutidira – †L. roquetus – L. schreibersii – L. semilineatus – L. sixtoi – L. stictigaster – L. varius – L. vinculum
Name

Leiocephalus Gray, 1827: 207

Type species: Leiocephalus carinatus Gray, 1827, by monotypy.

References
Primary references

Gray, J.E. 1827. A description of a new genus and some new species of saurian reptiles; with a revision of the species of chameleons. The Philosophical Magazine. Series II 2(9): 207–214. DOI: 10.1080/14786442708675648 Paywall; BHL Reference page.

Additional references

Köhler, G., Bobadilla, M.J.R. & Hedges, S.B. 2016. A new dune-dwelling lizard of the genus Leiocephalus (Iguania, Leiocephalidae) from the Dominican Republic. Zootaxa 4121(5): 517–532. DOI: 10.11646/zootaxa.4121.5.2. Reference page.
Bochaton, C., Charles, L. & Lenoble, A. 2021. Historical and fossil evidence of an extinct endemic species of Leiocephalus (Squamata: Leiocephalidae) from the Guadeloupe Islands. Zootaxa 4927(3): 383–409. DOI: 10.11646/zootaxa.4927.3.4 Paywall Reference page.

Links

Uetz, P. & Hallermann, J. 2022. Leiocephalus . The Reptile Database. Accessed on 25 May 2021.
Leiocephalus – Taxon details on Integrated Taxonomic Information System (ITIS).

Vernacular names
English: Curly-tailed Lizards
español: Lagartos de cola rizada

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 29 known species, all in the genus Leiocephalus.

Taxonomy

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.[2] 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.[3]

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.[3]
Distribution

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.[4][5] Additionally, Leiocephalus carinatus and Leiocephalus schreibersii have been introduced to Florida.[6]
Former distribution

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.[7] They went extinct in most of this range during the Quaternary extinction (with some such as the Jamaican taxon L. jamaicensis going extinct during the Late Pleistocene, well before the arrival of the first Amerindians to the area),[8] but some members of the Lesser Antillean radiation survived to more 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.[3]
General anatomy

The curly-tailed lizards vary in size depending on species, but typically are approximately 9 cm (3.5 in) 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.[4]
Behavior

The curly-tailed lizards mostly forage on arthropods such as insects, but also commonly take flowers and fruits.[4][9] Large individuals will eat small vertebrates, including anoles.[9][10]

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.[11][12] Although it has been suggested that it also functions as a territorial display,[6] 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.[12]
Conservation status and extinctions

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.[5] 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).[13][14][15]

Several species of Leiocephalus are already extinct, including all of the Jamaican, Puerto Rican and Lesser Antillean members of the genus.[7][16] 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.[3]

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.[14][15][17]
Newly discovered species

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.[18]
Species and subspecies
Leiocephalus carinatus
Leiocephalus cubensis
Leiocephalus macropus
Leiocephalus personatus

The following species and subspecies, listed alphabetically by scientific name, are recognized as being valid by the Reptile Database.[19]
Extant and recently extinct species

Leiocephalus barahonensis Schmidt, 1921 – orange-bellied curlytail
L. b. altavelensis Noble & Hassler, 1933 – Alto Velo curly-tailed lizard, Alto Velo curlytail (likely better regarded as a separate species)[20][21]
L. b. aureus Cochran, 1934
L. b. barahonensis Schmidt, 1921
L. b. beatanus Noble, 1923
L. b. oxygaster A. Schwartz, 1967
Leiocephalus carinatus Gray, 1827 – 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
Leiocephalus cubensis (Gray, 1840) – Cuban brown curlytail, Cuban curlytail lizard
L. c. cubensis (Gray, 1840)
L. c. gigas A. Schwartz, 1959
L. c. minor Varona & Garrido, 1970
L. c. pambasileus A. Schwartz, 1959
L. c. paraphrus A. Schwartz, 1959
Leiocephalus endomychus A. Schwartz, 1967 – Hinche curlytail, Central Haitian curlytail (possibly extinct, last seen in 1976)
†Leiocephalus eremitus (Cope, 1868) – Navassa curlytail lizard (extinct, 19th century)
Leiocephalus greenwayi Barbour & Shreve, 1935 – East Plana curlytail, Plana Cay curlytail lizard
†Leiocephalus herminieri (A.M.C. Duméril & Bibron, 1837) – Martinique curlytail lizard (extinct, 19th century)
Leiocephalus inaguae Cochran, 1931 – Inagua curlytail lizard
Leiocephalus loxogrammus (Cope, 1887) – San Salvador curlytail, Rum Cay curlytail lizard
L. l. loxogrammus (Cope, 1887)
L. l. parnelli Barbour & Shreve, 1935
Leiocephalus lunatus Cochran, 1934 – Hispaniolan maskless curlytail, Santo Domingo curlytail lizard
L. l. arenicolor Mertens, 1939
L. l. lewisi A. Schwartz, 1967
L. l. louisae Cochran, 1934
L. l. lunatus Cochran, 1934
L. l. melaenoscelis A. Schwartz, 1967
L. l. thomasi A. Schwartz, 1967
Leiocephalus macropus (Cope, 1863) – Cuban side-blotched curlytail, Monte Verde curlytail lizard
L. m. aegialus A. Schwartz & Garrido, 1967
L. m. asbolomus A. Schwartz & Garrido, 1967
L. m. felinoi Garrido, 1979
L. m. hoplites Zug, 1959
L. m. hyacinthurus Zug, 1959
L. m. immaculatus Hardy, 1958
L. m. koopmani Zug, 1959
L. m. lenticulatus Garrido, 1973
L. m. macropus (Cope, 1863)
L. m. phylax A. Schwartz & Garrido, 1967
L. m. torrei Garrido, 1979
Leiocephalus melanochlorus (Cope, 1863) – Tiburon curlytail, Jérémie curlytailed lizard
L. m. hypsistus Schwartz, 1966
L. m. melanochlorus (Cope, 1863)
Leiocephalus onaneyi Garrido, 1973 – Guantanamo striped curlytail, Guantanamo striped curly-tailed lizard, Sierra curlytail lizard
Leiocephalus personatus (Cope, 1863) – Hispaniolan masked curlytail, Haitian curlytail lizard
Leiocephalus pratensis (Cochran, 1928) – Haitian striped curlytail, Atalaye curlytail lizard (possibly extinct, last seen in 1966)
L. p. chimarus A. Schwartz, 1979
L. p. pratensis (Cochran, 1928)
Leiocephalus psammodromus Barbour, 1920 – Turks and Caicos curlytail, Bastion Cay curlytail lizard
L. p. aphretor A. Schwartz, 1967
L. p. apocrinus A. Schwartz, 1967
L. p. cacodoxus A. Schwartz, 1967
L. p. hyphantus A. Schwartz, 1967
L. p. mounax A. Schwartz, 1967
L. p. psammodromus Barbour, 1920
Leiocephalus punctatus Cochran, 1931 – Crooked Acklins curlytail, spotted curlytail lizard
Leiocephalus raviceps (Cope, 1863) – pallid curlytail, mountain curlytail lizard
L. r. delavarai Garrido, 1973
L. r. jaumei A. Schwartz & Garrido, 1968
L. r. kilinikowski A. Schwartz, 1960
L. r. raviceps (Cope, 1863)
L. r. uzzelli A. Schwartz, 1960
Leiocephalus rhutidira A. Schwartz, 1979 – Haitian black-throated curlytail, Lapierre curlytail lizard (possibly extinct, last seen in 1978)
†Leiocephalus roquetus Bochaton, Charles, and Lenoble, 2021 – La Désirade curlytail lizard, curlytail roquet (extinct, late 19th century)
Leiocephalus schreibersii (Gravenhorst, 1838) – red-sided curlytail, red-sided curly-tailed lizard
L. s. nesomorus A. Schwartz, 1968
L. s. schreibersii (Gravenhorst, 1838)
Leiocephalus semilineatus Dunn, 1920 – Hispaniolan pale-bellied curlytail, Thomazeau curlytail lizard, Pale-bellied Hispaniolan curlytail
Leiocephalus sixtoi Kohler, Bobadilla, & Hedges, 2016 – Hispaniolan dune curlytail
Leiocephalus stictigaster A. Schwartz, 1959 – Cuban striped curlytail, Cabo Corrientes curlytail lizard
L. s. astictus A. Schwartz, 1959
L. s. celeustes A. Schwartz & Garrido, 1968
L. s. exotheotus A. Schwartz, 1959
L. s. gibarensis A. Schwartz & Garrido, 1968
L. s. lipomator A. Schwartz & Garrido, 1968
L. s. lucianus A. Schwartz, 1960
L. s. naranjoi A. Schwartz & Garrido, 1968
L. s. ophiplacodes A. Schwartz, 1964
L. s. parasphex A. Schwartz, 1964
L. s. septentrionalis Garrido, 1975
L. s. sierrae A. Schwartz, 1959
L. s. stictigaster A. Schwartz, 1959
Leiocephalus varius Garman, 1887 – Cayman curlytail, Cayman curly-tailed lizard
Leiocephalus vinculum Cochran, 1928 – Gonave curlytail, Cochran's curlytail lizard

Fossil species

†Leiocephalus cuneus Etheridge, 1964 – 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 Pregill, 1981 – Morovis curlytail (extinct, Late Pleistocene of Puerto Rico)
†Leiocephalus jamaicensis Etheridge, 1966 – Jamaican curlytail (extinct, Late Pleistocene of Jamaica)
†Leiocephalus partitus Pregill, 1981 – 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.
References

"Leiocepalus ". Dahms Tierleben. www.dahmstierleben.de.
Zheng, Yuchi; Wiens, John J. (1 January 2016). "Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species". Molecular Phylogenetics and Evolution. 94 (Pt B): 537–547. doi:10.1016/j.ympev.2015.10.009. ISSN 1055-7903. PMID 26475614.
Bochaton, Corentin; Charles, Laurent; Lenoble, Arnaud (15 February 2021). "Historical and fossil evidence of an extinct endemic species of Leiocephalus (Squamata: Leiocephalidae) from the Guadeloupe Islands". Zootaxa. 4927 (3): 383–409. doi:10.11646/zootaxa.4927.3.4. ISSN 1175-5334. PMID 33756701. S2CID 232337806.
J., Vitt, Laurie (2014). Herpetology : an introductory biology of amphibians and reptiles. Caldwell, Janalee P. (4th ed.). Amsterdam. ISBN 9780123869197. OCLC 839312807.
Hedges, B. "All Islands". CaribHerp. Retrieved 2 July 2019.
Conant, R.; J.T. Collins (1998). A Field Guide to the Reptiles and Amphibians of Eastern and Central North America (3 ed.). pp. 244–246. ISBN 978-0395904527.
"Search results | The Reptile Database". reptile-database.reptarium.cz. Retrieved 1 March 2021.
"Leiocephalus jamaicensis". The Reptile Database. Retrieved 1 March 2021.
Kircher, B.L.; C.D. Robinson; M.A. Johnson (2014). "Herbivory in the Northern Curly-tailed Lizard (Leiocephalus carinatus)". Caribbean Herpetology. 50: 1–2. doi:10.31611/ch.50.
Steinberg; Losos; Schoener; Spiller; Kolbe; Leal (2004). "Predation-associated modulation of movement-based signals by a Bahamian lizard". PNAS. 11 (25): 9187–9192. doi:10.1073/pnas.1407190111. PMC 4078856. PMID 24843163.
Cooper, W.E. (2007). "Escape and its relationship to pursuit‐deterrent signaling in the Cuban curly‐tailed lizard Leiocephalus carinatus". Herpetologica. 63 (2): 144–150. doi:10.1655/0018-0831(2007)63[144:EAIRTP]2.0.CO;2.
Kircher, B.K.; M.A. Johnson (2017). "Why do curly tail lizards (genus Leiocephalus) curl their tails? An assessment of displays toward conspecifics and predators". Ethology. 123 (5): 342–347. doi:10.1111/eth.12603.
Fong, A. (2017). "Leiocephalus onaneyi". IUCN Red List of Threatened Species. 2017: e.T178278A7512731. doi:10.2305/IUCN.UK.2017-2.RLTS.T75310301A75607524.en.{{cite iucn}}: error: |doi= / |page= mismatch (help)
"57. Central Haitian Curlytail". EDGE species. Retrieved 2 July 2019.
"73. Lapierre Curlytail Lizard". EDGE species. Retrieved 2 July 2019.
Meiri, S.; et al. (2017). "Extinct, obscure or imaginary: The lizard species with the smallest ranges" (PDF). Diversity and Distributions. 24 (2): 262–273. doi:10.1111/ddi.12678.
"75. Atalaye Curlytail Lizard". EDGE species. Retrieved 2 July 2019.
Köhler, Gunther; Bobadilla, Marcos J. Rodríguez; Hedges, S. Blair (13 June 2016). "A new dune-dwelling lizard of the genus Leiocephalus (Iguania, Leiocephalidae) from the Dominican Republic". Zootaxa. 4121 (5): 517–32. doi:10.11646/zootaxa.4121.5.2. ISSN 1175-5334. PMID 27395240. S2CID 9386834.
Leiocephalus at the Reptarium.cz Reptile Database. Accessed 2 July 2019.
Inchaustegui, S.; Landestoy, M.; Powell, R. & Hedges, B. (2017) [errata version of 2016 assessment]. "Leiocephalus altavelensis". IUCN Red List of Threatened Species. 2016: e.T75306189A115482003. doi:10.2305/IUCN.UK.2016-3.RLTS.T75306189A75607464.en. Retrieved 14 September 2021.

Hedges, S. Blair (2021). "Isla Alto Velo". Caribherp: Amphibians and reptiles of Caribbean Islands. Retrieved 14 September 2021.