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Spilotes sulphureus Wagler, 1824 (Squamata: Colubridae): Review of distribution and first record in the state of Sergipe, Northeastern Brazil

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We present the first record of Spilotes sulphureus for the state of Sergipe. In Brazil, this species is distributed Amazon Rainforest, Atlantic Forest and Caatinga biomes. This large-bodied snake has arboreal habit, is diurnal, has oviparous reproduction, feeds on birds, eggs, lizards, bats, amphibians and small mammals and presents the defensive behaviour of lateral compression of the body.
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Spilotes sulphureus Wagler,  (Squamata: Colubridae):
review of distribution and rst record in the state of Sergipe,
northeastern Brazil
Hugo Andrade, , Rony Peterson Santos Almeida,  & Eduardo José dos Reis Dias
1 Universidade Federal de Sergipe, Departamento de Biociências, Laboratório de Biologia e Ecologia de Vertebrados, Avenida Vereador Olímpio
Grande s/n, Centro, CEP 49500-000, Itabaiana, SE, Brazil
2 Programa de Capacitação Institucional, Museu Paraense Emílio Goeldi, Coordenação de Ciências da Terra e Ecologia. Avenida Perimetral, Terra
Firme, CEP 66077-830, Belém, PA, Brazil
3 Corresponding author. E-mail: hugoandrade915@gmail.com
Abstract: We present the rst record of Spilotes sulphureus
for the state of Sergipe. In Brazil, this species is distributed
Amazon Rainforest, Atlantic Forest and Caatinga biomes.
is large-bodied snake has arboreal habit, is diurnal, has
oviparous reproduction, feeds on birds, eggs, lizards, bats,
amphibians and small mammals and presents the defen-
sive behaviour of lateral compression of the body.
Key words: Pseustes; gular ination; pung snake; geographical
distribution; Atlantic Forest
e colubrid snake Spilotes sulphureus (Wagler & Spix, )
is one of the larger species of the family (F ).
is species was previously placed in the genus Pseustes,
 and was later allocated to the present genus (J et
al. ). is arboreal snake has cryptic coloration and is
characterized by the defensive behavior of lateral compres-
sion of the body (M et al. ).
Spilotes sulphureus has wide distribution in rainforests of
Brazil, Ecuador, Peru, the Guianas, and Trinidad (P
 O-M ). In Brazil, this species is
distributed in Amazon Rainforest (C et al. ;
F et al. ; Á-P et al. ; B et
al. ), Atlantic Forest (M  C ;
A ; B-N et al. ; V
et al. ) and Caatinga (G et al. ).
On  May  and  January  two specimens of
Spilotes sulphureus, female ( mm SVL) and male (
mm SVL) (Figures  and , respectively), were collected
on the edge of an Atlantic Forest fragment in Refúgio
de Vida Silvestre Mata do Junco (RVSMJ) (°ʹʺ S,
°ʹʺ W;  m above sea level) in the municipality of
Capela, Sergipe state, Brazil.
e specimens were collected under permit -
issued by ICMBio and are housed in the snake collection
of the Laboratório de Biologia e Ecologia de Vertebrados
Check List 13(1): 2055, 21 February 2017 doi: https: //doi.org/10.15560/13.1.2055
ISSN 1809-127X © 2017 Check List and Au thors
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21 February 2017
Figures 1 and 2. Live adults of Spilotes sulphureus. 1. LABEVS32 female, 1075 mm. 2. LABEVS44 male, 1030 mm. Photos by RPSA.
1 2
Notes oN GeoGraphic DistributioN
Andrade et al. | First record of Spilotes sulphureus in Sergipe
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(LABEV), Universidade Federal de Sergipe (UFS) (LABEVS
 and LABEVS ).
Spilotes sulphureus presents the following characters:
dorsal scale rows , scales keeled and with one pair apical
pits; subcaudals paired, ; ventrals ; anal plate divided;
internasals in contact with rostral, anterior and posterior
nasal and prefrontals; prefrontals in contact with frontal,
posterior nasal, loreal, preocular, and supraocular; frontal
in contact with prefrontals, supraocular and parientals;
supraocular ; postorbitals ; preorbital ; upper labials
; lower labials ; anterior mentals in contact with the
mental groove and posterior mentals separated by small
gular scales (L et al. ).
Spilotes sulphureus may exhibit polymorphism in color.
Some individuals are yellow-brown with variable black bars
on the dorsum, head and upper labials, and yellow venter
(C et al. ) (Figure ). Other individuals are
immaculate yellow anteriorly, becoming darker posteriorly
(Figure ), as in B-N et al. (). Most records
of S. sulphureus are from the Amazon and Atlantic Forest
biomes (Figure ), likely associated with the arboreal habit
of the species (M et al. ). is snake is diurnal
(F ), feeds on birds (R  K ;
A et al. ), bats (R  B ),
lizards, small mammals, and amphibians (M et al.
; F ).
is paper presents the rst record of this species from
the state of Sergipe (Figure ). e new location is  km
Figure 3. Geographic distribution of Spilotes sulphureus in Brazil, based on data from Table 1.
Andrade et al. | First record of Spilotes sulphureus in Sergipe
Check List | www.biotaxa.org/cl Volume 13 | Issue 1 | Article 2055 3
Table 1. Localities of Spilotes sulphureus Wagler, 1824 in Brazil with coordinates and references.
State Municipality Collection site Latitude Longitude Reference/collection number
SE Capela Refúgio de Vida de Silvestre Mata do Junco 10°32’23” S 037°03’43” W This research
BA Esplanada not reported 11°47’46” S 037°45’42” W IBSP 49248
BA Porto Seguro Reserva Florestal da CVRD 16°26’59” S 039°03’53” W MBML 396
BA Itagibá Centro da Cidade 14°17’00” S 039°50’40” W Lisboa et al. (2009)
BA Ituberá Centro da cidade 13°43’31” S 039°08’53” W Lisboa et al. (2009)
BA Mata de São João Reserva Imbassaí 12°28’45” S 037°57’33” W Marques et al. (2011)
BA Mucuri Centro da cidade 18°04’33” S 039°33’37” W Lisboa et al. (2009)
BA Ilhéus Centro da cidade 14°47’37” S 039°02’46” W Lisboa et al. (2009)
AC Cruzeiro do Sul Centro da Cidade 07°37’40” S 072°40’33” W siLva et al. (2010)
AL Campo Alegre Fazenda Pindoba 09°45’32” S 036°14’09” W Lisboa et al. (2009)
AM Coari Base Operacional Geólogo Pedro de Moura 04°53’07” S 065°21’00” W Prudente et al. (2010)
AM Coari Porto Urucu 04°02’09” S 064°04’43” W MPEG.HOP 023878
AM Fazenda Scheer Rio Ituxi 08°20’00” S 065°43’00” W aviLa-Pires et al. (2009)
AM Humaitá km 80 - BR - Humaitá a Lábrea 07°30’22” S 063°01’15” W IBSP 40818
AM Manaus Itacoatiara 03°06’07” S 060°01’30” W IBSP 40442
AM not reported Floresta marginal ao longo do rio Japurá 03°08’02” S 065°06’00” W (Type locality) Cunha et al. (1985)
AM not reported Río Japurá 01°37’00” N 068°00’00” W FPR 60223
AM Presidente Figueiredo U.H.E. Balbina 02°02’04” S 060°01’30” W IBSP 51986
AM Presidente Figueiredo Balbina. Área de inundação da UHE. 01°13’15” S 060°01’24” W MPEG.HOP 017585
AM Tefé Porto Urucú. 04°24’51” S 065°31’23” W MPEG.HOP 018225
AP Laranjal do Jari Cachoeira Santo Antônio. 00°46’39” N 053°24’27” W MPEG.HOP 017739
AP Serra do Navio Distrito Maganífero, ICOMI 01°36’02” N 052°36’45” W MPEG.HOP 000333
CE Pacoti Maciço de Baturité 04°14’09” S 038°55’01” W borges-nojosa et al. (2006)
CE Ubajara Parque Nacional de Ubajara 03°50’25” S 040°54’28” W borges-nojosa et al. (2006)
ES Cachoeiro de Itapemirim Estação Ambiente Ilha dos Meirelles 20°50’56” S 041°06’46” W deLfino & Rabello (2007)
ES Colatina not reported 19°32’22” S 040°37’50” W IBSP 49534
ES Guarapari Rodovia do Sol. 20°40’00” S 040°29’51” W MBML 2187
ES Ibiraçu not reported 19°49’55” S 040°22’11” W IBSP 49983
ES Linhares Degredo 19°23’28” S 040°04’20” W MBML 2140
ES Santa Teresa Santa Lúcia. 19°56’08” S 040°36’01” W MBML 449
ES São Mateus not reported 18°44’00” S 039°51’00” W MCZ R 3770
ES Viana not reported 20°23’25” S 040°29’46” W MBML 1946
GO Luziânia Pôrto Nacional 16°13’56” S 048°01’51” W IBSP 19681
MA Junco do Maranhão Rio Gurupi. Nova Vida. 02°13’18” S 046°30’58” W MPEG.HOP 013664
MG Caratinga RFMA (Trilha acesso KK-Q) 19°47’23” S 042°08’21” W UFMG 113
MG Caratinga RPPN Feliciano Miguel Abdala 19°43’00” S 041°49’00” W PaLMuti et al. (2009)
MG Jequitinhonha Reserva Biológica da Mata Escura 16°20’00” S 041°05’00” W Duarte (2012)
MG Marliéria Parque Estadual do Rio Doce 19°42’44” S 042°43’56” W UFMG 112
MG Teólo Otoni Próximo à captação São José. 17°51’27” S 041°30’19” W FUNED 3173
MT Barra do Bugres Est. Ecológica da Serra das Araras (SEMA) 15°03’21” S 057°11’25” W MPEG.HOP 017062
MT Barra do Garças Fazenda Três Marias - norte de MT 15°53’24” S 052°15’24” W IBSP 30420
MT Barra dos Bugres Estação Ecológica da Serra das Araras 15°39’00” S 057°01’00” W nasCiMento et al. (1988)
MT Nova Ubiratã Margem esquerda do Rio Von den Steinen 12°59’35” S 054°45’59” W UFMT 5374
MT Paranaíta Sete quedas - Rio Teles Pires 09°35’40” S 056°40’55” W UFMT 8225
MT Pontes e Lacerda Fazenda Palmar 15°13’34” S 059°20’07” W IBSP 45429
MT Tangará da Serra not reported 14°37’10” S 057°29’09” W IBSP 50588
MT Xavantina 264 - km norte 14°40’29” S 052°19’33” W IBSP 29395
PA Acará Ipitinga. Estrada Mojú-Acará. 02°01’35” S 048°27’37” W MPEG.HOP 015519
PA Alenquer not reported 00°18’31” S 054°53’12” W MPEG.HOP 023677
PA Almeirim REBIO Maicuru 00°24’17” N 053°48’09” W MPEG.HOP 023721
PA Belém not reported 01°27’00” S 048°29’00” W MCZ R 53215
PA Benevides Estrada de Genipauba 01°21’40” S 048°14’36” W MPEG.HOP 015466
PA Capitão Poço Santa Luzia 01°52’13” S 047°13’23” W MPEG.HOP 002914
PA Castanhal Boa Vista 01°17’24” S 047°56’34” W MPEG.HOP 010528
PA Conceição do Araguaia Codespar, Fz. 08°15’28” S 049°15’53” W IBSP 26762
PA Curuçá Marauá 00°44’01” S 047°51’07” W MPEG.HOP 007091
PA Dom Eliseu Sítio Bela Vista. 04°12’03” S 047°51’55” W MPEG.HOP 015534
PA Inhangapi Arraial do Carmo 01°27’15” S 047°56’43” W MPEG.HOP 001566
PA Itaituba Parque Nacional Amazônia 04°15’07” S 055°59’42” W MPEG.HOP 021139
PA Jacundá Rio Tocantins. 04°35’34” S 049°14’05” W MPEG.HOP 016719
PA Juruti Estrada Juruti - Beneciamento 02°09’46” S 056°05’44” W MPEG.HOP 023258
Continued
Andrade et al. | First record of Spilotes sulphureus in Sergipe
Check List | www.biotaxa.org/cl Volume 13 | Issue 1 | Article 2055 4
State Municipality Collection site Latitude Longitude Reference/collection number
PA Marabá Serra Norte. Carajás. Área do manganês do Azul. 05°20’15” S 049°06’04” W MPEG.HOP 017092
PA Maracanã km 23 da estrada de Maracanã 00°46’28” S 047°27’15” W MPEG.HOP 004839
PA Melgaço Floresta Nacional de Caxiuanã. Itaperu. 01°32’08” S 051°08’12” W MPEG.HOP 022080
PA Óbidos ESEC Grão Pará 01°54’05” S 055°31’17” W MPEG.HOP 023749
PA Oriximiná Serra do Acarai 01°46’01” S 055°51’36” W MPEG.HOP 023705
PA Ourilândia do Norte not reported 06°45’25” S 051°04’54” W MPEG.HOP 025308
PA Palestina do Pará Porto Jarbas Passarinho. Transamazônica. 05°44’26” S 048°19’07” W MPEG.HOP 013411
PA Paragominas Margem do rio Jaboti - Km 70. 02°59’45” S 047°21’10” W IBSP 32411
PA Parauapebas Núcleo Urbano. Acesso Sul. Carajás 06°04’33” S 049°53’56” W MPEG.HOP 023009
PA Peixe-Boi not reported 01°14’55” S 047°18’51” W MPEG.HOP 002588
PA Portel Centro da Cidade 01°56’08” S 050°49’16” W IBSP 45869
PA Rio Parauapebas Serra do Carajás 06°33’00” S 050°34’00” W Cunha et al. (1985)
PA Santa Bárbara do Pará Parque Ecológico de GUNMA. 01°14’56” S 048°17’27” W MPEG.HOP 021323
PA Senador José Porfírio UHE-Belo Monte 02°48’18” S 051°50’14” W MPEG.HOP 020985
PA Tomé-Açu Ipitinga. Estrada Mojú-Acará. 02°25’06” S 048°09’16” W MPEG.HOP 013356
PA Tucuruí Margem esquerda do Rio Tocantins 03°45’58” S 049°40’21” W IBSP 47060
PA Vigia Santa Rosa, Estrada da Vigia. 00°56’44” S 048°04’42” W MPEG.HOP 009309
PA Viseu Santa Luzia 01°33’50” S 046°27’23” W MPEG.HOP 014124
PA Vitória do Xingú UHE-Belo Monte. 03°11’29” S 051°57’40” W MPEG.HOP 019876
PB João Pessoa APP Mata do Buraquinho 07°08’42” S 034°51’54” W santana et al. (2008)
PB João Pessoa Centro da Cidade 07°07’08” S 034°50’42” W Lisboa et al. (2009)
RJ Cachoeiras de Macacu Estação Ecológica Estadual do Paraíso 22°26’00” S 042°50’00” W vrCibradiC et al. (2011)
RJ Duque de Caxias Parque Natural Municipal de Taquara 22°35’00” S 043°14’00” W saLLes et al. (2010)
RJ Maricá Litoral da cidade 22°57’19” S 042°49’56” W Lisboa et al. (2009)
RJ Miracema not reported 21°24’44” S 042°11’48” W IBSP 22754
RJ Petropolis not reported 22°23’20” S 043°11’14” W MNHN(RA) 1877.180
RJ Rio de Janeiro Serra do Mendanha 22°48’00” S 043°28’00” W Pontes et al. (2009)
RJ Rio de Janeiro not reported 22°54’00” S 043°14’00” W MCZ R 1381
RO Porto Velho UHE JIRAU. Jací-Paraná 08°45’23” S 063°51’18” W MPEG.HOP 023989
RO Ji-Paraná Centro da Cidade 10°52’41” S 061°56’06” W bernarde et al. (2012)
RO Porto Velho Centro da Cidade 08°45’40” S 063°54’06” W bernarde et al. (2012)
SP Paranapiacaba Centro da Cidade 23°46’36” S 046°17’57” W Lisboa et al. (2009)
SP Paranapiacaba not reported 23°48’00” S 046°03’00” W Marques & CaLLeffo (1997)
* State abbreviations: AC, Acre; AL, Alagoas; AM, Amazonas; BA, Bahia; CE, Ceará; ES, Espírito Santo; GO, Goiás; MA, Maranhão; MG, Minas Gerais; MT, Mato Grosso;
PA, Pará; PB, Paraíba; RJ, Rio de Janeiro; RO, Rondônia; SP, São Paulo; SE, Sergipe.
Not exact geographical position.
Collections acronyms: Foundation Puerto Rastrojo (FPR- Colômbia); Foundation Ezequiel Dias (FUNED); Instituto Butantan (IBSP- São Paulo); Museum of Biology
Prof. Mello Leitão (MBML); Museum of Comparative Zoology, Harvard University (MCZ); Muséum National d’Histoire Naturelle Reptiles and Amphibians Collection
(RA) of the Muséum National d’Histoire Naturelle (MNHN - Paris); Museum Paraense Emilio Goeldi - Ophidia Collection (MPEG.HOP); Museum of Zoology of the
University of São Paulo (MZUSP); Universidade Federal de Mato Grosso (UFMT); University Federal de Minas Gerais (UFMG).
Table 1. Continued.
south of the nearest locality at Pindoba Farm, municipality
of Campo Alegre, Alagoas (L et al. ), and  km
north of the nearest locality in Bahia (IBSP ). e
location is  km from the type locality on the Japurá
River, Amazonas (C et al. ).
is new record lls a gap in the distribution of the
species, and contributes to knowledge of the biological
diversity in Sergipe state, which is relatively poorly known.
ACKNOWLEDGEMENTS
We thank J.M.G. Santos-Júnior, D.N. Costa, E.L. Alves,
A.C. Lima and M.L. Mota for help with eldwork; A.L.C.
Prudente for records this species in the Herpetological
Collection of the Museum Paraense Emílio Goeldi;
Secretaria de Meio Ambiente e Recursos Hídricos of the
state of Sergipe; and ICMBio for logistic support and
collecting permission in the RVSMJ.
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Authors’ contributions: HA wrote the text; RPSA collected the
specimens and wrote the text; and EJRD identied the specimens
and wrote the text.
Received:  June 
Accepted:  February 
Academic editor: Ross MacCulloch
http://hdl.handle.net//
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... The third most abundant 3FTx in S. sulphureus venom, sulmotoxin 2, did not show taxon specificity. Spilotes sulphureus are arboreal snakes that are generalist predators, feeding on birds, lizards, amphibians, and small mammals ( Andrade et al., 2017). The venom of S. sulphureus has two taxa specific 3FTxs present, and this correlates with its diet, which includes a diversity of prey. ...
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The genus Pseustes Fitzinger, 1843 is composed of three recognized species, Pseustes poecilonotus, P. shropshirei and P. sulphureus, which may be the largest sized colubrid snake in the New World. The group has a complex systematic history that has yet to be untangled using modern molecular phylogenetic approaches. The systematic position, within-group diversity and distribution are therefore uncertain. We obtained samples of four species from multiple specimens across their distribution and analysed one nuclear and two mitochondrial genes to determine the phylogenetic placement of the genus and infer relationships among Pseustes lineages. We find strong support for the paraphyly of Pseustes with respect to the monotypic genus Spilotes, both of which are nested within a clade of at least 23 other New World Colubrinae genera. Based on our results, we formally revise the taxonomy of P. poecilonotus and P. sulphureus, resurrecting the taxon P. polylepis for populations of P. poecilonotus from South America and allocating P. sulphureus to the genus Spilotes which renders both genera monophyletic. Additionally, we identify two lineages that are putatively new and currently unrecognized species. Finally, the placement of P. sulphureus, the type species of Pseustes, in the genus Spilotes, requires the allocation of the senior synonym Phrynonax be considered for the remaining Pseustes taxa.