Zootaxa 4097 (4): 511529 ISSN 1175-5326 (print edition)

http://www.mapress.com/j/zt/
Copyright 2016 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://doi.org/10.11646/zootaxa.4097.4.4
http://zoobank.org/urn:lsid:zoobank.org:pub:9D1538CB-C539-4540-8DA0-BD91CE0292F9

Another new and threatened species of lancehead genus Bothrops

(Serpentes, Viperidae) from Ilha dos Franceses, Southeastern Brazil

FAUSTO E. BARBO1,5, JOO LUIZ GASPARINI2, ANTONIO P. ALMEIDA3, HUSSAM ZAHER1,

FELIPE G. GRAZZIOTIN1, RODRIGO B. GUSMO4, JOS MRIO G. FERRARINI4
& RICARDO J. SAWAYA4
1
Museu de Zoologia, Universidade de So Paulo, P.B. 42494, So Paulo 04218970, So Paulo, Brazil
2
Departamento de Ecologia e Oceanografia, Universidade Federal do Esprito Santo, Vitria 29075910, Esprito Santo, Brazil
3
ICMBIO, Reserva Biolgica de Comboios. P.B. 105, CEP 29900-970, Linhares, Esprito Santo, Brazil
4
Departamento de Cincias Biolgicas, Universidade Federal de So Paulo, Diadema 09972270, So Paulo SP, Brazil
5
Corresponding author. E-mail: faustoebarbo@gmail.com

Abstract

A new insular species of the genus Bothrops is described from Ilha dos Franceses, a small island off the coast of Esprito
Santo State, in southeastern Brazil. The new species differs from mainland populations of B. jararaca mainly by its small
size, relative longer tail, relative smaller head length, and relative larger eyes. The new species is distinguished from B.
alcatraz, B. insularis and B. otavioi by the higher number of ventral and subcaudal scales, relative longer tail and smaller
head. The new species is highly abundant on the island, being nocturnal, semiarboreal, and feeding on small lizards and
centipeds. Due its unique and restricted area of occurrence, declining quality of habitat, and constant use of the island for
tourism, the new species may be considered as critically endangered.

Key words: Atlantic Forest, Bothrops, Conservation, Island rule, Heterochrony, taxonomy

Resumo

Uma nova espcie insular de jararaca descrita para a Ilha dos Franceses, localizada na costa do estado do Esprito Santo,
no sudeste do Brasil. A nova espcie difere das populaes continentais de B. jararaca principalmente pelo menor tama-
nho, maior comprimento relativo de cauda, menor comprimento relativo da cabea e olhos relativamente maiores. Dife-
rencia-se tambm de B. alcatraz, B. insularis e B. otavioi pelo maior nmero de escamas ventrais e subcaudais, maior com-
primento relativo de cauda e menor comprimento relativo de cabea. A nova espcie encontrada em grande abundncia
na ilha, sendo noturna, semi arborcola, e se alimenta de pequenos lagartos e centopeias. Devido sua reduzida rea de
ocorrncia, declnio da qualidade do habitat, e acesso constante de turistas ilha, a nova espcie deve ser considerada
como criticamente ameaada de extino.

Palavras-chave: Mata Atlntica, Bothrops, Conservao, Regra das ilhas, Heterocronia, Taxonomia

Introduction

Pitvipers diversity, morphology, and ecology are remarkable (Campbell & Lamar 2004). Approximately 230
species of crotalines are currently recognized, with at least 14 new species being described in the last four years
only (see Uetz & Hoek 2015). Bothrops is the most diverse genus within crotalines, including about 50 species
splitted into six different monophyletic groups: B. alternatus, B. atrox, B. jararaca, B. jararacussu, B. neuwiedi,
and B. taeniatus (Martins et al. 2001, 2002; Arajo & Martins 2006; Fenwick et al. 2009; Carrasco et al. 2012).
The genus is distributed from Central to South America, throughout several types of landscapes, including
rainforests and open habitats (Martins et al. 2002; Campbell & Lamar 2004).

Accepted by P. Passos: 17 Feb. 2016; published: 4 Apr. 2016 511

 The Bothrops jararaca complex comprising four known species: Bothrops jararaca (Wied, 1824), B. alcatraz
Marques, Martins & Sazima, 2002, B. insularis (Amaral, 1921), and B. otavioi Barbo, Grazziotin, Sazima, Martins
& Sawaya, 2012. The only mainland species, B. jararaca, is widespread throughout the Brazilian Atlantic forest,
from the northeastern highland forested enclaves of Bahia to the southern Atlantic forest of the state of Rio Grande
do Sul (Campbell & Lamar 1989; Martins et al. 2002), also occurring in disturbed environments such as borders of
forests and urbanized areas (Marques et al. 2009; Barbo et al. 2011). The latter three species are all endemic from
small continental islands of the Southeastern coast of Brazil (Ilha dos Alcatrazes, Ilha da Queimada Grande, and
Ilha da Vitria, respectively). Those islands correspond to emerging mountain picks with relatively high elevation
(more than 200 m above sea level; asl hereafter), located more than 20 km off the coast, and surrounded by
relatively deep waters (more than 30 m below sea level; hereafter bsl).
Recent studies have shown that the number of species for the group is probably underestimated (Grazziotin et
al. 2006; Barbo et al. 2012). Known mainland populations of B. jararaca possibly form a complex of species since
several highly genetically structured populations were identified by mitochondrial DNA analyses (Grazziotin et al.
2006). At the same time, the insular populations have shown several apparently fixed autapomorphic
morphological characters of coloration and scalation that distinguish them from their mainland relatives. These
unique morphotypes suggest the presence of distinct evolutionary trajectories that have not been hitherto
comprehensively studied throughout the entire distribution of the group (Marques et al. 2002).
Here, we describe a fifth species of the B. jararaca group from Ilha dos Franceses (maximum altitude 36 m
asl), a small island located less than five kilometers off the coast of the state of Esprito Santo and surrounded by
very shallow waters (less than 5 m bsl). The fact that this species lives in a small island easily accessible from the
shore makes it unique for ecological and evolutionary studies, but at the same time highly threatened and
dependent of governmental actions for its preservation.

Materials and methods

We examined 58 specimens of the new species, being five collected in the field by our team, nine photographed
and released in the field, and 44 previously available in scientific collections. We compared them with 154
specimens of Bothrops jararaca from mainland localities distributed throughout the states of Esprito Santo (n =
82), So Paulo (n = 30), Paran (n = 23), Santa Catarina (n = 9), and Rio Grande do Sul (n = 10). We also compared
the new species with eight specimens of B. alcatraz, 26 B. insularis, and 31 B. otavioi. Because only a few
specimens of B. alcatraz were available for this analysis, we complemented its sample with data retrieved in
Marques et al. (2002), totaling 32 specimens. Additional specimens of B. insularis from literature and unpublished
data (Amaral 1921; K. Kasperoviczus, pers. com.) were also included in some analyses (n = 374). Specimens
examined are cited in the Appendix and institutional abbreviations follow Sabaj-Prez (2015).
Linear morphometric analyses. Samples available for this analysis included a total of 260 males and 311
females (Table 1). We measured snoutvent length (SVL) and caudal length (CL) using a flexible ruler to the
nearest millimeter (mm). We also measured the head length (HL; from snout to extreme posterior portion of
mandible), with calipers to the nearest 0.01 mm. We determined the trunk length (TR) as SVL subtracting up the
HL. The relative tail length (RTL) corresponded to CL/SVL, whereas the relative head length (RHL) was obtained
by HL/TR. We counted ventral scales (from the first scale wider than long), intersupraocular scales (lines of
anterior, middle, and posterior scales between supraoculars), and interrictal scales (line of scales linking last
supralabials). Lateral scales of head were also counted: interoculabials (between upper 34th supralabials and
suboculars), circumorbitals (scales contacting eye), pre and postfoveals, temporals, and supra and infralabials.
Statistical tests were performed in R (R Core Team 2013). Adults and juveniles were analyzed separately due
to possible bias in RTL and RHL throughout ontogeny. Males and females from Ilha dos Franceses were
considered adults when the specimen was longer than 450 mm SVL and 540 mm SVL, respectively (see Natural
History Section below). Males from the mainland population of the state of EspritoSanto were considered adults
when they were longer than 610 mm SVL, while adult females were longer than 740 mm SVL (K. Kasperoviczus,
pers. comm.). We tested for the normality and homoscedasticity of samples using Kolmogorov-Smirnov and
Bartlett tests, respectively. We used Students t tests for morphological comparisons, when normal distributions
assumption applied (p-value > 0.05), with RTL and RHL as dependent variables. For non normal distributions we
applied Kruskal-Wallis' test.

512 Zootaxa 4097 (4) 2016 Magnolia Press BARBO ET AL.

 Geometric morphometric analysis. Samples available for this analysis included a total of 78 males and 84
females from the mainland populations of B. jararaca (N = 134) and Ilha dos Franceses (N = 28). Only the right
side of the head of each specimen was photographed with a camera Nikon D700 coupled to an AF-S VR Micro-
Nikkor 105 mm f/2.8G IF-ED lenses, fixed in a copystand with a standardized distance from the camera body of
550 mm. Images were digitalized with landmarks type I (Bookstein 1991) using tpsDig2 software (Rohlf 2005).
Sixteen landmarks were recognized in the right plane. Landmarks 1 to 5 refer to the region of the eye, 6 to 8 to the
loreal pit, 9 and 10 to the rostral region, and 11 to 16 to the mouth. (Fig. 1). We performed a Canonical Variate
Analyses (CVA) separately for mature males and females, with MorphoJ (Klingenberg 2011) and R (R CoreTeam
2013) softwares. The CVA was performed with the residuals of the regression between the shape and centroid size
aiming to correct the allometric differences.

Results

Our morphological comparisons using linear morphometrics identified a set of characters that distinguished the
population of Ilha dos Franceses from the mainland populations of B. jararaca and from all three insular species of
the group (B. alcatraz, B. insularis, and B. otavioi). We further identified significant shape differences between the
population from Ilha dos Franceses and the mainland populations of B. jararaca by performing a landmark-based
geometric morphometric analysis of head shape in a total of 162 specimens. Results from our analyses are provided
below.

Linear morphometric analyses

Our linear morphometric analyses distinguished the population of Ilha dos Franceses from the three insular species
of the group and the closest mainland population of B. jararaca.
The population from Ilha dos Franceses showed the following differences to the closest population of B.
jararaca from mainland Esprito Santo (Table 1): smaller adult size in males (H = 30.29, p < 0.001) and in females
(H = 29.14, p < 0.001); lower number of ventral scales in males (t = -2.803, df = 54.92, p = 0.006) and lower
numbers of subcaudals in females (H = 3.43, p = 0.064); relatively longer tail (TL/SVL) in males (t = 2.9075, df =
42.98, p = 0.006); relatively smaller heads (HL/TR) in males (t = 2.525, df = 38.223, p = 0.01).
The population from Ilha dos Franceses differed from Bothrops alcatraz by the following morphometric
characteristics (Table 1): larger adult size in males; higher number of ventral scales in males and females; higher
number of subcaudals in males and in females; relatively longer tail in males and in females; relatively smaller
head in males. We did not present statistical comparisons because all vouchers of B. alcatraz were lost in 2010
during the fire of herpetological collection of Instituto Butantan. Therefore, comparisons of measurments and
counts were made using data available in Marques et al. (2002).
The population from Ilha dos Franceses differed from Bothrops otavioi by the following morphometric
characteristics (Table 1): larger adult size in males (H = 14.819, p = 0.0001), and in females (H = 7.286, p = 0.007);
higher number of ventrals in males and females; higher number of subcaudals in males and females; relatively
smaller head in males (H = 15.418, p < 0.001), and females (H = 12.265, p <0.001).
Finally, the population from Ilha dos Franceses differed from Bothrops insularis mainly by its grayish-
brownish coloration vs. pale or yellowish in B. insularis; smaller adult size in males (t = -3.266, df = 27.362, p =
0.003), and females (H = 21.452, p < 0.001); higher number of ventral scales in males (H = 52.744, p < 0.001), and
females (H = 58.064, p < 0.001); higher number of subcaudal scales in males (H = 49.145, p < 0.001), and females
(H = 44.829, p < 0.001); relatively smaller tail in males (H = 13.28, p = 0.0002), and females (H = 4.504, p = 0.03);
relatively smaller head in males (H = 37.606, p < 0.001), and females (H = 39.917, p < 0.001).

NEW BOTHROPS FROM ATLANTIC COAST OF BRAZIL Zootaxa 4097 (4) 2016 Magnolia Press 513
 7$%/(6HOHFWHGPHDVXUHPHQWVPHULVWLFDQGHFRORJLFDOYDULDEOHVIRUWKHILYHVSHFLHVEHORQJLQJWRWKHBothrops jararacaVSHFLHVJURXS/HJHQG69/ 6QRXWYHQWOHQJWK&/ &DXGDOOHQJWK
+/ +HDGOHQJWK57/ 5HODWLYHWDLOOHQJWK5+/ 5HODWLYHKHDGOHQJWK9( 9HQWUDOVFDOHV6& 6XEFDXGDOVFDOHV$' $QWHULRUGRUVDOV0' 0LGERG\GRUVDOV3' 3RVWHULRUGRUVDOV
,62 ,QWHUVXSUDRFXODUVVFDOHV6/ VXSUDODELDOV,/ LQIUDODELDOV7WLS FRQVSLFXRXVWDLOWLSLQMXYHQLOHV0HDVXUHPHQWVLQPLOOLPHWHUVPPDQGVDPSOHVL]HLQSDUHQWKHVLV$GGLWLRQDOGDWDIRUB.
alcatrazDQGB. insularisZHUHREWDLQHGLQ0DUTXHVet al DQG$PDUDO 

Bothrops jararaca 
 Bothrops sazimai VSQRY Bothrops alcatraz* Bothrops insularis** Bothrops otavioi
PDLQODQG(VStULWR6DQWR

 PDOHV IHPDOHV PDOHV IHPDOHV PDOHV IHPDOHV PDOHV IHPDOHV PDOHV IHPDOHV

               

514 Zootaxa 4097 (4) 2016 Magnolia Press

69/ 
               
PP
n  n  n  n  n  n  n  n  n  n 

             

&/  
             
PP  
n  n  n  n  n   n  n  n 

              

+/  
           
PP  
n  n  n  n  n  n  n  n 

         
57/              
n  n  n  n    n  n  n  n 

         
5+/              
n  n  n  n     n  n  n 
n 
         
9(               
n  n  n  n  n  n  n  n   n   n 
        continued on the next page

BARBO ET AL.
 7$%/(&RQWLQXHG        
 Bothrops jararaca 
Bothrops sazimai VSQRY Bothrops alcatraz* Bothrops insularis** Bothrops otavioi
PDLQODQG(VStULWR6DQWR
 PDOHV IHPDOHV PDOHV IHPDOHV PDOHV IHPDOHV PDOHV IHPDOHV PDOHV IHPDOHV

     

   
6&            
n  n  n  n 
n  n  n  n  n  n 

NEW BOTHROPS FROM ATLANTIC COAST OF BRAZIL

    
$'  
n  n  n 

   

0'  
n  n  n 

  

3'  
n  n  n 

    

,62
n  n  n  n  n 

n n n  n n n n n n n n n  n n n 
6/
n n n  n n n   n n  n n 

n n n  n n n 

n n 
n n  n n n  n n  n n 
,/ n n 
n n n  n n n  n  n n 
n 
n  n n n 

Zootaxa 4097 (4) 2016 Magnolia Press

515
Geometric morphometric analysis

Our geometric morphometric analysis identified significant shape differences between the population of Ilha dos
Franceses and the remaining mainland populations of Bothrops jararaca distributed throughout the range of the
species. Indeed, the points of dispersion in the CVA did not overlap in insular and mainland populations, with the
ellipses of 95% confidence intervals overlapping only marginally in males (Fig. 1). The first canonical axes of CVA
explained 67.6% and 86.1% of variation for males and females, respectively. When compared with mainland
populations of B. jararaca, specimens of Ilha dos Franceses presented shorter distances between landmarks 11 to
16 and longer distances between landmarks 12 and 15 (Fig. 1), indicating a smaller head length and higher head
height at the level of the eyes, respectively. Landmarks 15 were distributed farther away from each other in
specimens of Ilha dos Franceses, instead of closer to each other as in all mainland populations, reflecting the larger
eyes in the new species. Landmarks 23, and 9 showed that canthals and the tip of the snout were projected
downward in the population of Ilha dos Franceses, whereas mainland populations had a snout projected upward.
Specimens from Ilha dos Franceses also differed from the mainland populations of B. jararaca in respect to their
rostral region that appeared to be sunk (landmarks 9 to11), and the modified size of their loreal pit that were nearer
to the nostril (landmarks 67 relative to 9) (Fig. 1).
In synthesis, our quantitative analyses (linear and geometrical) allowed us to recognize the population from
Ilha dos Franceses as a new species, described below.

Bothrops sazimai sp. nov.

Figs. 24

Bothrops jararacaCampbell & Lamar, 2004. Venomous Reptiles of the Western Hemisphere. Vol. 1, 1476:391. (in part).

Holotype. An adult male, MZUSP 22228, collected by our team on May 14, 2013, at Ilha dos Franceses
(2055'36"S, 4045'15"W), municipality of Itapemirim, Itaoca beach, state of Esprito Santo, Brazil (Figs. 23).
Paratypes. Fourteen specimens: IBSP 86673 (female), IBSP 8667475 (males), MBML 3319 (male with
hemipenis prepared), MBML 332021 (females), MBML 3322 (male), MBML 3323 (female), MZUSP 22229
(male), MZUSP 22230 (female), MZUSP 22231 (male), MZUSP 22232 (female), ZUEC 3383 (male), ZUEC 3384
(female).
Diagnosis. Bothrops sazimai is distinguished from other species of B. jararaca group by the following
combination of characters: (1) larger eyes; (2) shorter and higher head; (3) slender body; (4) relative longer tail; (5)
dorsum predominantly grayish and/or brownish; (6) venter creamish white, speckled in gray; (7) postorbital stripes
with the same color of lateral saddles; (8) usually two postoculars; (9) 2225 interictals; (10) 2024 anterior
dorsals; (11) 2023 midbody dorsals; (12) 198214 ventrals in females, 193206 in males; (13) 5465 subcaudals
in females, 6270 in males.
Bothrops sazimai differs from the mainland populations of B. jararaca by its smaller adult size (SVL), relative
longer tail (RTL), slender body, and larger eyes (Fig. 1; Table 1). The new species can be easily distinguished from
the other three species of the B. jararaca group (B. alcatraz, B. insularis, and B. otavioi) by a higher number of
ventrals and subcaudals (Table 1). It further differs from B. insularis by its grayish or brownish ground color
pattern, smaller length of adults, relative smaller head, and relative smaller tail. The new species is also
distinguished from B. alcatraz and B. otavioi, by its larger adult size, presence of conspicuous yellowish tail tip in
juveniles, and additionally from the latter species by the presence of two postoculars and higher number of
intersupraocular scales (Table 1).
Description of the holotype. Adult male; total length 738 mm; SVL 630 mm; TL 108 mm (17% of total
length); TR 604.1 mm; head length 25.9 mm (0.49% of trunk); head width 14.9 mm; mass 52.3 g (after draining
preservative). Rostral 2.9 mm wide, 4.2 mm high; nasal divided anterior and posterior to nostril; loreal trapezial
and single; prefoveals 2/2; postfoveals 2/2; prelacunal fused with second supralabial forming lacunolabial on both
sides; preoculars 2/2; postoculars 2/2; supralabials 8/8; interoculabials 3/2; circumorbitals 6/6; temporals 5/6;
infralabials 10/10, first three pairs contacting chin shields; four gulars between chin shields and first ventral scales;
six rows of gulars separating first ventral scales from 8th infralabial; canthals 2/2; five anterior intercanthals; four

516 Zootaxa 4097 (4) 2016 Magnolia Press BARBO ET AL.

FIGURE 1. Canonical analysis of females (A) and males (B) of Bothrops jararaca species group. Limits of variation in head
shape were defined by 95% confidence ellipses and computed by parametric bootstrap. Blue = population from Ilha dos
Franceses; Red = population of B. jararaca from mainland portions of the state of Esprito Santo; Green = other mainland
populations of B. jararaca, from the states of So Paulo, Paran, Santa Catarina, and Rio Grande do Sul.

NEW BOTHROPS FROM ATLANTIC COAST OF BRAZIL Zootaxa 4097 (4) 2016 Magnolia Press 517
FIGURE 2. Dorsal (A), lateral (B), and ventral (C) views of head of the holotype of Bothrops sazimai (MZUSP 22228). Scale
bar = 10 mm.

518 Zootaxa 4097 (4) 2016 Magnolia Press BARBO ET AL.

FIGURE 3. Dorsal (above) and ventral (below) views of the holotype of Bothrops sazimai (MZUSP 22228). Scale bar = 20
mm.

NEW BOTHROPS FROM ATLANTIC COAST OF BRAZIL Zootaxa 4097 (4) 2016 Magnolia Press 519
FIGURE 4. Active specimens of Bothrops sazimai at night in different macrohabitats on the island. Over fallen trunks and
twigs (A); coiled over trees (B) (MZUSP 22230); two individuals close to one another and moving on the ground (C); on the
rocks near the sea (D); coiled on the ground (MZUSP 22232) (E). Photographs by R. Sawaya (A, E), R. Zorzal (B), F. Barbo
(C), and T. Portillo (D).

posterior intercanthals; rows of anterior, central and posterior intersupraoculars 7/7/12; interrictals 22; dorsals
reducting posteriorly 21/21/17; ventrals 196; cloacal plate single; divided subcaudals 64. Posterior cephalic scales
longer than wide and strongly keeled; intersupralabials scales weakly keeled; temporal scales keeled; internasals,
canthals, and supraoculars smooth.
Coloration in life was grayish on dorsal surface with 15/14 lateral trapezoidal markings (saddles) irregularly
defined, dark brown-gray with well-defined borders, weakly pale-grayish edged, opposite and alternate to each
other in middle of dorsum; dorsum of head grayish, spotted with two small well-defined dark blotches between
occipitaltemporal portion and neck (Figs. 23); postorbital stripe is dark brown, bordered below by a thin white
line. Extends from behind eye, covering superior portion of 6th, 7th and 8th supralabials, up to three scales long
behind rictual region and three scales downwards to ventral direction; gular region mostly whitish-creamish, with

520 Zootaxa 4097 (4) 2016 Magnolia Press BARBO ET AL.

infralabials and symphysial speckled of gray; venter mostly whitish anteriorly, speckled in gray posteriorly and
covering cloacal plate; tail grayish dorsally, covered with small dark gray lateral blotches, with subcaudals
speckled in ventral plan.
Variation (Table 1). Dorsum of head can vary from entirely brownish or grayish, with or without brownish
stripes and blotches between supraoculars and the neck. Second supralabial fused with prelacunal forming the
lacunolabial (n = 23), partially fused (n = 6), or separated (n = 6). Besides differences in pholidosis, adult males
have longer tail than females (Table 1; H = 15.55, p < 0.001).
Hemipenis (n = 8). There is no evident variation in hemipenial morphology regarding shape and
ornamentation. Fully everted and maximally expanded organ moderately bilobed, subcylindrical, and bicapitate;
small and medium ossified spines covering proximal region of hemipenis; hemipenial body and intralobular region
asymmetrically covered by spinules on proximal portion, and medium and large spines on median and distal
portions on both sulcate and asulcate sides of organ; ossified spinules bordering sulcus spermaticus up to
capitulum, excepting croach; border of proximal calyces forming capitulum spinulate; sulcus spermaticus
bifurcating at level of croach and extending to tips of lobes (Fig. 5).

FIGURE 5. Sulcate (left) and asulcate (right) views of the hemipenis of B. sazimai (MBML 3319 paratype). Scale bar = 5 mm.

Distribution. The new species is known only from the type-locality, Ilha dos Franceses, (Fig. 6). This island
has about 15 ha with maximum elevation of 36 m asl covered by secondary Atlantic forest remnants (Ferreira et al.
2007), and located 3.6 Km eastwards from Itaoca beach (Fig. 7).
Natural History. Bothrops sazimai is abundant in Ilha dos Franceses as we have found approximately two
snakes per hour-person by visual search. The beginning of daily activity was observed in late afternoon. Specimens
were observed coiled (n = 13), moving (n = 8) or stationary (n = 4), on the ground (n = 15), as well as in lower
portions of shrubs and trees (n = 9) (Fig. 4). Except for the five collected individuals (holotype and four paratypes),
all snakes were observed and/or photographed and released in the field. Juveniles and adults of the new species

NEW BOTHROPS FROM ATLANTIC COAST OF BRAZIL Zootaxa 4097 (4) 2016 Magnolia Press 521
feed on ectothermic prey. Twenty-six out of 58 individuals examined in scientific collections had prey remnants in
their stomach and gut contents, including lizards (Gymnodactylus darwinii and Hemidactylus mabouia, n = 14),
centipedes Scolopendromorpha (n = 6), and a conspecific snake (n = 1). Youngs have yellowish tail-tip (n = 13),
suggesting caudal-luring behavior (see Andrade et al. 2010; Sazima 1991). The smallest mature male with enlarged
testes and opaque efferent ducts had 451 mm SVL, whereas the smallest mature females with follicles or embryos
in the oviduct measured 551 mm SVL (K. Kasperoviczus, pers. com.).

FIGURE 6. Coastal region of the state of Esprito Santo (above), with the Ilha dos Franceses (below), the type locality of
Bothrops sazimai. Legend of states: Bahia (BA); Esprito Santo (ES), Minas Gerais (MG), and Rio de Janeiro (RJ).

522 Zootaxa 4097 (4) 2016 Magnolia Press BARBO ET AL.

FIGURE 7. Aerial view of Ilha dos Franceses (A); forest remnants in the light-house area (B); vegetation on rocks of the tide
zone (C); dense vegetation predominant in island (D); view of Ilha dos Franceses from Itaoca Beach (E). Photographs by
Google Earth Pro (A), F. Barbo (B, C, D), and R. Sawaya (E).

NEW BOTHROPS FROM ATLANTIC COAST OF BRAZIL Zootaxa 4097 (4) 2016 Magnolia Press 523
 Etymology. The specific epithet sazimai represents a patronymic name dedicated to the friend and professor
Ivan Sazima, for his invaluable contribution to the study of natural history and conservation of Brazilian fauna, and
for his inspiring and pioneering field studies on Bothrops jararaca. Professor Ivan advised and mentored various
generations of students and researchers that presently work with diverse systems and organisms, including snakes,
amphibians, fishes, mammals, birds, and plants. He published hundreds of scientific contributions, among articles,
book chapters, and educational texts. We suggest the standard English name Franceses island-Lancehead for the
new species.

Discussion

Lanceheads of the Bothrops jararaca species group have been collected in continental islands along the Brazilian
coast since the 1910's. These were initially determined as B. jararaca due to their general morphological
similarities. An exception at that time was Bothrops insularis, which was described as a distinct species due to its
outstanding differences in coloration and ecological attributes when compared to the remaining species of the B.
jararaca complex (Amaral 1921). In the past few decades, studies on natural history and ecology within this group
of Lanceheads allowed to refine our understanding of its hidden ecological diversity (Martins et al. 2008; Andrade
et al. 2010; Marques et al. 2012, 2013; Guimares et al. 2014). However, the taxonomic status of several isolated
populations of B. jararaca remains poorly understood (but see Marques et al. 2002; Barbo et al. 2012). Recent
studies showed that two additional insular populations, previously recognized as belonging to B. jararaca, are
actually distinct species (Marques et al. 2002; Barbo et al. 2012). Bothrops sazimai represents the fourth island
endemic to be described for the group.
Island species usually show different patterns of body size and growth regime when compared to their related
mainland species (Foster 1964). Van Valen (1973) termed the phenomenon related to gigantism and dwarfism in
insular populations of mammals as the island rule (see also Lomolino 2005, 2012). According to this rule, large
mainland species become dwarf on islands and small mainland species become giant on islands. Some studies have
demonstrated that snakes also follow this rule, a phenomenon that could be triggered by reduced prey availability
and/or reduced prey size (Case 1978; Keogh et al. 2005), as suggested by the diet alteration hypothesis sensu
Boback (2003). The diet alteration hypothesis, related to island dwarfism in snakes, could explain the distinct body
size patterns observed in the insular members of the B. jararaca complex. All three island species of the group
present change on diet, because none of those islands retain natural populations of small mammals, the main prey
item of adult B. jararaca (Sazima 1992).
Bothrops insularis does not have any tendency to dwarfism or gigantism, and its diet resembles that of
mainland populations, with juveniles feeding upon ectothermic prey whereas adults became bird specialists,
shifting their diet towards the largest prey available in the island. Bothrops insularis also shows some variation in
body form that is apparently related to its more arboreal habits (see Martins et al. 2001). Bothrops alcatraz and B.
otavioi, on the other hand, are considered dwarf forms within the B. jararaca complex, the former feeding mainly
on centipedes, but also on lizards, while the latter preys upon frogs (Marques et al. 2002; Barbo et al. 2012).
Although Bothrops sazimai is larger than B. alcatraz and B. otavioi, it is definitely smaller than B. insularis and the
mainland populations of B. jararaca, and is considered here another dwarf form that feeds on ectothermic prey
(lizards, centipedes, and conspecific snakes). These three dwarf forms show a body form similar to that of young
mainland specimens of B. jararaca that also feed exclusively on ectothermic prey. We could then consider this
repeated and parallel evolution of dwarf forms as an evidence of the diet alteration hypothesis, and one possible
causal mechanism related to the well-known island rule pattern.
We observed a significant number of specimens (n = 13) of Bothrops sazimai in the field that were coiled on
the soil with their head directed to the base of tree trunks in which a large number of centipedes and small lizards
could also be observed. Since prey-type selection is known to have a strong effect on the evolution of head size
(Vanhooydonck et al. 2007), we consider that the overall larger eyes and more ventrally directed and shorter nostril
of B. sazimai (Fig. 1) may represent adaptations to this specific microhabitat and prey items. Small ectothermic
prey could be probably more effectively detected and subjugated by snakes with larger eyes and smaller heads,
with more anteriorly positioned sensorial organs. Although most field observations related to foraging B. sazimai
were made on specimens located on the ground, it is likely that the species also forages on the trees since nine

524 Zootaxa 4097 (4) 2016 Magnolia Press BARBO ET AL.

specimens were observed at 1.52 meters above the ground, inside the forest. Additionally, B. sazimai shows a
relatively longer tail length when compared to B. alcatraz and B. otavioi, a feature that is associated to arboreal
habits. Finally, the two lizards that are preyed by B. sazimai are arboreal species, and are the dominant prey found
in gut contents of preserved specimens.
Marine oscillations of the Brazilian coast are already known and mapped, with the intervals of time of
retractions and transgressions well defined (Martin et al. 1986). The islands of Alcatrazes, Queimada Grande, and
Vitria are located about 35 to 40 km off the coast of the state of So Paulo, whose depths varying between 4060
m. During the Pleistocene, between 12,00010,000 ya, the sea level was ca. 60 m lower (Martin et al. 1986),
indicating that these islands and the mainland were connected in this period. These pleistocenic oscillations of sea
level are the most acceptable hypothesis for separation between the mainland B. jararaca-like ancestor and these
closely related island species (see Rohling et al. 1998). For B. sazimai this process has probably occurred more
recently when comparing to the islands mentioned above.
Due to the short distance between the mainland and Ilha dos Franceses (ca. 4 km), and the relatively low depth
of the sea in this region (ca. 4 m), island and mainland should be probably connected during the medium Holocene
(ca. 3,8004,000 ya; Martin et al. 1986; Massad et al. 1996), when the sea level was ca. 46 m lower than observed
nowadays. Even so, this short time could be enough to promote some notable modifications in morphology and
ecology of the new species. These modifications could be produced by a strong genetic drift derived from the
genetic bottleneck caused by the insularization process, or instead by a strong selection driven by the restrictions of
the insular environment (absence of small mammals, population density, etc.). Alternatively, other factors as
heterochrony or more specifically allometry can be playing an important role in shaping the morphology of B.
sazimai. We claim that a more detailed study on this subject is needed to better understand the evolutionary process
involved in such speciation events.
The probable recent isolation of Bothrops sazimai (ca. 3,8004,000) in Ilha dos Francesessuggested by the
cycles of the marine oscillations of the Brazilian coastcan bring different sort of problems for the methods of
species delimitation based on the coalescent process (Maddison & Knowles 2006; Knowles & Carstens 2007). As
shown by Grazziotion et al. (2006) and Barbo et al. (2012) there is no genetic difference between the insular
species of the B. jararaca group and the mainland populations of B. jararaca. These authors also shown that B.
insularis, B. alcatraz and B. otavioi are nested within the B. jararaca mitochondrial lineages, presenting the same
mitochondrial haplotype found in the mainland B. jararaca. This mitochondrial DNA pattern indicates the
existence of shallow polymorphism among these species and suggests that the incomplete lineage sorting is
producing discrepancies between the gene tree and the species tree (Edwards 2008). Therefore, even not providing
genetic data for B. sazimai, we argue that the species probably does not present reciprocal monophyly in relation to
the mainland populations of B. jararaca. Based on the results of Grazziotin et al. (2006) and Barbo et al. (2012) we
also claim that sequencing a handful of loci will not offer sufficient molecular evidence for helping in the
delimitation of this species. Consequently, we based the description on morphological characters and the unique
combination of characters states (Davis & Nixon 1992) that makes B. sazimai diagnosable.
Island endemics are unique entities, evolving apart from mainland populations under different time and
evolutionary scenarios. Bothrops alcatraz and B. insularis were included in IUCN red-lists (see Marques et al.
2004a,b), and were included in a pioneering initiative of protection proposed by the governmental institution
ICMBio. This program was developed exclusively for the study of threatened species, and aimed to provide
detailed information on their natural history, ecology and population dynamics that would be used in management
and conservation strategies. In our opinion, such program should be extended to B. sazimai that faces potential
threats due to the short distance of the Ilha dos Franceses to the shore, and the presence of a small beach, which
encourage a constant transit of tourists into the island. The continuous flux of visitation results in uncontrolled
littering and even occasional fires, increasing the risk of pollution and destruction of significant portions of the
island. Those threats could affect drastically or even drive to extinction this unique insular species.
The encounter rate of Bothrops sazimai is about 15 snakes per day of visual search, and could be compared to
the one observed in Ilha da Queimada Grande for B. insularis (between 15 and 33 snakes a day; Martins et al.
2008), thus representing a second case of highly unusual density of an insular Lancehead population along the
Brazilian coast. According to IUCNs definitions (CR B1a,b [iii]), B. sazimai falls in the criterion of a "critically
endangered species" since it is endemic to an area with less than 100 km2 (0,15 km2, Figs. 67). Therefore,
Brazilian government should define a protected area that would encompass Ilha dos Franceses. The new species

NEW BOTHROPS FROM ATLANTIC COAST OF BRAZIL Zootaxa 4097 (4) 2016 Magnolia Press 525
could be included in the next future to the global and national red lists as Critically Endangered as well as in ex-situ
conservation actions.

Acknowledgments

We are grateful to K. Kasperoviczus for providing reproductive data for the new species and B. insularis, to G.
Sanches (MZUSP) and J. Paulo (MBML) for their support in respective collections, and to H.T. Pinheiro, T.E.
Simon (in memorian), C. Neto, T. Guedes, J.T. Portillo, and R. Zorzal for their help in the field. FEB and FGG
benefitted from a post-doctoral fellowship from the Fundao de Amparo Pesquisa do Estado de So Paulo
(FAPESP grant numbers 2012/09156-0 and 2012/08661-3, respectively). This research was supported by grants
from Fundao de Amparo Pesquisa do Estado de So Paulo (BIOTA/FAPESP 2011/50206-9 and FAPESP 2008/
54472-2) and Conselho Nacional de Desenvolvimento Cientfico e Tecnolgico (CNPq 565046/2010-1, 303545/
2010-0) to HZ and (CNPq 305911/2012-0) to RJS. RJS also thanks FADA-UNIFESP for financial support. This
research was carried out in strict accordance with federal laws in Brazil. Collecting permits of holotype and four of
paratypes were provided by the Instituto Chico Mendes de Conservao da Biodiversidade (ICMBio; Permit
numbers 148582 and 422701).

References

Amaral, A. (1921) Contribuio para o conhecimento dos ophidios do Brasil. Parte I. Quatro novas espcies de serpentes
Brasileiras. Memrias do Instituto Butantan, 1, 137.
Andrade, D.V., Marques, O.A.V., Gavira, R.S.B., Barbo, F.E., Zacariotti, R.L. & Sazima, I. (2010) Tail luring by the golden
lancehead (Bothrops insularis), an island endemic from southeastern Brazil. South American Journal of Herpetology, 5,
175180. http://dx.doi.org/10.2994/057.005.0302
Arajo, M.S. & Martins, M. (2006) Defensive behavior in pitvipers of the genus Bothrops (Serpentes, Viperidae).
Herpetological Journal, 16, 297303.
http://dx.doi.org/10.1643/0045-8511(2003)003[0081:BSEISE]2.0.CO;2
Barbo, F.E., Marques, O.A.V. & Sawaya, R.J. (2011) Diversity, natural history, and distribution of snakes in the municipality of
So Paulo. South American Journal of Herpetology, 6, 135160.
http://dx.doi.org/10.2994/057.005.0302
Barbo, F.E., Grazziotin, F.G., Sazima, I., Martins, M. & Sawaya, R.J. (2012) A new and threatened species of lancehead from
Southeastern Brazil. Herpetologica, 68, 418429.
http://dx.doi.org/10.1655/HERPETOLOGICA-D-12-00059.1
Boback, S.M. (2003) Body size evolution in snakes: evidence from island populations. Copeia, 2003, 8194.
http://dx.doi.org/10.1643/0045-8511(2003)003[0081:BSEISE]2.0.CO;2
Bookstein, F.L. (1991) Morphometric tools for landmark data: geometry and biology, Cambridge: Cambridge University Press.
New York, 485 pp.
Campbell, J.A. & Lamar, W.W. (1989) The Venomous Reptiles of Latin America. Cornell University Press. Ithaca, 425 pp.
Campbell, J.A. & Lamar, W.W. (2004) The Venomous Reptiles of the Western hemisphere. Vol. 1. Cornell University Press.
Ithaca, 476 pp.
Carrasco, P.A., Mattoni, C.I., Leynaud, G.C. & Scrocchi, G.J. (2012) Morphology, phylogeny and taxonomy of South American
bothropoid pitvipers (Serpentes,Viperidae). Zoologica Scripta, 41, 115.
http://dx.doi.org/10.1111/j.1463-6409.2011.00511.x
Case, T.J. (1978) A general explanation for insular body size trends in terrestrial vertebrates. Ecology, 59, 118.
http://dx.doi.org/10.2307/1936628
Davis, J.I. & Nixon, K.C. (1992) Populations, genetic variation, and the delimitation of phylogenetic species. Systematic
Biology, 41, 421435.
http://dx.doi.org/10.1093/sysbio/41.4.421
Edwards, S.V. (2008) A smrgsbord of markers for avian ecology and evolution. Molecular Ecology, 17, 945946.
http://dx.doi.org/10.1111/j.1365-294X.2007.03644.x
Fenwick, A.M., Gutberlet, R.L. Jr., Evans, J.A. & Parkinson, C.L. (2009) Morphological and molecular evidence for phylogeny
and classification of South American pitvipers, genera Bothrops, Bothriopsis, and Bothrocophias (Serpentes: Viperidae).
Zoological Journal of the Linnean Society, 156, 617640.
http://dx.doi.org/10.1111/j.1096-3642.2008.00495.x
Ferreira, A.L., Coutinho, B.R., Pinheiro, H.T. & Thomaz, L.D. (2007) Composio florstica e formaes vegetais da Ilha dos
Franceses, Esprito Santo. Boletim do Museu Biolgico Mello Leito, 22, 2544.

526 Zootaxa 4097 (4) 2016 Magnolia Press BARBO ET AL.

Foster, J.B. (1964) Evolution of mammals on islands. Nature, 202, 234235.
http://dx.doi.org/10.1038/202234a0
Grazziotin, F.G., Monzel, M., Echeverrigaray, S. & Bonatto, S.L. (2006) Phylogeography of the Bothrops jararaca complex
(Serpentes: Viperidae): past fragmentation and island colonization in the Brazilian Atlantic Forest. Molecular Ecology, 1
14.
http://dx.doi.org/10.1111/j.1365-294X.2006.03057.x
Guimares, M., Mungua-Steyer, R., Doherty, P.F. Jr., Martins M. & Sawaya, R.J. (2014) Population dynamics of the critically
endangered golden lancehead pitviper, Bothrops insularis: stability or decline? Plos One, 9, e95203.
http://dx.doi.org/http://dx.doi.org/10.1371/journal.pone.0095203
IUCN (2015) The IUCN Red List of Threatened Species. Version 2015-3. Available from: http://www.iucnredlist.org.
(Accessed 15 October 2015)
Keogh, J.S., Scott, I.A.W. & Hayes, C. (2005) Rapid and repeated origin of insular gigantism and dwarfism in Australian tiger
snakes. Evolution, 59, 226233.
http://dx.doi.org/10.1111/j.0014-3820.2005.tb00909.x
Klingenberg, C.P. (2011) MorphoJ: an integrated software package for geometric morphometrics. Molecular Ecology
Resources, 11, 353357.
http://dx.doi.org/10.1111/j.1755-0998.2010.02924.x
Knowles, L.L. & Carstens, B.C. (2007) Delimiting species without monophyletic gene trees. Systematic Biology, 56, 887895.
http://dx.doi.org/10.1080/10635150701701091
Lomolino, M.V. (2005) Body size evolution in insular vertebrates: generality of the island rule. Journal of Biogeography, 32,
16831699.
http://dx.doi.org/10.1111/j.1365-2699.2005.01314.x
Lomolino, M.V. (2012) Of mice and mammoths: evaluations of causal explanations for body size evolution in insular
mammals. Journal of Biogeography, 39, 842854.
http://dx.doi.org/10.1111/j.1365-2699.2011.02656.x
Maddison, W.P. & Knowles, L.L. (2006) Inferring phylogeny despite incomplete lineage sorting. Systematic Biology, 55, 21
30.
http://dx.doi.org/10.1080/10635150500354928
Marques, O.A.V., Martins, M. & Sazima, I. (2002) A new insular species of pitviper from Brazil, with comments on
evolutionary biology and conservation of the Bothrops jararaca group. Herpetologica, 58, 303312.
http://dx.doi.org/10.1655/0018-0831(2002)058[0303:ANISOP]2.0.CO;2
Marques, O.A.V., Martins, M. & Sazima, I. (2004a) Bothrops insularis. The IUCN Red List of Threatened Species 2004,
e.T2917A9493475.
http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T2917A9493475.en
Marques, O.A.V., Martins, M. & Sazima, I. (2004b) Bothrops alcatraz. The IUCN Red List of Threatened Species 2004,
e.T46344A11050246.
http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T46344A11050246.en
Marques, O.A.V., Kasperoviczus, K.N. & Almeida-Santos, S.M. (2013) Reproductive ecology of the threatened pitviper
Bothrops insularis from Queimada Grande island, Southeast Brazil. Journal of Herpetology, 47, 393399.
http://dx.doi.org/10.1670/11-267
Marques, O.A.V., Pereira, D.N., Barbo, F.E., Germano, V.J. & Sawaya, R.J. (2009) Os Rpteis do municpio de So Paulo:
diversidade e ecologia da fauna pretrita e atual. Biota Neotropica, 9, 139150.
http://dx.doi.org/10.1590/S1676-06032009000200014
Marques, O.A.V., Martins, M., Develey, P.F., Macarro, A. & Sazima, I. (2012) The golden lancehead Bothrops insularis
(Serpentes: Viperidae) relies on two seasonally plentiful bird species visiting its island habitat. Journal of Natural History,
46, 885895.
http://dx.doi.org/10.1080/00222933.2011.654278
Martin, L., Mrner, N.A., Flexor, J.M. & Suguio, K. (1986) Fundamentos e reconstruo de antigos nveis marinhos do
Quaternrio. Boletim do Instituto de Geocincias, Publicao Especial, 4, 1161.
Martins, M., Marques, O.A.V. & Sazima, I. (2002) Ecological and phylogenetic correlates of feeding habits in Neotropical
pitvipers of the genus Bothrops. In: Schuett, G., Hoggren, M. & Greene, H.W. (Eds.), Biology of the Vipers. Eagle
Mountain Publishing, Utah, 307328 pp.
Martins, M., Sawaya, R.J. & Marques, O.A.V. (2008) A first estimate of the population size of the critically endangered
lancehead, Bothrops insularis. South American Journal of Herpetology, 3, 168174.
http://dx.doi.org/10.2994/1808-9798(2008)3[168:AFEOTP]2.0.CO;2
Martins, M., Araujo, M.S., Sawaya, R.J. & Nunes, R. (2001) Diversity and evolution of macrohabitat use, body size and
morphology in a monophyletic group of Neotropical pitvipers (Bothrops). Journal of Zoology, 254, 529538.
http://dx.doi.org/10.1017/S0952836901001030
Massad, F., Suguio, K. & Perez, F.S. (1996) Propriedade geotcnica de sedimentos argilosos como evidncia de variaes do
nvel relativo do mar em Santos. In: Congresso Brasileiro de Geologia de Engenharia, 8., Rio de Janeiro, 1996. Anais. Rio
de Janeiro, ABGE, v. 1, pp. 163176.

NEW BOTHROPS FROM ATLANTIC COAST OF BRAZIL Zootaxa 4097 (4) 2016 Magnolia Press 527
R Development Core Team (2013) R: a language and environment for statistical computing. R Foundation for Statistical
Computing, Vienna, Austria. Avaliable from: http://www.R-project.org/ (Accessed 31 March 2016)
Rohlf, F. (2005) TpsDig. Digitize landmarks and outlines. Ver. 2.05. Department of Ecology and Evolution. State University of
New York at Stony Brook, New York.
Rohling, E.J., Fenton, M., Jorissen, E.J., Bertrand, P., Ganssen, G. & Caulet J.P. (1998) Magnitudes of sea-level lowstands of
the past 500,000 years. Nature, 394, 162165.
http://dx.doi.org/10.1038/28134
Sabaj Prez, M.H. (2014) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an
Online Reference. Version 5.0 (22 September 2014)
American Society of Ichthyologists and Herpetologists, Washington, DC. Available from: http://www.asih.org (accessed 01
October 2015)
Sazima, I. (1991) Caudal luring in two Neotropical pitvipers, Bothrops jararaca and B. jararacussu. Copeia, 1991, 245248.
http://dx.doi.org/10.2307/1446274
Sazima, I. (1992) Natural history of the jararaca pitviper, Bothrops jararaca, in southeastern Brazil. In: Campbell, J.A. &
Brodie, E.D. (Eds.), Biology of Pitvipers. Selva, Tyler, 199216 pp.
Uetz, P. & Hoek, J. (2015) The Reptile Database. Available from: http://www.reptile-database.org. (accessed on 15 September
2015)
Van Valen, L. (1973) Pattern and the balance of nature. Evolutionary Theory, 1, 3149.
Vanhooydonck, B., Herrel, A. & Van Damme, R. (2007) Interactions between habitat use, behaviour and the trophic niche of
lacertid lizards. In: Reilly, S.M., McBrayer, L.D. & Miles, D.B. (Eds), Evolutionary Consequences of Foraging Mode in
Lizards. Cambridge University Press, Cambridge, 427449 pp.
Wied-Neuwied, M. Prinz zu. (1824) Abbildungen zur Naturgeschichte Brasiliens. Isis von Oken, 14, 1103.

APPENDIX. Material examined.

Countries are given in bold capitals, states in upper capitals, municipalities in italics, and localities in plain text.
Bothrops alcatraz (n = 8). BRAZIL: SO PAULO: So Sebastio, Ilha dos Alcatrazes (IBSP 13031, 13126, 13183,
55578, 56133, 61853; MZUSP 1453, 21640).
Bothrops insularis (n = 26). BRAZIL: SO PAULO: Itanham: Ilha da Queimada Grande (IBSP 666, 683, 686, 1731,
1857, 1866, 1871, 1881, 1888, 1890, 1892, 1900, 1911, 1925, 192829, 1932, 1939, 1944, 1946, 196768, 1970, 1975, 1984,
2006).
Bothrops jararaca (n = 154). BRAZIL: ESPRITO SANTO: Alfredo Chaves: Vila Nova Maravilha (MBML 1921);
Anchieta (MBML 2272); Aracruz (MBML 468); Cariacica: Pitanga (MBML 1918); Domingos Martins: So Paulinho do
Arac (MBML 1716); Fundo (MBML 502); Guarapari (MBML 2005); Stio Modesto, Lameiro (MBML 1769); Ibirau:
Fazenda Morro das Palmas, Picuan (MBML 2900); Ibitirama: Pedra Rocha, Parque Estadual do Capara (MBML 1770, 1777,
1779); Itarana: Barra do Sossego (MBML 1691), Centro (MBML 716), Crrego Penedo (MBML 2101), Limoeiro de Santo
Antnio (MBML 2850); Linhares (MBML 1788; MZUSP 4448), Reserva Biolgica de Sooretama (MBML 2124, 2127),
Reserva Florestal da CVRD (MZUSP 5865; ZUEC 1788); Marechal Floriano (MBML 732), Stio Amarildo (MBML 1985),
Stio Trs Marias (MBML 73335, 752, 79194, 1791), Stio Zamprogno (MBML 1771, 1790); Nanuque (MZUSP 4459);
PARNA Capara: (MZUSP 14401); Santa Leopoldina (MZUSP 1491; ZUEC 118990), Encantado (MBML 280), Rio da Prata
(MBML 341, 346), Santo Antnio (MBML 38183, 417, 458); Santa Maria de Jetib (MBML 156; MZUSP 13171), Baixo
Rio Posmauser (MBML 1980), Rio Nove (MBML 397, 429, 45153); Santa Teresa: (MZUSP 5114), Aparecidinha (MBML
49, 1281, 2015), road to Santa Lucia (MBML 485), Goiapaba Au (MBML 86566), Museu Biolgico Mello Leito (MBML
469), Patrimnio de Santo Antnio (MBML 446, 474), Rio Bonito (MBML 123334, 1138), So Loureno (MBML 85, 563),
Stio Duas guas, Valo de So Pedro (MBML 720), Stio Max Loureiro Penha (MBML 497); Vale de So Pedro (MBML
2848), Valsugana Velha (MBML 1277); Serra: Centro Industrial de Vitria (ZUEC 543), Lagoa Humait, Carapebus (MBML
1925), Stio Gasparini, Carapebus (MBML 550, 1776, 192627); Vila Velha: Barra do Jacu (MBML 2014); Vitria (MBML
1772), Fonte Grande (MBML 2215), Restinga do Camburi (MBML 1917), Viveiro de Mudas da CST (MBML 1916);
PARAN: Castro (UFRGS 1377); Colnia Ouro Verde (UFRGS 1845); Cruz Machado (UFRGS 1694); Mallet (UFRGS
1640); Moreira Sales (UFRGS 1769); Palmas (UFRGS 1367, 1632); Porto Unio (UFRGS 1828); Porto Unio da Vitria
(UFRGS 168788, 1697, 1699, 1739, 1825, 1837); Rio Azul (UFRGS 1767); Unio da Vitria (UFRGS 1390, 1829, 1851,
1853); Vales (UFRGS 1827, 1849); Vera Guarani (UFRGS 1711); RIO GRANDE DO SUL: Campo Bom (UFRGS 219);
Canela (UFRGS 1764); Carlos Barbosa (UFRGS 1368); Caxias do Sul (UFRGS 1702); Erechim (UFRGS 1631); Gramado
(UFRGS 1373); Gravata (UFRGS 1461); So Leopoldo (UFRGS 1463); Torres (UFRGS 1755); Viamo (UFRGS 1751); SO
PAULO: Bertioga (MZUSP 4637, 12382), Guaratuba (MZUSP 732425); Guaruj (MZUSP 358384); Iguape: Barra do
Ribeira (MZUSP 406263); Ilhabela (MZUSP 4054); Juquitiba (MZUSP 1277071); Miracatu (MZUSP 12107); Mogi das
Cruzes (MZUSP 2265); Perube (MZUSP 12852); Salespolis: Estao Biolgica de Boraceia (MZUSP 446668, 488384,
11576); Santo Andr: PNMN Paranapiacaba (MZUSP 17938); So Jos do Barreiro: Serra da Bocaina (MZUSP 4910); So
Miguel Arcanjo: Parque Estadual Carlos Botelho (MZUSP 1524142, 17202); So Sebastio (MZUSP 140304), Barra do Una
(MZUSP 13167), Praia do Engenho (MZUSP 15137), Praia de Juquehy (MZUSP 12819); SANTA CATARINA: Canoinhas

528 Zootaxa 4097 (4) 2016 Magnolia Press BARBO ET AL.

(UFRGS 1846); Criciuma (UFRGS 1453); Garopaba (UFRGS 3487); Itaiopolis (UFRGS 1830); Lucerna (UFRGS 1710);
Porto Belo (UFRGS 239); So Bento do Sul (UFRGS 1884, 1895); So Domingos (UFRGS 6529).
Bothrops otavioi (n = 31). BRAZIL: SO PAULO: Ilhabela municipality-archipelago: Ilha da Vitria (IBSP 1886669,
1887182, 78572 holotype); MZUSP 3949, 395152, 557785; ZUEC 355051).
Bothrops sazimai (n = 34). BRAZIL: ESPRITO SANTO: Itapemirim: Ilha dos Franceses: IBSP 8667677 (females),
MBML 3318 (male with everted hemipenis), MBML 332425 (females), MBML 332627 (males), MZUSP 2228587
(females), MZUSP 22288 (male with everted hemipenis), MZUSP 22289 (males), MZUSP 22290 (female), MZUSP 2229192
(males), MZUSP 22293 (female), MZUSP 22294 (male with everted hemipenis), MZUSP 2229596 (females), MZUSP 22297
(male), MZUSP 2229899 (females), MZUSP 22300 (male), MZUSP 2253439 (females), MZUSP 2254043 (males),
MZUSP 22544 (female). The following nine specimens were only measured and photographed in the field, but not collected:
RJS 5001 (male), RJS 5002 (male), RJS 5003 (female), RJS 5004 (female), RJS 5007 (male), JMF 39 (male), JMF 40 (female),
JMF 4243 (males).

NEW BOTHROPS FROM ATLANTIC COAST OF BRAZIL Zootaxa 4097 (4) 2016 Magnolia Press 529