The Snake Fossil Record from Brazil
Brazilian fossil snakes have had a rich and diverse record since the Late Cretaceous. Here, we provide an overview of the snake fossil record from Brazil, gathering all available data to reconstruct the diversity and distribution of taxa through time. Mesozoic snakes are scarce, represented by the putative ophidian Tetrapodophis amplectus, the snake Seismophis septentrionalis, and some undescribed material referred to ‘Anilioidea.' The Paleocene of São José do Itaboraí holds the richest and most-diverse Brazilian snake fauna composed of several taxa of madtsoids, ‘aniloids,' boids, ‘booids,' and caenophidians. Fossils from the Brazilian Neogene are reported for the Late Miocene of the Solimões Formation including boids, Colubroides, and an alethinophidian of uncertain relationships, Colombophis. Quaternary deposits yield a rich snake fauna represented by extant species such as boids and colubroids. The Brazilian snake fossil record is crucial for understanding issues about the early evolution of snakes and provides valuable insights into paleobiogeography, paleoenvironmental, and morphological studies. O Brasil possui um registro fossilífero rico e diversificado de serpentes desde o Cretáceo tardio. Neste trabalho nós fornecemos um panorama geral do registro fossilífero de serpentes no território brasileiro, compilando todos os dados disponíveis para reconstruir a diversidade e distribuição dos táxons no decorrer do tempo. Serpentes Mesozoicas são escassas, ocorrendo o possível ofídio de quatro patas Tetrapodophis amplectus, a Seismophis septentrionalis e alguns materiais ainda não descritos atribuídos a 'Anilioidea'. O Paleoceno de São José do Itaboraí detém a mais rica e diversa fauna de serpentes que é composta por espécies de madtsoideos, 'anilioideos', boídeos, 'booideos' e um cenofídeo. Fósseis do Neógeno são reportados no Mioceno da Bacia do Acre, incluindo boídeos, Colubroides e um aletinofídeo de relações incertas, Colombophis. Os depósitos do Quaternário detêm uma rica fauna de serpentes de espécies viventes, como boídeos e um amplo registro de Colubroideos. O registro fossilífero de serpentes brasileiras constitui uma chave crucial para o entendimento de questões sobre a evolução inicial das serpentes, além de, propiciar estudos sobre paleobiogeografia, paleoambiente e morfologia.Abstract
Resumo

Distribution of the Mesozoic snakes. (A) Tetrapodophis amplectus from Nova Olinda Member, Crato Formation, Ceará State (Martill et al., 2015); (B) Seismophis septentrionalis from Falésia do Sismito of Cajual Island, Maranhão State (Hsiou et al., 2014); (C) Indeterminate snake from Adamantina Formation, São Paulo State, the symbol ‘?' denotes the unknown locality in the state (Bertini and Bonfim-Júnior, 1988); (D) Indeterminate ‘Anilioidea' from General Salgado, São Paulo state (Zaher et al., 2003); and (E) Indeterminate ‘Anilioidea' from Monte Alto city, Adamantina Formation (Fachini and Iori, 2009; Fachini and Hsiou, 2011).

Main fossil snakes species from Brazil showing the typical preservation found in the country, isolated precloacal vertebrae. (A) Seismophis septentrionalis; (B) schematic drawing of Madtsoia camposi (modified from Rage, 1988); (C) schematic drawing of Hoffstetterella brasiliensis (modified from Rage, 1988); (D) Colombophis spinosus (modified from Hsiou et al., 2010); and (E) Colombophis portai (modified from Hsiou et al., 2010). In (1) anterior; (2) posterior; (3) lateral; (4) ventral; and (5) dorsal views.

Distribution of the Paleogene snakes (square symbol). All material comes from the Middle Paleocene of São José do Itaboraí, Rio de Janeiro. (A) Madtsoia camposi (Rage, 1998); (B) Coniophis cf C. precedens (Rage, 1998); (C) Hoffstetterella brasiliensis (Rage, 1998); (D) Hechtophis austrinus (Rage, 2001); (E) cf. Hechtophis (Rage, 2001); (F) Corallus pricus (Rage, 2001); (G) cf. Corallus (Rage, 2001); (H) Waincophis pressulus (Rage, 2001); (I) Waincophis cameratus (Rage, 2001); (J) cf. W. pressulus (Rage, 2001); (K) cf. W. cameratus (Rage, 2001); (L) cf. Waincophis (Rage, 2001); (M) Boinae A (Rage, 2001); (N) Boinae B (Rage, 2001); (O) Indeterminate boine (Rage, 2001); (P) Paraungaliophis pricei (Rage, 2008); (Q) cf. Paraungaliophis (Rage, 2008); (R) Itaboraiophis depressus (Rage, 2008); (S) Paulacoutophis perplexus (Rage, 2008); and (T) ?Russellophiidae (Rage, 2008). Distribution of the Neogene snakes (circle symbol).
All material comes from the Late Miocene of Solimões Formation. (A) Colombophis portai from the Morro do Careca and Patos, Acre State; Talismã locality, Amazonas state locality (Hsiou et al., 2010); (B) Colombophis spinosus from Talismã locality, Purus River, Amazonas State (Hsiou et al., 2010); (C) aff. Epicrates from Talismã locality, Purus River, Amazonas State (Hsiou and Albino, 2010); (D) Waincophis sp. from Belford locality, Upper Juruá River, Belford site (Hsiou and Albino, 2010); (E) Eunectes sp. from Talismã locality, Purus River, Amazonas State (Hsiou and Albino. 2009); (F) Indeterminate ‘Colubridae' A from Lula locality near Sena Madureira city, Amazonas State (Hsiou and Albino, 2010); and (G) Indeterminate ‘Colubridae' B from Talismã locality, Purus River, Amazonas State (Hsiou and Albino, 2010).

Distribution of the Quaternary snakes. (A) Eunectes murinus from Gruta do Urso, Tocantins (Hsiou et al., 2013) and Lapa dos Brejões, Morro do Chapéu, Bahia (Camolez and Zaher, 2010); (B) Eunectes sp. from Buraco do Japonês, Bonito city, Mato Grosso do Sul (Camolez and Zaher, 2010); (C) Boa constrictor from Caverna Carneiro, Goiás state; Morro do chapéu, Bahia state, and Lapa do Santo, Lapa das Boleiras, and Lapa do Boquete, Minas Gerais state (Camolez and Zaher, 2010); (D) Epicrates cenchria from Lapa do Santo, Lapa das Boleiras, and Lapa do Boquete from Minas Gerais (Camolez and Zaher, 2010); (E) cf. Epicrates from speological province of Ubajara, Ceará (Hsiou and Albino, 2010); (F) Corallus cf. C. hortullanus from Caverna Carneiro, Goiás and Lapa do Boquete, Minas Gerais state (Camolez and Zaher, 2010); (G) Chironius sp. from Gruta Cuvieri, Lapa do Santo and Lapa das Boleiras, Minas Gerais states (Camolez and Zaher, 2010); (H) Tantilla sp. from Abismo Ponta de Flecha, Iporanga city, São Paulo state (Camolez and Zaher, 2010); (I) cf. Pseustes sulphureus from Gruta Cuvieri, Matozinhos city, Minas Gerais state (Camolez and Zaher, 2010); (J) cf. Mastigodryas bifossatus from Caverna Carneiro, Goiás state, Lapa dos Brejões, Bahia state, and Lapa do Santo, Minas Gerais state (Camolez and Zaher, 2010); (K) Indeterminated ‘Colubridae' from Abismo Ponta de Flecha, Iporanga, São Paulo state and Lapa do Santos, Matozinhos, Minas Gerais state (Camolez and Zaher, 2010); (L) Helicops leopardinus from Lapa das Boleiras, Matozinhos, Minas Gerais state (Camolez and Zaher, 2010); (M) cf. Philodryas from Lapa do Santo, Matozinhos, Minas Gerais state (Camolez and Zaher, 2010); (N) Indeterminate Pseudoboini from Caverna Nossa Senhora Aparecida, Goiás state and Abismo Ponta de Flecha, Iporanga, São Paulo (Camolez and Zaher, 2010); (O) Indeterminate Xenodontinae from Abismo Ponta de Flecha, Iporanga, São Paulo state and Lapa do Santo, Matozinhos, Mina Gerais state (Camolez and Zaher, 2010); (P) Indeterminate Dipsadinae from Caverna Nossa Senhora de Aparecida, Goiás state (Camolez and Zaher, 2010); (Q) Micrurus corallinus from Abismo Ponta de Flecha, Iporanga, São Paulo state (Camolez and Zaher, 2010); (R) Micrurus sp. from Abismo Ponta de Flecha, Iporanga, São Paulo state (Camolez and Zaher, 2010); (S) Bothrops sp. from speleological province of Ubajara, Ceará state and Caverna Carneiro, Goiás state, Lapa do Santo and Lapa do Boquete, Minas Gerais state (Camolez and Zaher, 2010); (T) Crotallus durissus from Lapa dos Brejões, Bahia state and Lapa do Santo, Lapa do Boquete from Minas Gerais state (Camolez and Zaher, 2010); and (U) Indeterminate Viperidae from Mississippi locality/Estirão do Mississipi, Upper Juruá river, Acre state (Hsiou and Albino, 2011).

Fossil snakes from Brazil plotted into the phylogenetic hypothesis of Martill et al. (2015) (Modified from Martill et al., 2015). Dashed lines indicate the dubious relationships based only on vertebral morphology. (1) Tetrapodophis amplectus; (2) Seismophis septentrionalis; (3) Indeterminate snake; (4) Indeterminate ‘Anilioidea'; (5) Indeterminate ‘Anilioidea'; (6) Madtsoia camposi; (7) Coniophis cf C. precedens; (8) Hoffstetterella brasiliensis; (9) Hechtophis austrinus (10) cf. Hechtophis; (11) Corallus pricus; (12) cf. Corallus; (13) Waincophis pressulus; (14) Waincophis cameratus; (15) cf. W. pressulus; (16) cf. W. cameratus; (17) cf. Waincophis; (18) Boinae A; (19) Boinae B; (20) Paraungaliophis pricei; (21) cf. Paraungaliophis; (22) Itaboraiophis depressus; (23) Paulacoutophis perplexus; (24) Colombophis portai; (25) Colombophis spinosus; (26) aff. Epicrates; (27) Waincophis sp.; (28) Eunectes sp.; (29) Indeterminate ‘Colubridae' A; (30) Indeterminate ‘Colubridae' B; (31) Eunectes murinus; (32) Eunectes sp.; (33) Boa constrictor; (34) Epicrates cenchria; (35) cf. Epicrates; (36) Corallus cf. C. hortullanus; (37) Chironius sp. (38) Tantilla sp.; (39) cf. Pseustes sulphureus; (40) cf. Mastigodryas bifossatus; (41) Indeterminate ‘Colubridae'; (42) Helicops leopardinus; (43) cf. Philodryas; (44) Indeterminate Pseudoboini; (45) Indeterminate Xenodontinae; (46) Indeterminate Dipsadinae; (47) Micrurus corallinus; (48) Micrurus sp.; (49) Bothrops sp.; (50) Crotallus durissus; and (51) Indeterminate Viperidae.
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