Leptotyphlopidae Stejneger, 1892

Adalsteinsson, Solny A., Branch, William R., Trape, Sébastien, Vitt, Laurie J. & Hedges, S. Blair, 2009, Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squamata), Zootaxa 2244, pp. 1-50 : 9-13

Family Leptotyphlopidae Stejneger, 1892

Stenosomata Ritgen, 1828: 255. Type genus: Stenosoma Wagler, 1824 . [Preoccupied by Stenosoma Latreille, 1810 : Coleoptera and Stenosoma Lamarck, 1817 : Mollusca.]

Stenostomi Wiegmann and Ruthe, 1832: 160.

Stenosomina Bonaparte, 1845: 377.

Stenosomatidae Günther, 1885: 85.

Stenostomidae Cope, 1886: 481 .

Glauconiidae Boulenger, 1890: 242 . Type genus: Glauconia Gray, 1845 .

Leptotyphlopidae Stejneger, 1892 [dated 1891]: 501.

Type genus. Leptotyphlops Fitzinger, 1843:24 .

Diagnosis. Small and thin snakes sharing with other members of Scolecophidia cylindrical bodies, ventral scales not enlarged, reduced eyes with a single visual cell type in the retina, and the absence of neural spines. They have solidly constructed skulls with toothless premaxillary, maxillary, and palatine bones sutured to the braincase along with the nasals and prefrontals. They lack a left lung, a tracheal lung, and a left oviduct ( Dowling & Duellman 1978; Underwood 1967; Vitt & Caldwell 2009). Except for two species having 16 midbody scale rows and two others having 14 or 16 rows, all of the other members of the family usually have 14 midbody scale rows. The maximum adult size of each species ranges from 104 mm ( Leptotyphlops carlae ) to 460 mm ( Rhinoleptus koniagui ) in total length; see discussion of body size in leptotyphlopid snakes ( Hedges 2008).

Content. Two subfamilies, five tribes, three subtribes, 12 genera, and 116 species ( Table 1).

Distribution. The family is distributed in the New World and Old World. In the New World it is distributed from North America (California, Utah, and Kansas) south through the Atlantic drainage of Middle and South America (exclusive of the high Andes) to Uruguay and Argentina. It also occurs on San Salvador Island ( Bahamas), Hispaniola, the Lesser Antilles, Cozumel Island ( Mexico), Islas de Bahia and Swan Islands ( Honduras), San Andres and Providencia Islands ( Colombia), Bonaire, Margarita Islands, and Trinidad. In the Old World it is distributed throughout Africa (north and south of the Sahara Desert), the Arabian Peninsula, and in southwest Asia ( Turkey, Iran, Pakistan, and northwest India); and on islands off the coast of Africa and Arabia (Bazaruto archipelago, Pemba, Manda, Lamu, Bioko, and Socotra) ( Fig. 1).

This study Previous classification

SUBFAMILY EPICTINAE

Tribe Epictini , Subtribe Epictina

Epictia albifrons ( Wagler 1824) SAM Leptotyphlops albifrons

Epictia albipuncta ( Jan 1861) SAM Leptotyphlops albipunctus

Epictia alfredschmidti ( Lehr, Wallach, Köhler & Aguilar 2002) SAM Leptotyphlops alfredschmidti

Epictia australis (Freiberg & Orejas-Miranda 1968) SAM Leptotyphlops australis

Epictia borapeliotes ( Vanzolini 1996) SAM Leptotyphlops borapeliotes

Epictia collaris ( Hoogmoed 1977) SAM Leptotyphlops collaris

Epictia columbi ( Klauber 1939) WI Leptotyphlops columbi

Epictia diaplocia ( Orejas-Miranda 1969) SAM Leptotyphlops diaplocius

Epictia goudotii ( Duméril & Bibron 1844) MAM Leptotyphlops goudotii

Epictia magnamaculata ( Taylor 1940) MAM Leptotyphlops magnamaculatus

Epictia melanurus ( Schmidt & Walker 1943) SAM Leptotyphlops melanurus

Epictia munoai ( Orejas-Miranda 1961) SAM Leptotyphlops munoai

Epictia nasalis ( Taylor 1940) MAM Leptotyphlops nasalis

Epictia peruviana ( Orejas-Miranda 1969) SAM Leptotyphlops peruvianus

Epictia rubrolineata ( Werner 1901) SAM Leptotyphlops rubrolineatus

Epictia rufidorsa ( Taylor 1940) SAM Leptotyphlops rufidorsus

Epictia signata ( Jan 1861) SAM Leptotyphlops signatus

Epictia striatula ( Smith & Laufe 1945) SAM Leptotyphlops striatula

Epictia subcrotilla ( Klauber 1939) SAM Leptotyphlops subcrotillus

Epictia teaguei ( Orejas-Miranda 1964) SAM Leptotyphlops teaguei

Epictia tenella ( Klauber 1939) SAM Leptotyphlops tenellus

Epictia tesselata ( Tschudi 1845) SAM Leptotyphlops tesselatus

Epictia tricolor ( Orejas-Miranda & Zug 1974) SAM Leptotyphlops tricolor

Epictia undecimstriata ( Schlegel 1839) SAM Leptotyphlops undecimstriatus

Epictia vellardi ( Laurent 1984) SAM Leptotyphlops vellardi

Siagonodon borrichianus ( Degerbøl 1923) SAM Leptotyphlops borrichianus

Siagonodon brasiliensis ( Laurent 1949) SAM Leptotyphlops brasiliensis

Siagonodon cupinensis (Bailey & Carvalho 1946) SAM Leptotyphlops cupinensis

Siagonodon septemstriatus ( Schneider 1801) SAM Leptotyphlops septemstriatus

Tribe Epictini , Subtribe Renina

Rena affinis ( Boulenger 1884) SAM Leptotyphlops affinis

Rena boettgeri ( Werner 1899) NAM Leptotyphlops humilis continued next page. Rena bressoni ( Taylor 1939) MAM Leptotyphlops bressoni

Rena dimidiata ( Jan 1861) SAM Leptotyphlops dimidiatus

Rena dissecta ( Cope 1896) MAM, NAM Leptotyphlops dissectus

Rena dulcis ( Baird & Girard 1853) MAM, NAM Leptotyphlops dulcis

Rena humilis ( Baird & Girard 1853) MAM, NAM Leptotyphlops humilis

Rena maxima ( Loveridge 1932) MAM Leptotyphlops maximus

Rena myopica ( Garman 1883) MAM, NAM Leptotyphlops myopicus

Rena nicefori ( Dunn 1946) SAM Leptotyphlops nicefori

Rena unguirostris ( Boulenger 1902) SAM Leptotyphlops unguirostris Tricheilostoma anthracinum ( Bailey 1946) SAM Leptotyphlops anthracinus Tricheilostoma brevissimum ( Shreve 1964) SAM Leptotyphlops brevissimus Tricheilostoma dugandi ( Dunn 1944) SAM Leptotyphlops dugandi

Tricheilostoma fulginosum ( Passos, Caramaschi & Pinto 2006) SAM Leptotyphlops fulginosus Tricheilostoma guayaquilensis ( Orejas-Miranda & Peters 1970) SAM Leptotyphlops guayaquilensis Tricheilostoma joshuai ( Dunn 1944) SAM Leptotyphlops joshuai

Tricheilostoma koppesi (Amaral 1955) SAM Leptotyphlops koppesi

Tricheilostoma macrolepis ( Peters 1857) SAM Leptotyphlops macrolepis Tricheilostoma salgueiroi (Amaral 1955) SAM Leptotyphlops salgueiroi

Tribe Epictini , Subtribe Tetracheilostomina

Mitophis asbolepis ( Thomas, McDiarmid & Thompson 1985) WI Leptotyphlops asbolepis

Mitophis calypso ( Thomas, McDiarmid & Thompson 1985) WI Leptotyphlops calypso

Mitophis leptepileptus ( Thomas, McDiarmid & Thompson 1985) WI Leptotyphlops leptepileptus

Mitophis pyrites ( Thomas 1965) WI Leptotyphlops pyrites

Tetracheilostoma bilineatum ( Schlegel 1839) WI Leptotyphlops bilineatus Tetracheilostoma breuili ( Hedges 2008) WI Leptotyphlops breuili

Tetracheilostoma carlae ( Hedges 2008) WI Leptotyphlops carlae

Tribe Rhinoleptini

Guinea bicolor ( Jan 1860) WAF Leptotyphlops bicolor

Guinea broadleyi ( Wallach & Hahn 1997) WAF Leptotyphlops broadleyi

Guinea greenwelli ( Wallach & Boundy 2005) WAF Leptotyphlops greenwelli

Guinea sundewalli ( Jan 1861) WAF Leptotyphlops sundewalli

Rhinoleptus koniagui Villiers 1956 WAF Rhinoleptus koniagui

Rhinoleptus parkeri ( Broadley 1999) EAF Leptotyphlops parkeri

SUBFAMILY LEPTOTYPHLOPINAE

Tribe Epacrophini

Epacrophis boulengeri ( Boettger 1913) EAF Leptotyphlops boulengeri

Epacrophis drewesi ( Wallach 1996) EAF Leptotyphlops drewesi

Epacrophis reticulatus ( Boulenger 1906) EAF Leptotyphlops reticulatus continued next page.

Tribe Myriopholini

Myriopholis adleri ( Hahn & Wallach 1998) WAF Leptotyphlops adleri

Myriopholis albiventer (Hallermann & Rödel 1995) WAF Leptotyphlops albiventer

Myriopholis algeriensis ( Jacquet 1895) WAF Leptotyphlops algeriensis

Myriopholis blanfordii (Boulenger 1890) AR, SWA Leptotyphlops blanfordii

Myriopholis boueti ( Chabanaud 1917) WAF Leptotyphlops boueti

Myriopholis braccianii ( Scortecci 1929) EAF Leptotyphlops braccianii

Myriopholis burii ( Boulenger 1905) AR Leptotyphlops burii

Myriopholis cairi ( Duméril & Bibron 1844) WAF, EAF Leptotyphlops cairi

Myriopholis dissimilis ( Bocage 1886) EAF Leptotyphlops dissimilis

Myriopholis erythraeus ( Scortecci 1929) EAF Leptotyphlops erythraeus

Myriopholis filiformis ( Boulenger 1899) SOC Leptotyphlops filiformis

Myriopholis ionidesi ( Broadley & Wallach 2007) EAF Leptotyphlops ionidesi

Myriopholis longicauda ( Peters 1854) SAF, EAF Leptotyphlops longicaudus

Myriopholis macrorhyncha ( Jan 1860) WAF, EAF, AR, SWA Leptotyphlops macrorhynchus Myriopholis macrura ( Boulenger 1899) SOC Leptotyphlops macrurus

Myriopholis narirostris ( Peters 1867) WAF Leptotyphlops narirostris

Myriopholis natatrix ( Andersson 1937) WAF Leptotyphlops natatrix

Myriopholis nursii ( Anderson 1896) EAF, AR Leptotyphlops nursii

Myriopholis perreti ( Roux-estéve 1979) WAF Leptotyphlops perreti

Myriopholis phillipsi ( Barbour 1914) AR Leptotyphlops phillipsi

Myriopholis rouxestevae ( Trape & Mane 2004) WAF Leptotyphlops rouxestevae

Myriopholis tanae ( Broadley & Wallach 2007) EAF Leptotyphlops tanae

Myriopholis wilsoni ( Hahn 1978) SOC Leptotyphlops wilsoni

Myriopholis yemenica ( Scortecci 1933) AR Leptotyphlops yemenicus

Tribe Leptotyphlopini

Leptotyphlops aethiopicus Broadley & Wallach 2007 EAF Leptotyphlops aethiopicus Leptotyphlops conjunctus ( Jan 1861) SAF Leptotyphlops conjunctus Leptotyphlops distanti ( Boulenger 1892) SAF Leptotyphlops distanti

Leptotyphlops emini (Boulenger 1890) CAF Leptotyphlops emini

Leptotyphlops howelli Broadley & Wallach 2007 EAF Leptotyphlops howelli

Leptotyphlops incognitus Broadley & Broadley 1999 SAF Leptotyphlops incognitus

Leptotyphlops jacobseni Broadley & Broadley 1999 SAF Leptotyphlops jacobseni

Leptotyphlops kafubi ( Boulenger 1919) CAF Leptotyphlops kafubi

Leptotyphlops keniensis Broadley & Wallach 2007 EAF Leptotyphlops keniensis

Leptotyphlops latirostris ( Sternfield 1912) EAF Leptotyphlops latirostris

Leptotyphlops macrops Broadley & Wallach 1996 EAF Leptotyphlops macrops

Leptotyphlops mbanjensis Broadley & Wallach 2007 EAF Leptotyphlops mbanjensis continued next page. Leptotyphlops merkeri ( Werner 1909) EAF Leptotyphlops merkeri

Leptotyphlops monticolus ( Chabanaud 1917) CAF Leptotyphlops monticolus

Leptotyphlops nigricans ( Schlegel 1839) SAF, EAF Leptotyphlops nigricans

Leptotyphlops nigroterminus Broadley & Wallach 2007 EAF Leptotyphlops nigroterminus Leptotyphlops pembae Loveridge 1941 EAF Leptotyphlops pembae

Leptotyphlops pitmani Broadley & Wallach 2007 CAF Leptotyphlops pitmani

Leptotyphlops pungwensis Broadley & Wallach 1997 SAF Leptotyphlops pungwensis Leptotyphlops scutifrons ( Peters 1854 SAF) , EAF Leptotyphlops scutifrons

Leptotyphlops sylvicolus Broadley & Wallach 1997 SAF Leptotyphlops sylvicolus

Leptotyphlops telloi Broadley & Watson 1976 SAF Leptotyphlops telloi

Namibiana gracilior ( Boulenger 1910) SAF Leptotyphlops gracilior

Namibiana labialis (Sternfeld 1908) SAF Leptotyphlops labialis

Namibiana latifrons (Sternfeld 1908) SAF Leptotyphlops latifrons

Namibiana occidentalis ( Fitzsimons 1962) SAF Leptotyphlops occidentalis

Namibiana rostrata ( Bocage 1886) SAF Leptotyphlops rostratus

Remarks. Hahn (1980) and Wallach (1998) reviewed the systematics of the family and McDiarmid et al. (1999) provided synonymies of the family and species. A more recent list of species, including synonymies, is provided by Uetz et al. (2009). Several changes in classification at the species level are discussed below. The two subfamilies recognized here correspond to the two major divisions within the family based on the phylogenetic relationships ( Figs. 3–4). The first is a mostly New World group, but includes six species from West Africa, and comprises mostly short-tailed species. The second is an entirely Old World assemblage comprising mostly long-tailed species.