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Indotyphlops braminus (DAUDIN, 1803)

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Higher TaxaTyphlopidae (Asiatyphlopinae), Typhlopoidea, Serpentes, Squamata (snakes)
Subspecies 
Common NamesE: Flowerpot Snake, Brahminy blindsnake, Bootlace Snake
G: Brahmanen-Wurmschlange
E: Khorat Worm Snake, Khorat Blind Snake [khoratensis]
G: Khorat-Blindschlange [khoratensis]
Chinese: 钩盲蛇
S: Culebrilla Ciega
Italian: tiflope bramino 
SynonymEryx braminus DAUDIN 1803: 279
Tortrix russelii MERREM 1820
Typhlops Braminus — FITZINGER 1826: 53
Typhlops braminus — CUVIER 1828: 103
Typhlops bramineus — SCHINZ 1834: 130 [unjustified emendation]
Typhlops russeli SCHLEGEL 1839
Typhlops Braminus — DUMÉRIL & BIBRON 1844: 309
Argyrophis truncatus GRAY 1845: 138
Argyrophis Bramicus — GRAY 1845: 138 [unjustified emendation]
Eryx bramicus — GRAY 1845: 138 [unjustified emendation]
Tortrix Bramicus — GRAY 1845: 279
Onychocephalus capensis SMITH 1846
Typhlops brame — DUMÉRIL 1853: 423
Typhlops Russellii — JERDON 1853: 527
Typhlops capensis — PETERS 1854: 621
Argyrophis bramicus [sic] — KELAART 1854: 137
Argyrophis Braminus — BLYTH 1856: 712
Argyrophis bramicus — TENNENT 1859: 203
Onychocephalus capensis — PETERS 1860: 83
Ophthalmidium tenue HALLOWELL 1861: 497
Typhlops (Typhlops) braminus — JAN 1863: 11
Typhlops (Typhlops) inconspicuus JAN 1863: 11
Typhlops accedens — JAN 1864
Typhlops braminus — JAN 1864
Tortrix russellii — GÜNTHER 1864: 175
Typhlops accedens JAN & SORDELLI 1864: 3
Typhlops capensis — PETERS 1869: 139
Typhlops avoterminatus GÜNTHER 1870: 139
Typhlops inconspicuus — BOETTGER 1879: 459
Typhlops bramini — BOETTGER 1881: 650
Typhlops (Typhlops) euproctus BOETTGER 1882: 479
Typhlops bramino — BOETTGER 1882: 479 [unjustified emendation]
Typhlops sp. (braminus var.?) — MÜLLER 1887: 259
Typhlops brahminus — PHIPSON 1888:49 [unjustified emendation]
Typhlops braminus — BOULENGER 1893: 16
Typhlops euproctus — BOULENGER 1893: 16
Tortrix russellii — BOULENGER 1893:16 [unjustified emendation]
Typhlops russellii — BOULENGER 1893:16 [unjustified emendation]
Cyphlops braminus — CARDEW 1897:592 [error typographicus]
Typhlops bramimus — MEHELY 1897:62 [error typographicus]
Typhlops bramina — HASSERT 1903:25 [unjustified emendation]
Typhlops braminus arenicola ANNANDALE 1906:192
Typhlops limbrickii ANNANDALE 1906
Typhlops braminus — STEJNEGER 1907: 260
Glauconia braueri STERNFELD 1910: 69
Tyhlops braminus — OSHIMA 1910: 186 [error typographicus]
Typhlops braueri — BOULENGER 1910: 29
Typhlopidae [sic] braminus — ROUX 1911
Typhlops braminis, MONTAGUE 1914: 643. [error typographicus]
Typhlops braminus — DE ROOIJ 1917: 5
Typhlops fletcheri WALL 1919: 556, pl. 1
Typhlops bramini — HOLTZINGER-TENEVER 1920:102. [unjustified emendation]
Typhlops braminus — BOULENGER 1920
Typhlops braminus — TAYLOR 1922
Typhlops sp. ? — GHARPUREY 1927: 224
Typhlops braminus braminus — MERENS 1930: 278
Typhlops sp. — PURACHATRA 1930: 60
Typhlops braminius — MAKI 1931
Typhlops braminius — GHARPUREY 1932: 272 [error typographicus]
Typhlopidae sp. — GHARPUREY 1935: 272
Typhlopus braminus — NAKAMURA 1938: 192 [error typographicus]
Typhlops braminus — SMITH 1943
Typhlops inconspiuous — LOVERIDGE 1957: 244 [unjustified emendation]
Typhlops khoratensis TAYLOR 1962: 248
Typhlops koratensis — TAYLOR 1965: 645 [error typographicus]
Ramphotyphlops braminus — ROBB 1966: 675
Typhlops pseudosaurus DRYDEN and TAYLOR 1969
Typhlops braminus — DEUVE 1970
Ehyx braminus — AGRAWAL 1970: 41 [error typographicus]
Typhlops braminae — BROWN & ALCALA [unjustified emendation]
Typhlops braminueus — GALLAGHER 1971: 27 [error typographicus]
Typhlops cf. braminus — CHEKE 1973: 469
Typhlina (?) bramina — MCDOWELL 1974: 22
Typhlina bramina — BROADLEY 1974: 24–26
Typhlops Graminas — AGARWAL 1979: 9 [error typographicus]
Typhlina braminus — MCDOWELL 1974
Typhlina bramina bramina — AUFFENBERG 1980: 113
Ramphotyphlops braminus — NUSSBAUM 1980
Ramphotyphlops bramina — WILSON & PORRAS 1983: 55
Ramphotyphlopus braminus — GIBBONS 1985: 139 [error typographicus]
Ramphotyphlops braminus — CONANT & COLLINS 1991: 144
Ramphotyphlops braminus — GLAW & VENCES 1994: 350
Ramphotyphlops braminus — MANTHEY & GROSSMANN 1997: 432
Ramphotyphlops braminus — COX et al. 1998: 12
Ramphotyphlops braminus — MCDIARMID, CAMPBELL & TOURÉ 1999: 59
Typhlops khoratensis — MCDIARMID, CAMPBELL & TOURÉ 1999: 106
Typhlops braminus — LAZELL 2002
Ramphotyphlops braminus — D’CRUZE et al. 2007
Ramphotyphlops braminus — BROADLEY & WALLACH 2009
Ramphotyphlops braminus — COX et al. 2012: 7
Typhlops khoratensis — COX et al. 2012: 25
Ramphotyphlops bramius — JEAN-BAPTISTE 2013 (in error)
Ramphotyphlops braminus — WALLACH et al. 2014: 626
Indotyphlops braminus — HEDGES et al. 2014
Ramphotyphlops braminus — COGGER 2014: 798
Indotyphlops khoratensis — HEDGES et al. 2014
Typhlops khoratensis — CHAN-ARD et al. 2015: 148
Ramphotyphlops braminus — CHAN-ARD et al. 2015: 145
Ramphotyphlops barminus — BALOUCHA et al. 2016 (in error)
Ramphotyphlops braminus — SUPSUP et al. 2016
Ramphotyphlops braminus — SANGUILA et al. 2016
Indotyphlops braminus — SPAWLS et al. 2018: 354
Virgotyphlops braminus — WALLACH 2020
Indotyphlops braminus — SPEYBROECK et al. 2020
Virgotyphlops braminus — CRUZ-ELIZALDE et al. 2022
Indotyphlops braminus — WICKRAMASINGHE et al. 2022
Rhamphotyphlops braminus — MENESES et al. 2022 (in error)
Ramphotyphops braminus — ICLEI 2023 (in error)
Virgotyphlops braminus — O’SHEA et al. 2023
Virgotyphlops braminus — SUNYER & MARTÍNEZ-FONSECA 2023 
DistributionAfrica: Egypt [HR 30: 236; HR 32: 11], Libya, Zanzibar, Tanzania, Mozambique, Republic of South Africa, Somalia, Cameroon, Benin, Ivory Coast, Senegal, Mauritania, Gabon, Central African Republic, Democratic Republic of the Congo (Trape 2019), Madagascar, Comoro Islands (Mayotte, Mohéli, Anjouan), Nossi Be = Nosy Bé, Mascarenes, Seychelles, Mauritius, Reunion, Rodrigues, Madeira (J. Jesus, pers. comm. 13 Dec 2012), Equatorial Guinea

Middle East: Saudi Arabia (introduced), Oman (introduced), United Arab Emirates (UAE), Kuwait, Iran, Iraq.

Asia: Vietnam, Laos, Thailand [HR 32: 279], Myanmar (= Burma), Cambodia, Malaysia (Pulau Tioman, Johor: Pulau Sibu, Pulau Besar etc.), Indonesia (Borneo, Sulawesi, Komodo, Sumatra, Java, Bali), Solomon Islands [McCoy 2000], Timor-Leste, S China (Hainan, Hunan, Guangdong: Nan Ao Island, Hong Kong, Sichuan/Chongqing, Fujian, Yunnan), Taiwan (incl. Lanyu), Japan (Ryukyu islands, incl. Senkaku), India (Madhya Pradesh, Himachal Pradesh, Odisha, Jammu and Kashmir, Kerala, Maharashtra, Telangana, Arunachal Pradesh [A. Captain, pers. Comm.], Punjab [Dino Aulakh, pers. comm.], Gujarat, Tamil Nadu, Mizoram), Andaman Islands, Nicobar Islands, Sri Lanka, Pakistan, Nepal, Bhutan, Philippines (Palawan: Calamian Islands, Panay, Luzon, Cebu, Camiguin Sur, etc.), New Guinea, Nauru, Palau, Singapore

Australia and Oceania: (Cook Islands, CKI, North Territory), New Caledonia, Toga Island, Vanuatu, Fiji, Micronesia, Guam (G. Rodda, pers. comm., 13 March 2016), French
Polynesia (Tahiti Island, Society Archipelago)

Europe: Portugal (Madeira Island), Spain (Canary Islands: Tenerife, Lanzarote, Gran Canaria, Fuerteventura, La Gomera), Italy

America: Mexico (introduced to Sinaloa, Querétaro, Coahuila, Michoacán, Aguascalientes [HR 32: 279], Jalisco, Baja California Sur, Morelos, Tamaulipas, Chiapas, Oaxaca, Hidalgo, Guanajuato, Nuevo León, Chihuahua, Veracruz, Zacatecas, Quintana Roo: Cozumel Island, Yucatán, Campeche, Nayarit, Puebla, Guerrero, Sonora, Durango), Guatemala (introduced), Belize ? (P. Stafford, pers. comm.), Honduras, Nicaragua

USA (introduced to Florida and Hawaii),
West Indies [HR 28: 210], Grand Cayman Islands [HR 34: 265], St. Martin, St. Barthélmy (BREUIL 2002), St. Kitts
Elevation 0-2000 m, Guadeloupe (La Désirade), Introduced to Barbados.

Caribbean: Introduced to Cuba, Saba (Dutch Caribbean), Bahamas (Abaco Island)

Type locality: Vizagapatam [Visakhapatnam], India.

khoratensis: C/E Thailand (Khorat Plateau area). Type locality: “Muak Lek, Friendship Highway, Sara Buri Province, Thailand”  
ReproductionOviparous. Seems to be the only parthenogenetic snake species known so far (another one may be Acrochordus arafurae). McDowell (1974) was the first to suggest that Ramphotyphlops braminus was an all-female species based upon a sample of 114 mostly Asian and Indonesian specimens, none of which was male. Reports of “male” Ramphotyphlops braminus (Wall, 1918, 1921, one from Assam, India; List, 1958, two from Sri Lanka) most likely stem from misidentifications of braminus-like members of the Typhlops pammeces group (Wallach 2009). Wynn et al. (1987) and Ota et al. (1991) demonstrated that R. braminus is a triploid parthenogenetic species. 
TypesLectotype: iconotype, Russell 1796, plate 43
Holotype: FMNH 178263 [khoratensis] 
DiagnosisDiagnosis: First, I. braminus is found in regions where there is no other blind snake.
Second, the condition of the nasal sutures can separate I. braminus from all but five scolecophidian species. In I. braminus the inferior or infranasal suture (INS) from the nostril (N) curves downward and contacts the preocular (PO), rather than the first or second supralabial (SL1 or SL2) as in most other typhlopids, whereas the superior or supranasal suture (SNS) extends upwards and backwards onto the top of the snout and makes contact with the rostral (R) to completely divide the nasal shield (rather than not extending onto dorsum of snout and not contacting the rostral as in most other typhlopids) (Fig. 1). The supranasal suture extends onto the dorsum of the snout and contacts the rostral in only three other species (Anilios erycinus of New Guinea, A. ganei and A. ligatus of western Australia) and the infranasal suture contacts the preocular in only two other species (Indotyphlops lankaensis and I. violaceus of Sri Lanka). Among these five problematic species, they differ from Indotyphlops braminus in having the following distinctions: Anilios erycinus of New Guinea has the INS contacting SL1 (vs. PO), more postoculars (2 vs. 1), a narrower rostral (0.15 vs. 0.30–0.40 head width), and is larger in size (LOA > 230 mm vs. < 200 mm); A. ganei of Australia has the INS in contact with SL2 (vs. PO), SNS invisible dorsally (vs. visible), more scale rows (24 vs. 20), more middorsals (> 440 vs. < 310), and a larger body size (> 235 mm vs. < 200 mm); A. micromma of Australia has the INS contacting SL 2 (vs. PO), fewer scale rows (18 vs. 20), more middorsals (493 vs. < 370), more postoculars (3 vs. 1), and a broader rostral (0.5 vs. 0.3–0.4 head width); I. lankaensis of Sri Lanka has the SNS invisible dorsally (vs. visible), fewer middorsals (< 261 vs. > 261), and a thicker body (L/W < 35 vs. > 30); I. violaceus of Sri Lanka has the SNS invisible dorsally (vs. visible). With the exceptions just noted for the above five species, the nasal suture condition of I. braminus will differentiate it from all of the other 475 scolecophidian species.
Third, the head of I. braminus exhibits prominent and distinct subcutaneous sebaceous glands arranged in rows beneath each head shield (also present in the I. pammeces species group). These gland lines run parallel to and along the margins of the anterior head shields. A unique situation involves the gland lines of the prenasal or anterior nasal (N1) shield and the median rostral (R) shield. As the gland line on the lower or anterior rostral (ARL) ascends towards the top of the snout it is interrupted by the supranasal gland line (SNL) on the prenasal (N1) shield, which is confluent with the gland line along the posterior rostral (PRL). A close inspection of the head will clearly reveal this condition and it is unique among all scolecophidians. This one feature will positively identify I. braminus. Another unusual gland line feature, also visible without the aid of a microscope, is that the posterior rostral (R) and supranasal (N2) shields have a bowlike gland line (CGL) that is convex rather than concave and parallel to the posterior rostral border. This unique feature on the top of the head just anterior to the level of the eyes is distinct and easily visible. These two features, 1) a continuous line of whitish glands extending along the nasal suture (SNL) and posterior rostral (PRL), and 2) a convexly curved gland line (CGL) across the posterior rostral (R) and supranasal (N2) shields, will positively identify I. braminus in an examination or photographs (figs. 1–2).
Fourth, in spite of the fact that the colour in I. braminus ranges from jet black through all shades of brown to tan and pink, it has a lighter colored snout with a white chin, cloacal region, and tail tip (from Wallach 2020, see figures there). 
CommentDescription in BROADLEY & WALLACH 2009.

Synonymy: following Wallach 2009 (who lists many more typos and variations).

Type species: Eryx braminus DAUDIN 1803 is the type species of the genus Virgotyphlops Wallach 2020 which was erected based on its parthenogenetic mode of reproduction (which may or may not be justified, given that other genera have mixed modes of reproduction, e.g. Darevskia or Aspidoscelis).

Distribution: Not listed for Togo by Segniagbeto et al. 2012. See map in Burriel-Carranza et al. 2019 for map in UAE. For a map see Sindaco et al. 2013. For a map with localities in Equatorial Guinea see SÁNCHEZ-VIALAS et al. 2022.

Phylogenetics: I. braminus as currently diagnosed appears to fall into several genetically distinct clades which may be separate species (Wickramasinghe et al. 2022). The monophyly of Indotyphlops including braminus was also confirmed by Sidharthan & Karanth 2021, hence we reject the establishment of a new genus, Virgotyphlops.

Karyotype: triploid karyotype of 3n=42 with seven macro- and seven microtriplets and NORs on a microtriplet (Mezzasalma et al. 2016).

Similar species: I. lankaensis, I. violaceus (both Sri Lanka).

Genome: Khedkar et al. 2022.

Transcriptome: Zhu et al. 2023. 
EtymologyNamed after the Latinized form of the word “Brahmin” = a caste among Hindus. 
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