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Accepted by P. Passos: 20 Mar. 2018; published: 10 May 2018
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4418 (5): 469
–
480
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Article
469
https://doi.org/10.11646/zootaxa.4418.5.4
http://zoobank.org/urn:lsid:zoobank.org:pub:D6EF6500-C1D8-4742-B5BC-268B3856C95D
Morphological variation of the rare psammophilous species Apostolepis gaboi
(Serpentes, Dipsadidae, Elapomorphini)
THAÍS B. GUEDES
1,4
, FAUSTO E. BARBO
2
, DANIELLA FRANÇA
2,3
& HUSSAM ZAHER
2
1
University of Gothenburg, Gothenburg Global Biodiversity Center and Department of Biological and Environmental Sciences, Box
461, SE-405 30, Göteborg, Sweden.
2
Museu de Zoologia, Universidade de São Paulo, Av. Nazaré 481, Ipiranga, 04263-000, São Paulo, SP, Brazil.
3
Programa de Pós-graduação em Ciências Biológicas, Instituto de Biociências, Universidade Estadual Paulista, Av. 24
a
1515, 13506-
900, Rio Claro, SP, Brazil.
4
Corresponding author. E-mail: thaisbguedes@yahoo.com.br
Abstract
Apostolepis gaboi was described based only on the holotype found in the Queimadas, state of Bahia, northeastern Brazil.
Since its original description, no additional specimens were reported in literature and the species was considered to be rare
and poorly known. Here, we provide a detailed description for the species based on the examination of the holotype and
34 additional specimens from the type locality and adjacent areas. Additional information is also provided on intraspecific
color variation and hemipenial morphology.
Key words: Caatinga, color pattern, endemism, hemipenial morphology, intraspecific variation
Resumo
Apostolepis gaboi foi descrita com base apenas no holótipo, encontrado em Queimadas, estado da Bahia, nordeste do Bra-
sil. Desde sua descrição, nenhum espécime adicional foi relatado na literatura e a espécie era considerada rara e pouco
conhecida. Fornecemos aqui uma descrição detalhada para a espécie com base no exame do holótipo e de 34 espécimes
adicionais oriundos da localidade-tipo e áreas adjacentes. Fornecemos também informações sobre a variação de coloração
intraespecífica e morfologia hemipeniana.
Palavras-chave: Caatinga, endemismo, morfologia hemipeniana, padrão de coloração, variação intra-específica
Introduction
The dipsadid snake genus Apostolepis Cope 1862 includes 35 species distributed throughout cis-Andean South
America (Uetz et al. 2018). The genus is considered to be monophyletic within the tribe Elapomorphini (Grazziotin
et al. 2012), being characterized by fused internasal and prefrontal shields, and by the presence of a black band on
the tip of the tail (Boulenger 1896; Ferrarezzi 1993; Lema 2001).
The lack of a taxonomically comprehensive review of the group since Strauch (1885) makes it difficult to
assess the taxonomic identity of most populations of Apostolepis (Rodrigues 1992). As result, available keys for
species identification (e.g., Peters & Orejas-Miranda 1970; Lema 2001; Ferrarezzi et al. 2005) have proven to be of
limited accuracy or restricted value, forcing taxonomists to undergo tenuous, geographically restricted comparisons
each time a new specimen needs identification (Vanzolini 1986). Furthermore, there are limitations imposed by the
homogeneous scale pattern known to occur throughout the group and the paucity of available samples in
collections for several currently recognized species (see below).
The last few years witnessed an unusually high number of species descriptions based on a single specimen that
are ultimately recognized solely by their color pattern (e.g., A. gaboi Rodrigues 1992; A. phillipsi Harvey 1999; A.
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christineae Lema 2002; A. cerradoensis Lema 2003; A. striata Lema 2004a; A. tertulianobeui Lema 2004b; A.
freitasi Lema 2004c; A. serrana Lema & Renner 2006; and A. roncadori Lema 2016). Along with these, a number
of older descriptions based on single specimens [e.g., A. longicaudata Gomes in Amaral 1921; A. vittata (Cope
1887)] help increase substantially the apparent state of chaos that characterizes the taxonomy of the genus.
However, recent efforts that clarified the identity of some of these species were welcome (e.g., Lema et al. 2005;
Lema & Renner 2005a,b; Curcio et al. 2011).
As for most species in the genus, Apostolepis gaboi was described based only in the holotype. Its secretive
psammophilous habits and apparently restricted distribution to the quaternary sand dunes of the left bank of the São
Francisco River (Rodrigues 1992; Guedes et al. 2014) reinforced the status of rare species. According to IUCN’s
definitions, A. gaboi was ultimately ranked in the Brazilian Red List of Threatened Species as Endangered (EN),
under the criteria B1ab(iii) (ICMBio 2016). However, during a survey in the zoological collections of Brazil, we
found a large number of specimens of A. gaboi that were collected at the type locality or close to it, within the sand
dunes region. Here, we provide additional information on meristic variation, color pattern, and hemipenial
morphology to the species.
Materials and methods
We examined a total of 35 specimens of Apostolepis gaboi, including its holotype, housed in the following
Brazilian collections: Museu de Zoologia, Universidade de São Paulo (MZUSP), São Paulo; Museu de Zoologia,
Universidade Federal da Bahia (MZUFBA), Salvador; and Museu de Zoologia, Universidade Estadual Feira de
Santana (MZUEFS), Feira de Santana. The specimens and localities are listed in the Appendix.
We analyzed characters from external quantitative (meristic and morphometric data) and qualitative traits
(pholidosis and color pattern) and hemipenial morphology. We took measurements to the nearest 0.01 mm with a
digital caliper, except for the total length (snout-vent length plus tail length), which were taken with a flexible ruler.
We converted most measurements to proportions for purpose of descriptions. The biometric information of
intraspecific variation comprises minimum, maximum and average ± single standard deviation, based on the 35
specimens examined (15 females and 20 males, including the holotype; Table 1). We performed a non-parametric
Kruskal-Wallis analysis to test for secondary sexual dimorphism in the sample (Zar 1999). We followed Dowling
(1951) method for the counting segmental scales. We prepared hemipenis following procedures described by
Pesantes (1994) for retracted or semi-everted organs from preserved specimens. We colored the calcified structures
of the hemipenis with red dye Alizarine S. We followed the terminology for the hemipenial morphology from
Dowling & Savage (1960), Zaher (1999), and Zaher & Prudente (2003). We obtained the images with a digital
camera or layer-by-layer camera and mounted in plates using Photoshop CS6 version 13.0. We retrieved
coordinates of localities based on direct visual inspection of layers (municipality, São Francisco river, and villages)
in the software QGIS version 2.14.10 (QGIS Development Team 2017) and Google Earth. We produced a
geographical distribution map in QGIS on the basis of refined coordinates.
Results
Redescription of the holotype of Apostolepis gaboi Rodrigues 1992 (Figs. 1–2, Table 1). Described based on a
single specimen (MZUSP 10290) from Queimadas (10° 23’ S, 42° 30’ W; 295 m a.s.l.), municipality of Pilão
Arcado, state of Bahia, Brazil. A slim juvenile female 215 mm SVL, 20 mm TL (8.5% of total length); body
diameter 2.5 mm (1.2% SVL); head length 5.64 mm (2.4% SVL), head width 2.26 mm at broadest point (40.1%
head length); head not distinct from neck; dorsal scales rows 15/15/15; ventrals 229, divided cloacal plate; paired
subcaudals 31; rostral plate slightly wider (0.84 mm) than long (0.66 mm), visible in dorsal view; prefrontals large,
slightly wider (0.98 mm) than long (0.89 mm), in broad median contact, shorter than frontal scale; frontal longer
(1.31 mm) than wide (1.01 mm), anteriorly exceeding anterior border of supraoculars; parietals very long (2.77
mm), almost two times longer than frontal scale; parietal laterally contacting 5
th
and 6
th
supralabials, in contact
anteriorly with frontal, supraocular and postocular, posteriorly with first rows of dorsals slightly increased;
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MORPHOLOGICAL VA RIATION OF APOSTOLEPI S GABOI
FIGURE 1. Head views (left, right, dorsal and ventral, respectively) (A), dorsal (B) and ventral body views (C), tail views
(dorsal and ventral, respectively), and dorsal midbody view (E) of the holotype of Apostolepis gaboi (MZUSP 10290) from
Queimadas, municipality of Pilão Arcado, state of Bahia, Brazil.
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FIGURE 2. General view of the holotype of Apostolepis gaboi (MZUSP 10290) in life from Queimadas, municipality of Pilão
Arcado, state of Bahia, Brazil. Photo by Miguel Rodrigues.
supraoculars smaller than frontal, with their anterior margin very straight; nasal plate entire, longer than high,
thinner posteriorly, in contact with rostral, prefrontal, 1
st
and 2
nd
supralabials; nostril centralized in nasal plate;
preocular one, very small; postocular one; supralabials six, 2
nd
and 3
rd
contact the eye, 5
th
and 6
th
increased in length;
temporal scales indistinct, parietal in contact with 5
th
and 6
th
supralabials; mental triangular, as long as high,
separated from chinshields by first pair of infralabials; chinshields in two pairs, second being longer (1.05 mm)
than first (0.68 mm); first pair of chinshields in contact with first four infralabials, second pair with 4
th
and 5
th
infralabials.
Coloration of holotype faded after preservation in 70% ethanol. Head dark brown, except for the snout light
cream; one cream blotch extending from 3
rd
to 5
th
supralabial scales; nostrils cream (Fig. 1A); neck with complete
white collar (three scales long) in dorsal and lateral views followed by dark brown collar (two to three scales long);
gular region cream anteriorly with incomplete dark brown band after chinshields (Fig. 1A); dorsum of body pale
brown or beige with five narrow longitudinal brown stripes (one vertebral and four dorsolateral lines on the 4
th
and
6
th
rows) extending until the tip of the tail (Fig. 1B and E); ventral surface of body immaculate cream from the neck
to subcaudals, except for the tip of the tail (Fig. 1C); tip of the tail dark brown in dorsal (12 scales long) and ventral
(11 scales long) views; terminal spine dark brown (Fig. 1D).
Pictures of Apostolepis gaboi in life (Fig. 2), show it has dorsum of the head dark brown; snout pale brown;
distinct oval blotch pale brown covering the 3
rd
to 5
th
supralabials; nuchal collar white, followed by dark brown to
black cervical collar; five black stripes arising in the black collar following to up the tail; vertebral and paraventral
stripes are narrowest while paravertebrals are widest; dorsal background is rust-colored; tip of the tail is black and
terminal spine is also black.
Intraspecific variation in continuous characters (Table 1). Largest female 517 mm total length (TL), 31 mm
of tail length; largest male has 424 mm and 38 mm. Total length 236–517 mm (mean = 349.3 ± 95.95 mm; n = 15)
in females, 234–424 mm (mean = 299.7 ± 37.08 mm; n = 20) in males. Tail length 18–35 mm (mean = 25.33 ± 5.44
mm; 12.13–21.25% TL) in females, 16–40 mm (mean = 29.3 ± 5.52 mm; 7.63–18.50% TL) in males; ventral scales
200–239 (mean = 221.6 ± 11.32) in females, 190–230 (mean = 208.7 ± 9.18) in males; subcaudals 27–34 (mean =
30.95 ± 4.2) in males, 24–31 (mean =27.4 ± 2.35) in females. We found a significant secondary sexual dimorphism
in the number of ventrals (H = 9.21; p < 0.001; n = 35) and subcaudals (H = 16.07, p < 0.001; n = 35); supralabials
six (2
nd
and 3
rd
contacting the eye). Preocular absent (n = 9), 1/1 (n = 23) or 1/0 (n = 2). Preocular usually fused to
prefrontals, except for the MZUFBA 1693 where it seems to be fused with the 2
nd
supralabial. Second pair of
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chinshields contacting 4–5 infralabials, except for the MZUFBA 1697 in which the first pair of chinshields contact
first three infralabials and the secondary chinshields 3–4 infralabials.
FIGURE 3. Color pattern variation of the preserved specimens of Apostolepis gaboi. Dorsal (A) and lateral (B) views of the
head, lateral (C) and dorsal (D) views of the midbody, and ventral view of the tail (E).
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FIGURE 4. Sulcate (left), asulcate (center), and lateral (right) views of the fully everted and maximally expanded hemipenis of
Apostolepis gaboi (MZUFBA 1683).
Intraspecific variation in coloration. White nuchal collar cover two (MZUFBA 1674, 1676, 1680–81, 1685,
1689, 1694, 1702–03) to five scales long (MZUFBA 1673, 1680, 1690, and 1697), most specimens with three
scales long (MZUFBA 1675, 1677–79, 1682, 1684, 1686–88, 1691–93, 1695–96, 1698–1701, 1704) (Fig. 2A);
dark brown collar one and half (MZUFBA 1690, 1693, 1703) to three scales long (MZUFBA 1673–76, 1678,
1680–81, 1683–85, 1687, 1691, 1694, 1699) (Fig. 3A), contacting ventrals or starts on third dorsal scale row (Fig.
3B); dorsal coloration usually with vertebral stripe and two dorsolateral stripes each; dorsal stripes continuous,
dark brown, speckled with small dots and arranged between the 4
th
and 5
th
dorsal rows (Fig. 3C and D); eight
individuals with five stripes along the body (MZUFBA 1675, 1678, 1683, 1688, 1692, 1696–97, 1701, and 1703)
and 12 with five stripes plus a pair of faint dashed brownish lines at the level of the 3
rd
dorsal
row (MZUFBA 1679,
1689, 1691, 1695, and 1700); six specimens presented vestigial additional stripes (MZUFBA 1673, 1677, 1682,
1684, 1694, 1699, and 1702) (Fig. 3C and D); tail tip surrounded by dark brown band, including terminal spine;
caudal band varies in its extent, covering ventral region of two (MZUFBA 1704), four (MZUFBA 1687, 1689, and
1696), five (MZUFBA 1679), six (MZUFBA 1982), seven (MZUFBA 1684) or eight subcaudals (MZUFBA 1695)
(Fig. 3E).
Hemipenial morphology. Organ of adult specimen (MZUFBA 1683) fully everted and maximally expanded
(Fig. 4); hemipenis short, stout and unilobed, unicaliculate and unicapitate; lobe extends for about one third of the
hemipenis; capitulum restricted to sulcate and lateral surfaces of the organ, covering half of the organ’s length at
the level of the sulcus spermaticus; capitulum formed by small papillate calyces disposed concentrically on sulcate
and lateral surfaces of the lobe, except for two outermost rows of calyces that are clearly larger in size and
spinulate; on asulcate surface, lobes ornamented with two parallel rows of three to four deep, mediolaterally
enlarged and papillate body calyces; body of hemipenis covered by spinules on sulcate surface and proximal region
of asulcate surface; spinules reduce in number and increase gradually in size towards lateral surface and distal
region of asulcate surface of the body, where they form one outer and distal row of greatly enlarged spines; the
enlarged spines disposed asymmetrically on each side of asulcate or lateral surfaces; distally to the last row of
enlarged lateral spines, hemipenial body is completely nude, except for a median row of spinules arranged along
midline of asulcate surface; sulcus spermaticus forks within the base of the capitulum where it continues in almost
parallel portions to reach distal surface of lobe on its sulcate side.
Distribution and natural history notes. Apostolepis gaboi is a psammophilous species endemic of the
Caatinga ecoregion (Rodrigues 1992, 2003; Guedes et al. 2014) with a restricted distribution known to encompass
only the municipalities of Barra and Pilão Arcado, in the eastern sandy bank of the São Francisco River, state of
Bahia, Brazil (Fig. 5). The vegetation of the area is bushy and herbaceous Caatinga that grows on sandy soil and
dunes (Rodrigues 1992; Guedes et al. 2014). Apostolepis gaboi is both diurnal and nocturnal (Rodrigues 1992;
Guedes et al. 2014), with a probable feeding preference for elongated preys, as suggested by the only prey item
registered so far and identified as Amphisbaena hastata (Damasceno 2002).
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MORPHOLOGICAL VA RIATION OF APOSTOLEPI S GABOI
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MORPHOLOGICAL VA RIATION OF APOSTOLEPI S GABOI
FIGURE 5. Landscape view from the Quaternary sandy dunes in the São Francisco River, state of Bahia, northeastern Brazil
(A) and the map showing the distribution of Apostolepis gaboi in the left side of the São Francisco River (B). Photo A by
Miguel Rodrigues.
Discussion
Rodrigues (1992) described Apostolepis gaboi on the basis of a single specimen from Queimadas, in the Caatinga
biome of the state of Bahia, northeastern Brazil. Ferrarezzi et al. (2005) included Apostolepis gaboi in the
Apostolepis assimilis species group along with A. ammodites, A. assimilis, A. cearensis and A. arenaria. All these
species share external similarities such as the presence of white cervical collar followed by a black nuchal collar
and entirely black terminal spine. Most species of the A. assimilis group present dorsal color pattern uniformly red
(as in A. flavotorquata), while only A. arenaria and A. gaboi show dorsal color pattern with stripes (as in A.
longicaudata and A. nigrolineata). Among the species of the A. assimilis group, two of them (A. gaboi and A.
cearensis) show a higher degree of rostral prominence, which is a putative synapomorphy also shared with several
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species referred to A. dimidiata group (e.g., A. dimidiata, A. ambiniger, A. vittata, A. goiasensis) (see Ferrarezzi et
al. 2005).
The hemipenes of the species from the A. assimilis group are similar (Fig. 6). All of them are short, stout and
unilobed, being unicaliculate and unicapitate and with a narrower base. The capitulum reaches the sulcate and
lateral surface of the organs, covering almost half of its length until the level of the sulcus spermaticus. On its
asulcate surface, the lobe is ornamented with two parallel rows of three to four deep, midlaterally enlarged and
papillate body calyces. The hemipenial body is covered by spinules on the sulcate surface and the proximal region
of the asulcate surface. Spinules reduce in number and increase gradually in size towards the lateral surface and
distal region of the asulcate surface of the body, where they form one outer and distal row of greatly enlarged
spines. However, the hemipenis of A. gaboi seems to share more similarities with the organs from the A. assimilis.
These hemipenial conditions refereed above for the A. assimilis group are also present in the A. longicaudata
(Curcio et al. 2011) and, at least partially, in the A. quinquelineata (T. Guedes pers. observ.); suggesting that these
might have evolved independently in both groups or may be more widespread in the genus. Nevertheless, no
definitive evidence supports a close phylogenetic affinity of A. gaboi with species of the A. assimilis group, except
for the geographical proximity and overall morphological similarities pointed out by Ferrarezzi et al. (2005).
FIGURE 6. Sulcate (left), asulcate (center), and lateral (right) views of the fully everted and maximally expanded hemipenis of
Apostolepis ammodites (MZUSP 12069-A), A. arenaria (MZUSP 15573-B), A. assimilis (IB 52754-C), A. cearensis (MZUSP
19549-D), and A. gaboi (MZUFBA 1683-E).
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MORPHOLOGICAL VA RIATION OF APOSTOLEPI S GABOI
Acknowledgments
We are grateful to Rejâne Lira-da-Silva (MZUFBA) and Ilka Biondi (MZUEFS) for permission to examine
specimens under their care, to Nelson R. Albuquerque and Paulo Passos for valuable criticism in the first version of
the manuscript. TBG and DF were supported by scholarships from the Fundação de Amparo à Pesquisa do Estado
de São Paulo (FAPESP 2013/04170-8, 2014/18837-7, and 2015/05047-0). Funding for this research was provided
by Fundação de Amparo à Pesquisa do Estado de São Paulo to TBG (FAPESP 2014/18837-7) and to HZ (BIOTA/
FAPESP 2011/50206-9).
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APPENDIX. Material examined.
Country is given in bold capital, state in upper capital, municipalities in italics, and localities in plain text.
Apostolepis gaboi (n = 35). BRAZIL: BAHIA: Ibiraba, Barra (MZUFBA 1673–1704), Icatú, Barra (MZUEFS 981),
Queimadas, Pilão Arcado (MZUSP 10290, holotype; MZUSP 10547, topotype).
