diff --git a/data/69/7A/F1/697AF1B87B5E5EFABEF602ADD4E48B4D.xml b/data/69/7A/F1/697AF1B87B5E5EFABEF602ADD4E48B4D.xml new file mode 100644 index 00000000000..7002ebdfe19 --- /dev/null +++ b/data/69/7A/F1/697AF1B87B5E5EFABEF602ADD4E48B4D.xml @@ -0,0 +1,6938 @@ + + + +A new four-pored Amphisbaena Linnaeus, 1758 (Amphisbaenia, Amphisbaenidae) from the north of Espinhaço Mountain Range, Brazil + + + +Author + +Ribeiro, Síria +0000-0002-2301-7089 +Universidade Federal do Oeste do Pará, Programa de Pós-Graduação em Biodiversidade, 68040 - 255, Santarém, Pará, Brazil + + + +Author + +Santos Jr, Alfredo P. +0000-0002-2829-718X +Universidade Federal do Oeste do Pará, Programa de Pós-Graduação em Biodiversidade, 68040 - 255, Santarém, Pará, Brazil + + + +Author + +Martins, Isabelly G. +https://orcid.org/0009-0002-4420-5713 +Universidade Federal do Oeste do Pará, Programa de Pós-Graduação em Biodiversidade, 68040 - 255, Santarém, Pará, Brazil + + + +Author + +Oliveira, Elaine C. S. +Laboratório de Ecologia e Comportamento Animal, Universidade Federal do Oeste do Pará, 68040 - 255, Santarém, Pará, Brazil + + + +Author + +Graboski, Roberta +0000-0002-9123-4819 +Instituto Chico Mendes de Conservação da Biodiversidade, Coordenação Regional Oeste do Pará, 68040 - 000, Santarém, Pará, Brazil + + + +Author + +Barbosa Da Silveira, Thiago +School of Zoology, Faculty of Life Sciences, the Steinhardt Museum of Natural History, Tel-Aviv University, Tel-Aviv, Israel + + + +Author + +Benício, Matheus H. M. +Tecsan – Tecnologia e Saneamento Ltda – Rua Dr. José Peroba, 275, Ed. Metrópolis Empresarial, Sala 201 / 202, Stiep, 41770 - 235, Salvador, Bahia, Brazil + + + +Author + +Vaz-Silva, Wilian +0000-0001-6235-5331 +Bahia Mineração S / A, Rodovia BA 156, s / n, Projeto Pedra de Ferro, 46400 - 000, Caetité, Bahia, Brazil + +text + + +ZooKeys + + +2024 + +2024-09-24 + + +1213 + + +1 +27 + + + +journal article +10.3897/zookeys.1213.122265 +77F68F11-081E-4693-94BD-F211FFCFED62 + + + + + +Amphisbaena amethysta + +sp. nov. + + + + +Figs 1 +, +2 +, +3 +, +4 + + + + +Type material. + + + + +Holotype + +: • +male +; + +CEPB +2311 + +; municipality of +Caetité +, state of +Bahia +, +Brazil +; [ + +14 ° 21 ' 31 " S +, +42 ° 32 ' 19 " W + +; + +1012 m +above sea level + +(a. s. l.)]; collected on + +1 November 2022 + +by +Faunal Rescue Team Tecsam +( +F. Santos +, +P. Belufi +, and +G. Nascimento +) + +. + +Paratypes + +: • All from +Caetité +, +Bahia +, +Brazil +; collected by +Faunal Rescue Team Tecsam +( +R. Assunção +, +A. Hirota +, +T. Silveira +, +F. Santos +, +P. Belufi +, and +G. Nascimento +) • + +Female +; + +CEPB +2301 + +; ( + +14 ° 21 ' 53 " S +, +42 ° 32 ' 20 " W + +; + +1013 m +a. s. l. + +); + +8 June 2022 + + +• + +Male +; + +CEPB +2302 + +; ( + +14 ° 01 ' 16 " S +, +42 ° 31 ' 06 " W + +; + +1082 m +a. s. l. + +); + +13 October 2021 + + +• + +Female +; + +CEPB +2303 + +; ( + +14 ° 21 ' 50 " S +, +42 ° 32 ' 20 " W + +; + +1013 m +a. s. l. + +); + +26 May 2022 + + +• + +Female +; + +CEPB +2308 + +; ( + +14 ° 19 ' 49 " S +, +42 ° 32 ' 44 " W + +; + +1011 m +a. s. l. + +); + +8 September 2022 + + +• + +Female +; + +CEPB +2327 + +; ( + +14 ° 19 ' 52 " S +, +42 ° 32 ' 42 " W + +; + +1011 m +a. s. l. + +); + +1 October 2022 + + +• + +Female +; + +CEPB +2331 + +; ( + +14 ° 19 ' 45 " S +, +42 ° 32 ' 45 " W + +; + +1011 m +a. s. l. + +); + +30 August 2022 + + +• + +Male +; + +CEPB +2346 + +; ( + +14 ° 22 ' 08 " S +, +42 ° 32 ' 17 " W + +; + +923 m +a. s. l. + +); + +6 October 2022 + + +• + +Male +; + +CEPB +2379 + +; ( + +14 ° 21 ' 27 " S +, +42 ° 32 ' 20 " W + +; + +1033 m +a. s. l. + +); + +26 January 2023 + + +• + +Female +; + +CEPB +2381 + +; ( + +14 ° 21 ' 27 " S +, +42 ° 32 ' 20 " W + +; + +1033 m +a. s. l. + +); + +26 January 2023 + + +. + + + + +Referred specimens. + + +All from +Caetité +, +Bahia +, +Brazil +; collected by +Faunal Rescue Team Tecsam +( +R. Assunção +, +A. Hirota +, +T. Silveira +, +F. Santos +, +P. Belufi +, and +G. Nascimento +) • + +Female +; + +CEPB +2298 + +; ( + +14 ° 21 ' 53 " S +, +42 ° 32 ' 20 " W + +; + +1013 m +a. s. l. + +); + +8 June 2022 + + +• + +Female +; + +CEPB +2299 + +; ( + +14 ° 21 ' 53 " S +, +42 ° 32 ' 20 " W + +; + +1013 m +a. s. l. + +); + +8 June 2022 + + +• + +Male +; + +CEPB +2300 + +; ( + +14 ° 21 ' 51 " S +, +42 ° 32 ' 21 " W + +; + +1011 m +a. s. l. + +); + +25 May 2022 + + +• + +Female +; + +CEPB +2304 + +; ( + +14 ° 21 ' 53 " S +, +42 ° 32 ' 20 " W + +; + +1013 m +a. s. l. + +); + +8 June 2022 + + +• + +Female +; + +CEPB +2305 + +; ( + +14 ° 21 ' 53 " S +, +42 ° 32 ' 20 " W + +; + +1013 m +a. s. l. + +); + +8 June 2022 + + +• + +Female +; + +CEPB +2306 + +; ( + +14 ° 21 ' 53 " S +, +42 ° 32 ' 20 " W + +; + +1013 m +a. s. l. + +); + +13 August 2022 + + +• + +Male +; + +CEPB +2307 + +; ( + +14 ° 21 ' 23 " S +, +43 ° 32 ' 07 " W + +; + +972 m +a. s. l. + +); + +23 November 2022 + + +• + +Undetermined +sex; + +CEPB +2309 + +; ( + +14 ° 19 ' 46 " S +, +42 ° 43 ' 20 " W + +; + +1011 m +a. s. l. + +); + +1 September 2022 + + +• + +Female +; + +CEPB +2310 + +; ( + +14 ° 19 ' 55 " S +, +42 ° 32 ' 44 " W + +; + +1011 m +a. s. l. + +); + +8 October 2022 + + +• + +Male +; + +CEPB +2312 + +; ( + +14 ° 19 ' 56 " S +, +42 ° 32 ' 43 " W + +; + +1011 m +a. s. l. + +); + +12 October 2022 + + +• + +Female +; + +CEPB +2313 + +; ( + +14 ° 22 ' 08 " S +, +42 ° 32 ' 17 " W + +; + +9215 m +a. s. l. + +); + +6 October 2022 + + +• + +Female +; + +CEPB +2314 + +; ( + +14 ° 22 ' 08 " S +, +42 ° 32 ' 17 " W + +; + +925 m +a. s. l. + +); + +6 October 2022 + + +• + +Female +; + +CEPB +2315 + +; ( + +14 ° 22 ' 08 " S +, +42 ° 32 ' 17 " W + +; + +925 m +a. s. l. + +); + +6 October 2022 + + +• + +Female +; + +CEPB +2316 + +; ( + +14 ° 19 ' 47 " S +, +42 ° 32 ' 43 " W + +; + +1011 m +a. s. l. + +); + +1 September 2022 + + +• + +Female +; + +CEPB +2317 + +; ( + +14 ° 19 ' 55 " S +, +43 ° 32 ' 44 " W + +; + +1011 m +a. s. l. + +); + +1 October 2022 + + +• + +Female +; + +CEPB +2318 + +; ( + +14 ° 19 ' 46 " S +, +42 ° 32 ' 43 " W + +; + +1011 m +a. s. l. + +); + +1 September 2022 + + +• + +Female +; + +CEPB +2319 + +; ( + +14 ° 20 ' 37 " S +, +42 ° 32 ' 13 " W + +; + +1076 m +a. s. l. + +); + +3 August 2022 + + +• + +Female +; + +CEPB +2320 + +; ( + +14 ° 20 ' 38 " S +, +42 ° 32 ' 12 " W + +; + +1013 m +a. s. l. + +); + +2 June 2022 + + +• + +Female +; + +CEPB +2321 + +; ( + +14 ° 19 ' 53 " S +, +42 ° 32 ' 43 " W + +; + +1011 m +a. s. l. + +); + +29 September 2022 + + +• + +Female +; + +CEPB +2322 + +; ( + +14 ° 14 ' 28 " S +, +42 ° 32 ' 47 " W + +; + +842 m +a. s. l. + +); + +17 September 2022 + + +• + +Female +; + +CEPB +2323 + +; ( + +14 ° 19 ' 49 " S +, +42 ° 32 ' 17 " W + +; + +1062 m +a. s. l. + +); + +13 August 2022 + + +• + +Male +; + +CEPB +2324 + +; ( + +14 ° 19 ' 48 " S +, +42 ° 32 ' 43 " W + +; + +1011 m +a. s. l. + +); + +6 September 2022 + + +• + +Female +; + +CEPB +2325 + +; ( + +14 ° 19 ' 55 " S +, +42 ° 32 ' 44 " W + +; + +1011 m +a. s. l. + +); + +1 October 2022 + + +• + +Male +; + +CEPB +2326 + +; ( + +14 ° 20 ' 52 " S +, +43 ° 32 ' 16 " W + +; + +1053 m +a. s. l. + +); + +13 August 2022 + + +• + +Female +; + +CEPB +2328 + +; ( + +14 ° 21 ' 53 " S +, +42 ° 32 ' 22 " W + +; + +1011 m +a. s. l. + +); + +28 September 2022 + + +• + +Male +; + +CEPB +2329 + +; ( + +14 ° 19 ' 47 " S +, +42 ° 32 ' 43 " W + +; + +1011 m +a. s. l. + +); + +1 September 2022 + + +• + +Male +; + +CEPB +2330 + +; ( + +14 ° 19 ' 55 " S +, +43 ° 32 ' 44 " W + +; + +1011 m +a. s. l. + +); + +1 October 2022 + + +• + +Male +; + +CEPB +2332 + +; ( + +14 ° 20 ' 50 " S +, +42 ° 32 ' 16 " W + +; + +1057 m +a. s. l. + +); + +13 August 2022 + + +• + +Female +; + +CEPB +2333 + +; ( + +14 ° 14 ' 28 " S +, +42 ° 32 ' 47 " W + +; + +842 m +a. s. l. + +); + +17 September 2022 + + +• + +Female +; + +CEPB +2334 + +; ( + +14 ° 19 ' 46 " S +, +42 ° 32 ' 43 " W + +; + +1011 m +a. s. l. + +); + +1 September 2022 + + +• + +Female +; + +CEPB +2336 + +; ( + +14 ° 15 ' 11 " S +, +45 ° 32 ' 16 " W + +; + +987 m +a. s. l. + +); + +3 August 2022 + + +• + +Female +; + +CEPB +2337 + +; ( + +14 ° 20 ' 38 " S +, +45 ° 32 ' 12 " W + +; + +1076 m +a. s. l. + +); + +3 August 2022 + + +• + +Female +; + +CEPB +2338 + +; ( + +14 ° 21 ' 31 " S +, +45 ° 32 ' 16 " W + +; + +1010 m +a. s. l. + +); + +13 August 2022 + + +• + +Female +; + +CEPB +2339 + +; ( + +14 ° 14 ' 28 " S +, +42 ° 32 ' 47 " W + +; + +842 m +a. s. l. + +); + +17 September 2022 + + +• + +Male +; + +CEPB +2356 + +; ( + +14 ° 21 ' 27 " S +, +42 ° 32 ' 21 " W + +; + +1033 m +a. s. l. + +); + +1 November 2022 + + +• + +Male +; + +CEPB +2379 + +; ( + +14 ° 21 ' 27 " S +, +42 ° 32 ' 20 " W + +; + +1033 m +a. s. l. + +); + +26 January 2023 + + +• + +Female +; + +CEPB +2380 + +; ( + +14 ° 21 ' 27 " S +, +42 ° 32 ' 20 " W + +; + +1033 m +a. s. l. + +); + +26 January 2023 + + +• + +Female +; + +CEPB +2381 + +; ( + +14 ° 21 ' 27 " S +, +42 ° 32 ' 20 " W + +; + +1033 m +a. s. l. + +); + +26 January 2023 + + +. + + + + +Diagnosis and comparisons with other south American amphisbaenians. + + + +Amphisbaena amethysta + +sp. nov. +is a medium-sized amphisbaenid ( +258 mm +maximum snout-vent length), and can be distinguished from its congeners by the following combination of characters (see details in Table +1 +): (1) snout convex in profile view, slightly compressed not keeled; (2) pectoral scales arranged in regular annuli; (3) four precloacal pores; (4) distinct cephalic shields; (5) 185–199 dorsal half-annuli; (6) 13–16 caudal annuli; (7) conspicuous autotomic site between 4 +th +– 6 +th +caudal annuli; (8) 16–21 dorsal and ventral segments at midbody; (9) 3 / 3 supralabials; (10) 3 / 3 infralabials; and (11) smooth and rounded tail tip. + + + +Amphisbaena amethysta + +sp. nov. +differs from + +Amphisbaena acrobeles +( +Ribeiro, Castro-Mello & Nogueira, 2009 +) + +, + +A. bilabialata +(Stimson, 1972) + +, + +A. kingi +(Bell, 1833) + +, + +A. anomala + +, + +Mesobaena huebneri +Mertens, 1925 + +; + +M. rhachicephala +Hoogmoed, Pinto, Rocha & Pereira, 2009 + +; and all + +Leposternon +species + +, mainly in having the snout convex in profile view, slightly compressed not keeled ( +vs +snout hardly compressed forming a sharp and prominent keel or snout depressed shovel-like). Differs still from + +A. anomala + +and all + +Leposternon +species + +by having pectoral scales arranged in regular annuli ( +vs +pectoral scales with an irregular form, and dermal annuli not regularly arranged). + + + +Amphisbaena amethysta + +sp. nov. +presents four precloacal pores, differing from all + +Amphisbaena +species + +except + +A. acangaoba +Ribeiro, Gomides & Costa, 2020 + +, + +A. alba + +, + +A. albocingulata +Boettger, 1885 + +, + +A. angustifrons +Cope, 1861 + +, + +A. arda + +, + +A. arenaria + +, + +A. arenicola +Perez & Borges-Martins, 2019 + +, + +A. bahiana + +, + +A. bakeri +Stejneger, 1904 + +, + +A. barbouri +Gans & Alexander, 1962 + +, + +A. bedai +( +Vanzolini, 1991 +) + +, + +A. bolivica +Mertens, 1929 + +, + +A. borellii +Peracca, 1897 + +, + +A. brasiliana +(Gray, 1865) + +, + +A. caeca +Cuvier, 1829 + +, + +A. camura +Cope, 1862 + +, + +A. carlgansi +Thomas & Hedges, 1998 + +, + +A. carioca +Rocha, Barros-Filho & Sluys, 2023 + +, + +A. carvalhoi + +, + +A. caudalis +Cochron, 1928 + +, + +A. cayemite +Thomas & Hedges, 2006 + +, + +A. cegei +Montero, Sáfadez, Álvarez, 1997 + +, + +A. cubana +Gundlach & Peters, 1879 + +, + +A. cuiabana +( +Strüssmann & Carvalho, 2001 +) + +, + +A. cunhai +Hoogmoed & Ávila-Pires, 1991 + +, + +A. darwini + +, + +A. elbakyanae +Torres-Ramírez, Angarita-Sierra & Vargas-Ramírez, 2021 + +, + +A. fenestrata +(Cope, 1861) + +, + +A. frontalis + +, + +A. gonavensis +Gans & Alexander, 1962 + +, + +A. gracilis +Strauch, 1881 + +, + +A. hastata + +, + +A. heathi + +, + +A. hogei +Vanzolini, 1950 + +, + +A. hoogmoedi +Oliveira, Vaz-Silva, Santos-Jr, Graboski, Teixeira Jr, Dal Vechio & Ribeiro, 2018 + +, + +A. hyporissor +Thomas, 1965 + +, + +A. innocens +Weinland, 1862 + +, + +A. kingi + +, + +A. kraoh +(Vanzolini, 1971) + +, + +A. leali +Thomas & Hedges, 2006 + +, + +A. lumbricalis + +, + +A. manni +Barbour, 1914 + +, + +A. medemi +Gans & Mathers, 1977 + +, + +A. metallurga + +, + +A. mongoyo + +, + +A. munoai +Klappenbach, 1960 + +, + +A. myersi +Hoogmoed, 1989 + +, + +A. nana +Perez & Borges-Martins, 2019 + +, + +A. nigricauda +Gans, 1966 + +, + +A. occidentalis +Cope, 1875 + +, + +A. pericensis +Noble, 1921 + +, + +A. plumbea +Gray, 1872 + +, + +A. polygrammica +Werner, 1901 + +, + +A. prunicolor +(Cope, 1885) + +, + +A. ridleyi +Boulenger, 1890 + +, + +A. rozei + +Lancini, 1963, + +A. sanctaeritae +Vanzolini, 1994 + +, + +A. saxosa +(Castro-Mello, 2003) + +, + +A. schmidti +Gans, 1964 + +, + +A. slateri +Boulenger, 1907 + +, + +A. slevini +Schmidt, 1936 + +, + +A. spurrelli +Boulenger, 1915 + +, + +A. steindachneri +Strauch, 1881 + +, + +A. supernumeraria +Mott, Rodrigues & Dos Santos, 2009 + +, + +A. talisiae +Vanzolini, 1995 + +, + +A. tyaraju +, +Perez & Borges-Martins, 2019 + +, + +A. townsendi +Stejneger, 1911 + +, + +A. tragorrhectes +Vanzolini, 1971 + +, + +A. uroxena + +, + +A. vanzolinii +Gans, 1963 + +, + +A. vermicularis + +, and + +A. xera +Thomas, 1966 + +. + + + +Amphisbaena amethysta + +sp. nov. +differs from + +Amphisbaena +species + +with four precloacal pores mainly by following combination of meristic characters (Table +1 +): cephalic shield distinct (vs frontals and parietals shields not distinct in + +A. supernumeraria + +, ocular and second supralabial not distinct in + +A. cubana + +); snout slightly compressed (vs hard compressed in + +A. kingi + +and rounded in all other species, except + +A. kraoh + +, + +A. brasiliana + +, + +A. bahiana + +, + +A. bedai + +, and + +A. saxosa + +); 185–199 dorsal half-annuli (vs <170 annuli in + +A. cayemite + +and> 200 annuli in + +A. acangaoba + +, + +A. arda + +, + +A. arenaria + +, + +A. bahiana + +, + +A. bakeri + +, + +A. barbouri + +, + +A. bedai + +, + +A. borellii + +, + +A. brasiliana + +, + +A. caeca + +, + +A. carlgansi + +, + +A. carvalhoi + +, + +A. cayemite + +, + +A. cuiabana + +, + +A. cunhai + +, + +A. elbakyanae + +, + +A. fenestrata + +, + +A. frontalis + +, + +A. gonavensis + +, + +A. gracilis + +, + +A. hastata + +, + +A. hoogmoedi + +, + +A. kingi + +, + +A. kraoh + +, + +A. lumbricalis + +, + +A. manni + +, + +A. medemi + +, + +A. mongoyo + +, + +A. munoai + +, + +A. myersi + +, + +A. nigricauda + +, + +A. occidentalis + +, + +A. plumbea + +, + +A. polygrammica + +, + +A. rozei + +, + +A. sanctaeritae + +, + +A. saxosa + +, + +A. slevini + +, + +A. spurrelli + +, + +A. steindachneri + +, + +A. supernumeraria + +, + +A. talisiae + +, + +A. tyaraju + +, + +A. townsendi + +, + +A. uroxena + +, + +A. vanzolinii + +, + +A. vermicularis + +, and + +A. xera + +); 13–16 caudal annuli (vs> 16 annuli in + +A. albocingulata + +, + +A. arda + +, + +A. arenaria + +, + +A. arenicola + +, + +A. bedai + +, + +A. bolivica + +, + +A. borellii + +, + +A. carvalhoi + +, + +A. caudalis + +, + +A. cegei + +, + +A. cuiabana + +, + +A. cunhai + +, + +A. darwini + +, + +A. elbakyanae + +, + +A. frontalis + +, + +A. gracilis + +, + +A. hastata + +, + +A. hoogmoedi + +, + +A. leali + +, + +A. lumbricalis + +, + +A. manni + +, + +A. medemi + +, + +A. metallurga + +, + +A. mongoyo + +, + +A. munoai + +, + +A. myersi + +, + +A. nana + +, + +A. nigricauda + +, + +A. occidentalis + +, + +A. polygrammica + +, + +A. prunicolor + +, + +A. rozei + +, + +A. sanctaeritae + +, + +A. saxosa + +, + +A. schmidti + +, + +A. slateri + +, + +A. slevini + +, + +A. spurrelli + +, + +A. steindachneri + +, + +A. supernumeraria + +, + +A. talisiae + +, + +A. tyaraju + +, + +A. townsendi + +, + +A. tragorrhectes + +, and + +A. vermicularis + +); conspicuous autotomic site between 4 +th +– 6 +th +caudal annuli (vs absent in + +A. acangaoba + +, + +A. alba + +, + +A. angustifrons + +, + +A. bakeri + +, + +A. barbouri + +, + +A. brasiliana + +, + +A. carioca + +, + +A. carlgansi + +, + +A. cayemite + +, + +A. cunhai + +, + +A. fenestrata + +, + +A. gonavensis + +, + +A. hastata + +, + +A. hoogmoedi + +, + +A. innocens + +, + +A. lumbricalis + +, + +A. occidentalis + +, + +A. ridleyi + +, + +A. saxosa + +, and + +A. uroxena + +; or from the 7 +th +caudal annuli in + +A. albocingulata + +, + +A. arda + +, + +A. arenicola + +, + +A. carvalhoi + +, + +A. cegei + +, + +A. cuiabana + +, + +A. darwini + +, + +A. heathi + +, + +A. kingi + +, + +A. metallurga + +, + +A. mongoyo + +, + +A. myersi + +, + +A. nana + +, + +A. prunicolor + +, + +A. schmidti + +, + +A. slateri + +, + +A. steindachneri + +, + +A. supernumeraria + +, + +A. talisiae + +, + +A. tyaraju + +, + +A. townsendi + +, + +A. tragorrhectes + +, and + +A. vanzolinii + +), smooth and rounded tail tip [vs bluntly ridged tail tip in + +A. bahiana + +; slightly dorsally compressed in + +A. acangaoba + +; with tubercles sit is depressed (compressed dorsoventrally) in + +A. uroxena + +; with modified conic pointed tubercles in + +A. caetitensis + +; and vertically keeled in + +A. borellii + +and + +A. steindachneri + +]; 16–21 dorsal segments at midbody (vs < +16 in + +A. albocingulata + +, + +A. arenaria + +, + +A. arenicola + +, + +A. barbouri + +, + +A. carlgansi + +, + +A. carvalhoi + +, + +A. cayemite + +, + +A. cuiabana + +, + +A. elbakyanae + +, + +A. fenestrata + +, + +A. heathi + +, + +A. hogei + +, + +A. elbakyanae + +, + +A. metallurga + +, + +A. munoai + +, + +A. nana + +, + +A. nigricauda + +, + +A. sanctaeritae + +, + +A. schmidti + +, + +A. slateri + +, + +A. slevini + +, + +A. supernumeraria + +, + +A. talisiae + +, + +A. tyaraju + +, + +A. tragorrhectes + +, + +A. uroxena + +, and + +A. vanzolinii + +); and> 21 segments in + +A. alba + +, + +A. arda + +, + +A. bolivica + +, + +A. camura + +, + +A. hoogmoedi + +, and + +A. kraoh + +); 16–21 ventral segments at midbody (vs < +16 in + +A. rozei + +, + +A. sanctaeritae + +, and + +A. tragorrhectes + +; and> 21 segments in + +A. alba + +, + +A. arda + +, + +A. bolivica + +, + +A. camura + +, + +A. cegei + +, + +A. gonavensis + +, + +A. hyporissor + +, + +A. kraoh + +, + +A. occidentalis + +, and + +A. townsendi + +); 3 / 3 supralabials (vs 2 / +2 in + +A. slevini + +and + +A. vanzolinii + +; and 4 / +4 in + +A. acangaoba + +, + +A. alba + +, + +A. angustifrons + +, + +A. arda + +, + +A. arenaria + +, + +A. bedai + +, + +A. camura + +, + +A. cayemite + +, + +A. occidentalis + +, + +A. plumbea + +, + +A. ridleyi + +, + +A. saxosa + +, + +A. townsendi + +, + +A. tragorrhectes + +, and + +A. vermicularis + +); and 3 / 3 infralabials (vs 2 / +2 in + +A. slevini + +and + +A. vanzolinii + +; and or 4 / +4 in + +A. occidentalis + +, + +A. plumbea + +, + +A. ridleyi + +, + +A. saxosa + +, + +A. townsendi + +, and + +A. tragorrhectes + +). + + + + + + +Diagnostic characters for the + +Amphisbaena +species + +with four precloacal pores. +PC +– precloacal pore, +CS +– cephalic shield, +DA +– dorsal half-annulus, +CA +– caudal annulus, +AS +– autotomic site, +DS +– dorsal segment, +VS +– ventral segment, +SL +– supralabial, +IL +– infralabial, +PM +– postmalar, +SN +– snout, +DG +– dorsal groove, +VG +– ventral groove, a – absent, +cp +– compressed, +n / a +– non-available data, p – present, +rd +– rounded, and +sc +– slightly compressed. Differences from the new species in bold. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
Species +PC + +CS + +DA + +CA + +AS + +DS + +VS + +SL + +IL + +PM + +SN + +DG + +VG +Reference
+ +A. amethysta + +sp. nov. +4distinct and paired185–19913–164–616–2116–2133a +sc +aapresent study
+ +A. acangaoba + +4–8distinct and paired +216–293 +13–17 +a +18 – 2418–24 +4 +3p +rd +aa +Ribeiro et al. (2020) +
+ +A. alba + +4–10distinct and paired198–24813–21a30 – 4235–4643p +rd +aa +Gans (1962 a +) +
+ +A. albocingulata + +4distinct and paired190–20424–278–912–1416–1833a + +rd + +aa +Perez et al. (2012) +
+ +A. angustifrons + +3–6distinct and paired190–21512–18 +a +20 – 3121–30 +4 +3p +rd +aa +Gans (1965 a +); +Gans and Diefenbach (1972) +
+ +A. arda + +4distinct and paired242308232343p +rd +aa +Rodrigues et al. (2003) +
+ +A. arenaria + +4distinct and paired285–30722–236–7 +12–14 +14–16 +4 +3a + +rd + +aaTeixeira Junior et al. (2016)
+ +A. arenicola + +4distinct and paired199–21620–228–912–1416–1833P +rd + +n / a + +n / a + +Perez and Borges-Martins (2019) +
+ +A. bahiana + +4distinct and paired +204–223 +14–164–512–1614–1633 +p + +sc +pp +Gans (1964 b +); +Dal Vechio et al. (2018) +
+ +A. bakeri + +4distinct and paired +239–255 +14–16 +a +14 – 1616–1733a + +rd + + +n / a + +n / a + +Gans and Alexander (1962) +
+ +A. barbouri + +4–6distinct and paired +226–240 +13–18a12 – 1416–1833a + +rd + + +n / a + +n / a + +Gans and Alexander (1962) +; +Thomas (1966) +
+ +A. bedai + +4distinct and paired272–28422–23618–2016–18 +4 +3 +p + +sc +aa +Oliveira et al. (2018) +
+ +A. bolivica + +4–6distinct and paired200–231 +18–26 +4–527–3626–3633p +rd +aa +Montero (1996) +
+ +A. borellii + +4distinct and paired239–24517–196–814–1616–2033 +n / a + + +rd + +aa +Gans (1964 b +) and +Oliveira et al. (2018) +
+ +A. brasiliana + +4distinct and paired +213–229 +11–15 +a +18 – 2118–2233 +p + +sc + +P +a +Oliveira et al. (2018) +
+ +A. caeca + +4–6distinct and paired +217–236 +13–184–813–1814–2033p +rd + +n / a + +n / a + +Gans and Alexander (1962) +
+ +A. caetitensis + +4distinct and paired186–19410–12a1619–2233a + +rd + +aa +Almeida et al. (2018) +
+ +A. camura + +4–6distinct and paired188–20714–194–528–4229–4643p +rd +aa +Gans (1965 a +); +Hoogmoed and Ávila-Pires (1991) +
+ +A. carioca + +4distinct and paired18621a10–12 +n / a +33p +rd +aa +Rocha et al. (2023) +
+ +A. carlgansi + +4distinct and paired +212–228 +14–16a1418–2033a + +rd + +aa +Thomas and Hedges (1998) +
+ +A. carvalhoi + +4distinct and paired231–24519–227–812–1416–1833a + +rd + +aa +Gans (1965 b +) +
+ +A. caudalis + +4distinct and paired193–217 +17–21 +612–1618–2133a + +rd + + +n / a + +n / a + +Gans and Alexander (1962) +; +Thomas and Hedges (1998) +
+ +A. cayemite + +4distinct and paired +150–164 +10–13a12 – 1318 +4 +3p +rd + +n / a + +n / a + +Thomas and Hedges (2006) +
+ +A. cegei + +4distinct and paired19822721–22 +22–23 +33a + +rd + + +n / a + +n / a + +Montero et al. (1997) +
+ +A. cubana + +4–6 + +ocular and second +SL +fused + +199–21810–166–912–1614–1833a + +rd + + +n / a + +n / a + +Gans and Alexander (1962) +; +Thomas (1966) +
+ +A. cuiabana + +4distinct and paired286–29218–209–10141633a +rd +pa +Strüssmann and Carvalho (2001) +
+ +A. cunhai + +4distinct and paired226–23925–26a14 – 1614–1833p +rd +aa +Hoogmoed and Ávila-Pires (1991) +
+ +A. darwini + +2–5distinct and paired174–19519–257–1013–1916–2333 +n / a + + +rd + +aa +Perez et al. (2012) +
+ +A. elbakyanae + +4distinct and paired245–25720–246–7 +13–15 +16–1833p +rd +paTorres-Ramírez et al. (2022)
+ +A. fenestrata + +4distinct and paired +236–249 +12–14a13 – 1414–1633p +rd + +n / a + +n / a + +Gans and Alexander (1962 +
+ +A. frontalis + +0–4distinct and paired235–27523–295–714–1814–1633a + +rd + +aa +Ribeiro-Júnior et al. (2022) +
+ +A. gonavensis + +4distinct and paired +214–225 +10–13 +a +15 – 18 +22–25 +33p +rd + +n / a + +n / a + +Gans and Alexander (1962) +
+ +A. gracilis + +4distinct and paired224–24821–246–713–1614–1733p +rd +paGonzales-Sponga and Gans (1971)
+ +A. hastata + +4distinct and paired266–27340a181633a + +rd + +ap +Vanzolini (1991) +
+ +A. heathi + +4distinct and paired183–187 +n / a +7 – 81218–2033a + +rd + +aa +Gans (1965 b +) +
+ +A. hogei + +4distinct and paired177–19115–194–7 +10–13 +14–1833p +rd +aa +Gans (1966) +; +Vanzolini (1950) +
+ +A. hoogmoedi + +4distinct and paired247–252277–822–2419–2133a +sc +aa +Oliveira et al. (2018) +
+ +A. hyporissor + +4distinct and paired199–22716–21514–18 +22–24 +33p +rd + +n / a + +n / a + +Thomas (1965) +; +Thomas and Hedges (2006) +
+ +A. innocens + +4distinct and paired185–22011–14 +a +13 – 1618–2133a + +rd + + +n / a + +n / a + +Gans and Alexander (1962) +; +Thomas and Hedges (1998) +
+ +A. kingi + +4distinct and paired +214–244 +15–23 +7 +12–1914–2233p +cp +aa +Gans and Rhodes (1964) +; +Oliveira et al. (2018) +
+ +A. kraoh + +4–6distinct and paired +270–281 +1552824–2733 +n / a + +sc + +n / a + +n / a + +Oliveira et al. (2018) +
+ +A. leali + +4distinct and paired188–206 +17–20 +614–1620–2233a + +rd + + +n / a + +n / a + +Thomas and Hedges (2006) +
+ +A. lumbricalis + +2–6distinct and paired225–24720–26a12 – 1616–2033 +n / a + + +rd + +aa +Vanzolini (1996) +
+ +A. manni + +4–9distinct and paired209–24317–225–712–1614–2033a + +rd + + +n / a + +n / a + +Gans and Alexander (1962) +
+ +A. medemi + +4distinct and paired230–23517–185–7161833a + +rd + +a +p + +Gans and Mather (1977) +
+ +A. metallurga + +2–4distinct and paired185–19923–257–912–1414–1633 +p +or a + + +rd + +aa +Costa et al. (2019) +
+ +A. mongoyo + +4distinct and paired2082510141633a + +rd + +aa +Teixeira Junior et al. (2019) +
+ +A. munoai + +4distinct and paired202–21818–235–9 +10–14 +13–1833p +rd +aa +Perez and Borges-Martins (2019) +
+ +A. myersi + +4distinct and paired221288161633 +n / a + + +rd + +aa +Hoogmoed (1989) +
+ +A. nana + +4distinct and paired195–21618–227–1012–1414–1733p +rd + +n / a + +n / a + +Perez and Borges-Martins (2019) +
+ +A. nigricauda + +4distinct and paired222–22619–246–9 +10 +1633a + +rd + +aa +Gans (1966) +
+ +A. occidentalis + +4distinct and paired261–27918–21a16 – 1922–2744p +rd +pp +Gans (1961) +
+ +A. pericensis + +4distinct and paired198–21816–196–812–1616–2033a + +rd + +aa +Gans (1963 a +) +
+ +A. plumbea + +4distinct and paired +233–282 +16–215–918–2720–3044p +rd +aa +Gans and Diefenbach (1972) +
+ +A. polygrammica + +4distinct and paired27020 +n / a +1816 +n / a + +n / a + +n / a + +n / a + +n / a + +n / a + +Vanzolini (2002) +
+ +A. prunicolor + +4distinct and paired181–21518–277–1010–1714–2033p +rd +aa +Perez et al. (2012) +
+ +A. ridleyi + +4distinct and paired172–19214–17 +a +16 – 1820–2844p +rd +aa +Gans (1963 b +) +
+ +A. rozei + +4distinct and paired205–209206 or a15–16 +14 +33p +rd +aa +Vanzolini (2002) +; +Costa et al. (2019) +
+ +A. sanctaeritae + +4distinct and paired +269–288 +18–206–7101433p +rd +aa +Costa et al. (2019) +
+ +A. saxosa + +4distinct and paired253–27217–21a18 – 2416–2144p +sc +aa +Oliveira et al. (2018) +
+ +A. schmidti + +4distinct and paired198–20220–227–81416–1733p +rd +aa +Gans (1964 a +) +
+ +A. slateri + +4distinct and paired176–21320–247–1010–1414–1633 +p +or a + + +rd + +aa +Gans (1967) +; +Costa et al. (2018) +
+ +A. slevini + +4distinct and paired204–21123–255–6 +10–14 +10–1222a +rd +papresent study
+ +A. spurrelli + +4distinct and paired218–22218–206–716–1816–1833p +rd +aa +Gans (1962 b +); +Costa (2020) +
+ +A. steindachneri + +4distinct and paired256–26617–18714–161633a +rd +pp +Gans (1964 b +) +
+ +A. supernumeraria + +4not distinct333–33722–2310–121417–1833 +n / a + + +rd + +a +n / a + +Mott et al. (2009) +
+ +A. talisiae + +4distinct and paired205–23417–29710–1414–1833a + +rd + +aa +Vanzolini (1995) +; +Costa et al. (2019) +
+ +A. tyaraju + +4distinct and paired204–22118–257–910–1413–1633p +rd + +n / a + +n / a + +Perez and Borges-Martins (2019) +
+ +A. townsendi + +4distinct and paired261–27922–267–816–1922–2744p +rd + +n / a + +n / a + +Gans (1961) +
+ +A. tragorrhectes + +4distinct and paired1963112121244p +rd +aapresent study
+ +A. uroxena + +0–4distinct and paired +210–213 +12–13a1414–1533p +rd +a +p + +Mott et al. (2008) +
+ +A. vanzolinii + +4distinct and paired +225–228 + +n / a +712–1316–1722p +rd +aa +Gans (1963 c +) +
+ +A. vermicularis + +4distinct and paired211–25423–345–718–2618–25 +4 +3 +n / a + + +rd + +aa +Gans and Amdur (1966) +
+ +A. xera + +4distinct and paired +225–234 +13 – 16512–1614–1633a + +rd + +aa +Thomas (1966) +
+
+
+ + + +Description of the +holotype +. + + + +medium-sized specimen; snout-vent length +233 mm +plus +0.50 mm +of cloacal portion; tail length +21.24 mm +, representing 9.1 % of snout-vent length; midbody diameter +8.2 mm +(3.5 % of snout-vent length); head relatively small, +6.90 mm +(~ 2.9 % of snout-vent length); snout compressed in dorsal view and slightly convex in profile view, hardly keratinised, rostrum projecting forward beyond the jaw (prognathous). Anterior portion of body is slightly narrower. Rostral subtriangular, visible in dorsal and ventral view (Fig. +1 +), almost as high ( +2.21 mm +) as wide ( +2.11 mm +), in contact with nasal and first supralabial lateroposteriorly. Nasals subrectangular, aligned at the midline (1.00 mm suture) (Fig. +1 A +), almost as long ( +2.05 mm +) as wide ( +1.99 mm +), in contact with first supralabial laterally and prefrontals posteriorly, with nostrils placed near the antero-inferior angle of the nasal shield (Fig. +1 B +). + + + + + + + +Amphisbaena amethysta + +sp. nov. +(Holotype, + +CEPB +2311 + +) +A +dorsal +B +lateral, and +C +ventral views of head. Scale bar: 3 mm. + + + +Prefrontals paired, relatively large (41.6 % of head length), with a shorter middorsal suture ( +2.01 mm +; 29.3 % of head length), longer than the nasal middorsal suture (1.00 mm, 14.6 % of head length), almost as long as frontal middorsal suture ( +2.10 mm +, 30.6 % of head length), anterior border convex, lateroposterior portion projected, in contact with second supralabial and ocular laterally, frontals posteriorly and in point contact with postocular posteriorly (Fig. +1 A +). Frontals subtriangular, longer (suture length) than wide ( +1.58 mm +), aligned at midline ( +2.10 mm +), in narrow contact with the oculars, and in broad contact posterolaterally with the postocular and parietal. Parietals in two larger irregulars segments, wider ( +1.48 mm +) than long ( +1.04 mm +), intercalated by four very smalls segments; not aligned at the midline, in narrow contact with postoculars laterally, and followed by the first dorsal half-annulus. Occipitals absent (Fig. +1 A +). Oculars almost diamond-shape, almost as long ( +1.57 mm +) as high ( +1.51 mm +), in contact with prefrontals and second supralabial anteriorly, third supralabial and postocular posteriorly, and in point contact with postsupralabial. Eyes slightly visible. Postocular longer than wide, sub-pentagonal, in contact with frontal, labial, parietal and the segments of the first dorsal half-annulus, and in point contact with prefrontal. Temporal subrectangular, higher ( +1.55 mm +) than long ( +0.92 mm +), in contact with third supralabial anteriorly, postocular and postsupralabial laterally and first dorsal half-annulus posteriorly (Fig. +1 B +). + + +Three supralabials, irregularly polygonal; first subtrapezoid, longer ( +2.05 mm +) than high ( +1.50 mm +), in contact with second supralabial posteriorly; second supralabial sub-pentagonal, higher ( +1.76 mm +) than long ( +1.59 mm +maximum length), in contact with prefrontal, ocular and third supralabial; third supralabial subtrapezoid, almost as high ( +1.37 mm +) as long ( +1.24 mm +), in contact with ocular and postsupralabial. Postsupralabial subquadrangular, representing almost 50 % of third supralabial high, in contact with temporal laterally and first half-annulus posteriorly (Fig. +1 B +). Mental longer ( +1.72 mm +) than wide, anterior border wider ( +1.72 mm +) than posterior ( +1.08 mm +), in contact with the first pair of infralabials and postmental. Postmental longer ( +2.01 mm +) than wide ( +1.65 mm +), in contact with the first and second infralabial, narrowly with malar, and two anterior postgenials. Postgenials with five shields irregularly distributed, in contact with malars and first ventral half-annuli (Fig. +1 C +). + + +Three infralabials, first medium sized, irregular polygonal, almost as long ( +1.55 mm +) as wide ( +1.56 mm +), in contact with second supralabial; second the largest, sub-pentagonal, wider ( +2.36 mm +) than long ( +1.81 mm +maximum length), in contact with third infralabial; third infralabial smallest, almost as long ( +1.24 mm +) as wide ( +1.37 mm +) (Fig. +1 C +). + + +Body annuli well demarcated, first and second annuli without enlarged dorsal segments. Segments become regularly rectangular toward posterior portion of body and progressively longer than wide, and smaller in size, and larger towards midventral areas starting from the fifth half-annulus. One hundred ninety-four dorsal and 195 ventral half-annuli, three lateral half-annuli, 14 caudal annuli plus tip rounded; tail relatively long with autotomy line on the fifth annulus, 18 / 18 dorsal and ventral segments at midbody, respectively and 28 segments in fourth caudal annulus. Lateral sulci clearly visible from the forty-ninth annulus; dorsal and ventral sulci absent. Cloacal plate with six segments increasing in size from towards midline, eleven postcloacal segments; four precloacal pores strongly visible on the row of segments on the last ventral half-annulus; each pore placed on the posterior half of a single segment, and distributed along a continuous series of segments, but pores in the medial scales placed laterally (Fig. +2 +). + + + + + + +Tail of + +Amphisbaena amethysta + +sp. nov. +(Holotype, + +CEPB +2311 + +). Detail of the autotomic site and four precloacal pores. Scale bar: 3 mm. + + + + + +Intraspecific variation. + + +The main variations in the type series for meristic and morphometric data are given in Table +2 +. Variation in the arrangement and contact of shields were also observed. + + +CEPB + +2280 + +presents frontal fused with the parietal and segments of first and second body annuli (Fig. +3 A +), + + +CEPB + +2309 + +present four parietals shields (Fig. +3 B +), and + + +CEPB + +2303 + +present supralabials fused in left side (Fig. +3 C +). + + + + + + + +Amphisbaena amethysta + +sp. nov. +in life (not identified specimen of type series) +A +lateral view and +B +dorsal. Photograph by T. B. S. + + + + + + + +Variation in meristic and morphometric (mm) data in the type series of + +Amphisbaena amethysta + +sp. nov. +S – sex, +DA +– dorsal half – annuli, +LA +– lateral half – annuli adjacent to cloacal region, +VA +– ventral half – annuli, +CA +– caudal annuli, +AS +– autotomic site, +DS +– dorsal segment in midbody, +VS +– ventral segment in midbody, +SCA +– segments of fourth caudal annulus, +PRCL +– precloacal segments, +POCL +– postcloacal segments, +PGE +– postgenials, +PA +– parietals, +SVL +– snout – vent length, +TL +– tail length, +BW +– body width in midbody, +bs +– brooked specimen, +lta +– last tail annuli, +n / a +– non-available data, + n – mutilated tail, * – tail cicatrised, ** – tail not cicatrised, +un +- unidentified. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
SpecimensS +DA + +LA + +VA + +CA + +AS + +DS + +VS + +SCA + +PRCL + +POCL +PGE 1 +PA + +SVL + +TL + +BW +
+ +CEPB +2311 + +(Holotype) +male194319514518182861151 / 123321.28.2
+ +CEPB +2301 + +(Paratype) +female19131915 + nlta 5 **19172971351 / 12063.5 + n6.4
+ +CEPB +2302 + +(Paratype) +male194419215520183171351 / 119917.87.2
+ +CEPB +2303 + +(Paratype) +female19441975 + nlta 5 *17162761451 / 11978.2 + n7.0
+ +CEPB +2308 + +(Paratype) +female1954196155181828 +n / a +1341 / 115013.05.1
+ +CEPB +2327 + +(Paratype) +female196319614520193161351 / 120516.56.0
+ +CEPB +2331 + +(Paratype) +female197419515518163061151 / 117514.45.8
+ +CEPB +2346 + +(Paratype) +male195419514518182961451 / 113811.34.3
+ +CEPB +2379 + +(Paratype) +male18541895 + nlta 5 **19182961351 / 11805.0 + n7.0
+ +CEPB +2381 + +(Paratype) +female189418915416162461251 / 116514.85.6
+ +CEPB +2298 + +female199319914 +n / a +18183071351 / 121516.77.4
+ +CEPB +2299 + +female19341936 + nlta 5 *18152961341 / 11867.6 + n5.1
+ +CEPB +2300 + +male194319514518172661251 / 121315.36.1
+ +CEPB +2304 + +female190419214518 +n / a +2661251 / 115511.63.5
+ +CEPB +2305 + +female192419215518162861361 / 117014.85.6
+ +CEPB +2306 + +female197419614418182881251 / 114510.84.2
+ +CEPB +2307 + +male19031906 + nlta 6 **18172761352 / 22206.8 + n7.9
+ +CEPB +2309 + + +un +19631976 + nlta 5 *1916 +n / a +61461 / 21257.9 + n4.5
+ +CEPB +2310 + +female193419414518182661120 / 119515.75.5
+ +CEPB +2312 + +female192319215518183061531 / 119015.16.6
+ +CEPB +2313 + +female189319015518152861451 / 113011.74.2
+ +CEPB +2314 + +female191319314518162661361 / 118511.95.7
+ +CEPB +2315 + +female193319314518192761251 / 117714.15.9
+ +CEPB +2316 + +female195319516620182871351 / 117315.26.4
+ +CEPB +2317 + +female19551965 + nlta 5 **18 +n / a + +n / a +61351 / 11803.0 + n5.3
+ +CEPB +2318 + +female196319515520183161151 / 112016.14.3
+ +CEPB +2319 + +female18641865 + nlta 5 **19182961251 / 1952.0 + n3.2
+ +CEPB +2320 + +female185418215519182671321 / 1927.72.6
+ +CEPB +2321 + +female194319216618183061251 / 116014.16.0
+ +CEPB +2322 + +female19641955 + nlta 5 **21182761141 / 11553.7 + n5.3
+ +CEPB +2323 + +female194419315516162461321 / 11128.83.0
+ +CEPB +2324 + +male193319316619183161351 / 116214.65.7
+ +CEPB +2325 + +female193419014519182761151 / 117414.26.1
+ +CEPB +2326 + +male198319716617172861051 / 119516.46.4
+ +CEPB +2328 + +female192319415618182861251 / 117315.56.5
+ +CEPB +2329 + +male195419615519183061361 / 117013.05.7
+ +CEPB +2330 + +male19441935 + nlta 5 **20183061351 / 11782.6 + n6.0
+ +CEPB +2332 + +male190 + n419015518162861551 / 1 +bs +11.14.0
+ +CEPB +2333 + +female196319415 +n / a +18172661021 / 120316.96.3
+ +CEPB +2334 + +female19431925 + n520183061461 / 11907.0 + n7.3
+ +CEPB +2335 + +female197319714520183161251 / 11359.73.9
+ +CEPB +2336 + +female190419015618172961341 / 11059.13.4
+ +CEPB +2337 + +female18641855 + nlta 5 **19182961351 / 11453.7 + n4.9
+ +CEPB +2338 + +female190 + n418613518172861251 / 1~ 16013.85.1
+ +CEPB +2339 + +female192419316519183261351 / 112710.64.0
+ +CEPB +2356 + +male188318916 +n / a +18182871331 / 119016.75.8
+ +CEPB +2380 + +female192319514517162561252 / 214011.55.6
Minimum-18531821341615246102-922.22.6
Maximum-19951991662119328156 +n / a +23321.28.2
Mean-192.93.6193.014.85.018.417.428.36.212.64.7 +n / a +167.313.45.5
Mode-19441951551818286135 +n / a +190 +n / a + +n / a +
Standard deviation-3.40.53.40.80.41.01.01.90.51.11.0 +n / a +34.13.51.3
+
+
+ + +Colour in life. + + +Dorsum and lateral parts with dark brown coloration on the segments, which is more pronounced in the vertebral (Fig. +4 A +) and dorsal section of the tail regions (Fig. +4 B +). Pink predominates in areas where the brown colour is less pronounced. We do not have photographs of the ventral region of the live specimen. + + + + + + +Variation (grey in the drawings) of cephalic shields of + +Amphisbaena amethysta + +sp. nov. +A +dorsal view of the head of + +CEPB +2280 + +with frontal variation +B +dorsal view of + +CEPB +2309 + +with four parietal shields, and +C +lateral view of left side of + +CEPB +2303 + +(paratype) with supralabials fused. Scale bar: 3 mm. + + + + + +Colour in preservative + + +(ethylic alcohol 70 %). +Dorsum cream, with brown colouring on the segments in the dorsum, lateral parts, and dorsal portions of the tail portions. Dorsal, lateral, and ventral portions of head pale brown, darker than the ventral portion. Venter cream coloured. + + + + +Etymology. + + +The specific epithet refers to the mineral amethyst that is a type of quartz and also the name of the region of the type locality “ Brejinho das Ametistas ”, a district located south of the municipality of Caetité, state of +Bahia +. This region has been an amethyst mining centre since the beginning of the 20 +th +century. +Spix and Martius (1938) +defined the mineral from the “ Brejinho das Ametistas ” mines as “ the beautiful amethysts ” on their trip through the “ Alto Sertão ” of +Bahia +at the beginning of the 19 +th +century ( +Cotrim 2015 +). The region currently has an economy based on mining and energy activities focused on wind energy production. The type series was collected during the execution of environmental programs within the scope of +Bahia +Mineração ( + +BAMIN + +), which operates in the exploration of iron ore in the “ Brejinho das Ametistas ” region. + + + + +Distribution and habitat. + + + +Amphisbaena amethysta + +sp. nov. +is known from municipality of Caetité municipality, state of +Bahia +, +Brazil +(Fig. +5 +). The region is in the northern portion of the Espinhaço Mountain Range, has an average altitude of +1000 m +a. s. l., and lies within the ecotone between two morphoclimatic domains, Caatinga and Cerrado. In the region there are patches of deciduous and semi-deciduous forests [“ Floresta Estacional Decidual ” and “ Floresta Semidecidual Montana ” sensu +IBGE (2023 +)] associated with valleys, slopes, and gallery forests, and containing floristic elements common to the vegetation of the Caatinga, Cerrado, and Atlantic Forest morphoclimatic domains. Areas of savannah vegetation with rock outcrops, typical of the woody Caatinga, occur at higher elevations (Fig. +6 +). + + + + + + +Geographic distribution of + +Amphisbaena amethysta + +sp. nov. +and others + +Amphisbaena +species + +from Espinhaço Mountain Range locality. Black arrows indicate the type localities of the species with more than one known geographic record. + + + + + + + +Caatinga site where the holotype of + +Amphisbaena amethysta + +sp. nov. +was collected in the in the northern portion of Espinhaço Mountain Range, in state of Bahia, Brazil. + + + + + +Phylogenetic relationships. + + +Our concatenated alignment totalled 4806 base pairs (1007 bp for 12 S, 528 bp for 16 S, 761 bp for +nd 2 +, 679 bp for +bdnf +, 574 bp for +c-mos +, and 1257 bp for +rag 1 +). Partition Finder identified a best-fit scheme composed of ten partitions with the GTR + G model. The resulting + +ML + +topology (Fig. +7 +) for the higher-level affinities was similar to those reported by previous studies ( +Mott and Vieites 2009 +; +Longrich et al. 2015 +; +Graboski et al. 2022 +) (Fig. +7 +). + +Amphisbaena amethysta + +sp. nov. +was recovered as a sister group of + +A. caetitensis + +, with 92 % of bootstrap support. The clade composed by + +Amphisbaena amethysta + +sp. nov. +and + +A. caetitensis + +was recovered as a sister group of + +A. angustifrons + +, + +A. darwini + +, + +A. kingi + +, + +A. leeseri + +, and + +A. munoai + +with low bootstrap support (21 %). The genetic distance ( +p-distance +) between + +Amphisbaena amethysta + +sp. nov. +and + +A. caetitensis + +is 6.1 % for 16 S. + + + + + + +Maximum likelihood tree zoomed in Central and South American +Amphisbaenidae +resulting from the RAxML analysis based on six concatenated genes, three nuclear (BDNF, c-mos, and RAG 1), and three mitochondrial (12 S, 16 S, and ND 2) genes. Red branches denote the clade composed of the new species and its sister species. Numbers on branches represent bootstrap values RAxML (> 70 %). The grey codes on the right side of the clades represent the South American subclades of + +Amphisbaena + +(G 01 – G 09) and West Indies clades (WIC 01 – WIC 02). + + + +
+
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