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New Polychromatic Species of Atractus (Serpentes: Dipsadidae) from the
Eastern Portion of the Colombian Andes
Authors: Elson Meneses-Pelayo, and Paulo Passos
Source: Copeia, 107(2) : 250-261
Published By: American Society of Ichthyologists and Herpetologists
URL: https://doi.org/10.1643/CH-18-163
Downloaded From: https://bioone.org/journals/Copeia on 09 May 2019
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New Polychromatic Species of Atractus (Serpentes: Dipsadidae) from the
Eastern Portion of the Colombian Andes
Elson Meneses-Pelayo
1,2
and Paulo Passos
3
We describe herein a new polychromatic species of the snake genus Atractus from the cloud forests of the northeastern
Andes of Colombia. The new species is distinguished from all congeners by having an exclusive combination of phenotypic
characters, such as: dorsal scale rows 17, loreal long, seven to ten maxillary teeth, ventrals 156–174 in females and 153–
169 in males, subcaudals 20–30 in females and 23–30 in males, dorsum with variable coloration, changing from dark green
to orange or red with a black nuchal band (three to four scales long) connected to a black vertebral line and two black
dorsolateral continuous stripes from the occipital region to tip of the tail, venter with irregular black blotches, relatively
small body size, small tail length in females and moderately long in males, hemipenis moderately bilobed, semicapitate
and semicalyculate. We compared the new species with all congeners occurring along the Cordillera Oriental in Colombia,
Sierra de Perija
´in the Colombia/Venezuela frontier and Cordillera de M´
erida in Venezuela. We discussed aspects related
to polychromatism and its implication toward a robust taxonomy for the genus Atractus.
Describimos una nueva especie de serpiente policroma
´tica del g´
enero Atractus para los bosques nublados del noreste de
los Andes de Colombia. La nueva especie se distingue de todos los cong´
eneres por tener una combinaci ´
on exclusiva de
caracteres fenot´
ıpicos, como: escamas dorsales en 17 hileras, loreales largas, siete a diez dientes maxilares, ventrales
156–174 en hembras y 153–169 en machos, subcaudales 20–30 en hembras y 23–30 en machos, dorso con coloraci´
on
variable, cambiando de verde oscuro a naranja o rojo con una banda nucal negra (largo de tres o cuatro escamas)
conectada a una l´
ınea vertebral negra y dos franjas continuas dorsolaterales negras desde la regi ´
on occipital hasta la
punta de la cola, superficie ventral del cuerpo con manchas irregulares negras, tama˜
no corporal relativamente peque˜
no,
longitud de la cola peque˜
na en las hembras y moderadamente larga en los machos, hemipene moderadamente
bilobado, semicapitado y semicaliculado. Comparamos las nuevas especies con todos los cong´
eneres que ocurren a lo
largo de la Cordillera Oriental en Colombia, Sierra de Perija
´en la frontera de Colombia/Venezuela y la Cordillera de
M´
erida en Venezuela. Discutimos aspectos relacionados con el policromatismo y su implicaci ´
on hacia una taxonom´
ıa
s´
olida para el g´
enero Atractus.
THE cryptozoic snake genus Atractus is widely distrib-
uted in the Neotropical region, occurring from central
Panama to northeastern Argentina (Giraudo and
Scrocchi, 2000; Myers, 2003). Atractus comprises 140 cur-
rently recognized species making it the most species-rich
extant snake genus (Passos et al., 2018a), with the highest
number of currently recognized taxa in the trans-Andean
region of Colombia (Passos et al., 2009a). However, factors
such as the small body size and cryptozoic lifestyle may be
responsible for the relative paucity of samples for some
species available in collections (Schargel and Garc´
ıa-P´
erez,
2002; Myers, 2003; Myers and Donnelly, 2008; Prudente and
Passos, 2010). On the other hand, there are a lot of specimens
currently misidentified in collections and public repositories
(Passos et al., in press). To date, 25 species in Atractus are
recognized to occur in the highlands of the Cordillera
Oriental, Serran´
ıa de Perija
´, Cordillera de M´
erida in the
Colombia and Venezuela boundary, but many of them still
remain with uncertain taxonomic status. Although a com-
prehensive review for the species of this region is underway,
we anticipate herein the formal description of a new
polychromatic Atractus from the western slopes of the
Cordillera Oriental of Colombia.
MATERIALS AND METHODS
Specimens examined of the new species are housed in the
herpetological collection of the Universidad Industrial de
Santander, department of Santander, Colombia (MHN-UIS),
and comparative material is listed in the Material Examined.
Data from additional specimens of Atractus previously
examined by PP can be found in: Passos et al. (2005,
2007a, 2007b, 2009a, 2009b, 2009c, 2009d ‘‘2008’’, 2009e,
2010a ‘‘2009’’, 2010b, 2010c, 2012, 2013a, 2013b, 2013c,
2013d, 2016a, 2016b, 2017, 2018a, 2018b, 2018c), Passos
and Fernandes (2008), Passos and Arredondo (2009), Passos
and Lynch (2011 ‘‘2010’’), Passos and Prudente (2012),
Prudente and Passos (2008, 2010), Schargel et al. (2013),
Almeida et al. (2014), Salazar-Valenzuela et al. (2014), and de
Fraga et al. (2017).
Measurements were taken with a dial caliper to the nearest
0.1 mm under a stereomicroscope, except for snout–vent
length (SVL) and caudal length (CL), which were taken with
a flexible ruler to the nearest 1.0 mm. Terminology for
cephalic shields in Atractus follows Savage (1960), except for
the loreal that follows the proposal of Passos et al. (2007b).
Method of counting ventral and subcaudal scales follows
follows Dowling (1951). Condition of the loreal scale follows
Passos et al. (2007b). Measurements and descriptions of
paired cephalic scales are strictly based on the right side of
head. We describe the color pattern of the new species based
on live specimens or on its digital photographs taken directly
in the field, since several specimens were released after the
measurement of general biometric data. Sex was determined
based on presence or absence of hemipenes verified through
a ventral incision at the base of the tail. We examined
maxillae in situ under a Nikon C-LEDS stereomicroscope
through a narrow lateral-medial incision between the supra-
1
Grupo de Estudios en Anfibios y Reptiles de Santander, Universidad Industrial de Santander, Carrera 27, Calle 9, Bucaramanga, Colombia.
2
Grupo de Estudios en Biodiversidad, Universidad Industrial de Santander, Carrera 27, Calle 9, Bucaramanga, Colombia.
3
Departamento de Vertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, Rio de Janeiro, RJ 20940-040,
Brazil; Email: atractus@gmail.com. Send reprint requests to this address.
Submitted: 3 December 2018. Accepted: 11 February 2019. Associate Editor: W. L. Smith.
Ó2019 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CH-18-163 Published online: 9 May 2019
Copeia 107, No. 2, 2019, 250–261
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labials and the maxillary arch. After removing tissues
covering the maxillary bone, we counted teeth and empty
sockets. The method for preparation of preserved hemipenis
was modified from Pesantes (1994) in replacing potassium
hydroxide (KOH) with distilled water (see Passos et al.,
2016a), observing precautions highlighted by Zaher and
Prudente (2003). Prior to inflation with petroleum jelly, the
organs remained 15–20 min in a 70% ethanol solution with
Alizarin red to stain the ornamented calcareous structures
according to Uzzel (1973), with adaptations from Harvey and
Embert (2008) and Nunes et al. (2012). Terminology for
hemipenial descriptions follows Dowling and Savage (1960)
and Zaher (1999) with a few minor adaptations. We follow
Passos et al. (2009e) and Passos et al. (2010a ‘‘2009’’) with
respect to conditions of the morphological characters used in
diagnosis and description.
Because scales counts are known to be sexually dimorphic
in Atractus (Savage, 1960; Passos et al., 2005), we used the
Mann-Whitney U-test to evaluate statistical differences
between sexes. Previously, we evaluated the assumptions of
univariate normality and homoscedasticity with the Kolmo-
gorov-Smirnov’s and Levene’s tests, respectively (Zar, 1999).
We used the following characters in the statistical analysis:
number of ventral scales, subcaudal scales, snout–vent length
(SVL), tail length (TL), and SVL/CL ratio. Additionally, we
performed a discriminant function analysis (DFA) for males
and females separately to evaluate differentiation between
chromatic patterns with an a priori definition of operational
groups based on each morphotype (Manly, 2005). We used a
cross-validation matrix to obtain the posterior classification
probabilities in which each individual is allocated to its own
group. For this analysis, meristic and morphometric data
were used for adult males and females. The minimal body
size for sexually mature individuals was based on the study of
Gualdr´
on-Dura
´n et al. (in press). We used the packages
‘‘MASS’’ (Ripley et al., 2016) and ‘‘ellipse’’ (Murdoch and
Chow, 2013) for performing statistical analysis. We used the
packages ‘‘ggplot2’’ (Wickham, 2009) and ‘‘colorspace’’
(Ihaka et al., 2013) for producing the graphics. All these
packages are in the R software (R Core Team, 2018).
Atractus marthae, new species
urn:lsid:zoobank.org:act:E7B6E8EB-0183-4624-B46F-
9E51B2CD7E64
Martha’s Groundsnake
Culebras Tierreras de Martha
Figures 1–6, Table 1
Holotype.—UIS-R 3027, adult male, Colombia, department of
Santander, municipality of Santa Ba
´rbara, Vereda Esparta,
0780105.38 00 N, 72853043.0400 W, ca. 2400 m above sea level
(hereafter asl), E. Meneses-Pelayo, 23 October 2014.
Paratypes.—(111 specimens, all collected or legated by first
author from the department of Santander, Colombia):
Reserva el Diviso, 0780306.1600 N, 72859011.9700 W, ca. 2400
m asl, municipality of Piedecuesta: UIS-R 1734, 5 May 2007;
Vereda Potrero Grande, 06850046.9600 N, 72851020.6700 W, ca.
2220 m asl, municipality of Guaca: UIS-R 3248–49, 20
November 2015; Vereda Esparta, 0780105.3800 N,
72853043.04 00 W, ca. 2400 m asl, municipality of Santa
Ba
´rbara: IAvH 9046 (formerly UIS-R 3021), IAvH-R 9045
(formerly UIS-R 3020), IUCN-R 13168 (formerly UIS-R
3022), IUCN-R 13169 (formerly UIS-R 3024), MHUA
15283–84 (formerly UIS-R 3018–19), UIS-R 3017, UIS-R
3023, UIS-R 3025–26, UIS-R 3028–35, UIS-R 3054, 23
October 2014; UIS-R 3038–44, UIS-R 3053, UIS-R 3055, 20
December 2014; UIS-R 3061–74, 22 February 2015; UIS-R
3090–98, UIS-R 3100, UIS-R 3262–63, 14 March 2015; UIS-R
3108–14, UIS-R 3120–25, 25 April 2015; UIS-R 3138–44, 23
May 2015; UIS-R 3148–54, UIS-R 3156, 28 June 2015; UIS-R
3172–78, UIS-R 3193, 25 July 2015; UIS-R 3194–97, 29
August 2015; UIS-R 3198–3205, 19 September 2015; UIS-R
3251–55, 20 November 2015.
Diagnosis.—Atractus marthae is distinguished from all conge-
ners by unique combination of the following characters: (1)
dorsal scale rows smooth in 17/17/17; (2) postoculars two; (3)
long loreal; (4) temporals usually 1þ2; (5) supralabials usually
seven, third and fourth contacting the orbit; (6) infralabials
usually seven, first three contacting chinshields; (7) maxil-
lary teeth seven to ten; (8) gular scale rows usually four; (9)
preventrals usually four; (10) ventrals 156–174 in females,
153–169 in males; (11) subcaudals 20–30 in females, 23–30 in
males; (12) dorsum with variable coloration and pattern,
presenting green to red ground color with broad black
vertebral line (four scales wide) or narrow vertebral line
(one scale wide) and two dorsolateral bands; (13) ventral
ground color varying from cream with irregular black spots to
belly mostly black with a few cream blotches anteriorly,
usually posterior region of body and ventral surface of tail
uniformly black; (14) moderately long body size, females
reaching 346 mm SVL, males 307 mm; (15) small tail length
in females (7.9–10.2% of SVL), small to moderately long in
males (8.1–14.4% SVL); (16) hemipenis moderately bilobed
(lobe size equivalent to the capitulum length), semicapitated
and semicalyculated.
Comparisons.—Among all currently recognized species of
Atractus from the Cordillera Oriental of Colombia, Sierra de
Perija
´in the Colombia/Venezuela frontier and Cordillera de
M´
erida in Venezuela, the new species shared conspicuous
and regular longitudinal stripes (lacking paravertebral con-
nections with spots and/or transversal blotches) only with
Venezuelan species A. emigdioi,A. mariselae, and A. taphorni.
Atractus marthae differs from A. taphorni by having 17/17/17
dorsal scale rows and two dorsal-lateral stripes (vs. 15/15/15
rows of dorsal scales and absence of dorsal-lateral lines in A.
taphorni); differs from A. mariselae by having three longitu-
dinal stripes, 20–30 subcaudals in females and 23–30 in
males (vs. reticulated brown dorsal coloration with two
narrow dorsolateral lines; 31–33 subcaudals in females and
36–39 in males); differs from A. emigdioi by having belly
mostly black with irregularly distributed and few dispersed
square cream spots and hemipenis moderately bilobed,
semicapitated and semicalyculated (vs. belly with a broad
black central stripe with cream lateral lines on the para-
ventral region or with midline divided in two black
longitudinal stripes cream bordered; hemipenis slightly
bilobed, non-capitated and non-calyculated).
Considering the congeners with overlapping ranges of
distribution, Atractus marthae occur sympatrically or para-
patrically only with A. pamplonensis,A. variegatus, and A.
wagleri along the Cordillera Oriental of Colombia. The new
species differs from all of them by having dorsum with three
continuous and conspicuous dorsal-lateral stripes from the
neck to the tip of the tail (vs. variable color pattern but
lacking longitudinal lines or stripes in A. pamplonensis,A.
variegatus, and A. wagleri).
Meneses-Pelayo and Passos—New polychromatic Atractus from Colombia 251
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Fig. 1. General view of the holotype
of Atractus marthae, new species
(UIS-R 3027) in life.
Fig. 2. Lateral (A), dorsal (B), and ventral (C) views of head, and dorsal (D) and ventral (E) views of body of the holotype of Atractus marthae (UIS-R
3027) preserved in 70% ethanol.
252 Copeia 107, No. 2, 2019
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Description of the holotype.—Adult male, SVL 277 mm, TL 29
mm (10.5% SVL); head rounded in dorsal view, flattened in
lateral view, 8.8 mm long (3.2% SVL), 5.6 mm wide (63.6%
head length); cervical constriction indistinct; body sub-
cylindrical, body diameter 6 mm (2.2% SVL); belly flattened;
tail moderately long, with terminal spine moderately long
and acuminate; snout rounded in dorsal view, truncated in
lateral view; rostrum–orbit distance 3.1 mm (35.2% head
length); nasal–orbit distance 2.3 mm (26.1% head length);
intraorbital distance 3.5 mm (62.4% head width); rostral
subtriangular in frontal view, twice as wide (1.9 mm) as high
(0.9 mm), visible dorsally; internasal slightly wider (1.3 mm)
than long (1.0 mm); internasal suture (0.9 mm long) sinister
with respect to prefrontal median suture; prefrontal 2.2 mm
long, 1.9 mm wide; frontal subtriangular, as wide (2.6 mm) as
long (2.6 mm); supraocular subtrapezoidal, slightly longer
(1.5 mm) than wide (1.2 mm); parietal about twice as long
(4.0 mm) as wide (2.4 mm); nasal divided; prenasal 0.9 mm
high, 0.6 mm long, contacting rostral, internasal, first
supralabial, and postnasal; postnasal slightly higher (1.0
mm) than long (0.7 mm), contacting prenasal, prefrontal,
loreal, and first and second pair of supralabials; loreal twice as
long (1.6 mm) as high (0.8 mm); loreal contacting eyes,
prefrontals, nasals, and second and third supralabials; eye
diameter 1.3 mm; pupil round; two postoculars, upper
postocular about as high (0.9 mm) as long (0.7 mm); lower
postocular twice as high (0.9 mm) as long (0.4 mm); upper
postocular contacting eye, lower postocular, supraocular,
parietal, and anterior temporal; lower postocular contacting
eye, fourth and fifth supralabials, and anterior temporal;
temporal formula 1þ2; anterior temporal twice as long (1.6
mm) as high (0.8 mm), contacting parietal, fifth supralabial,
sixth supralabial, postocular, and posterior temporal; poste-
rior superior temporal about three times as long (3.4 mm) as
Fig. 3. General view in life of the four distinctive color patterns of Atractus marthae observed in the type-locality.
Meneses-Pelayo and Passos—New polychromatic Atractus from Colombia 253
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high (0.9 mm); temporal posterior inferior 1.5 mm long, 0.8
mm high; seven supralabials, third and fourth contacting
orbit; second supralabial higher than first and lower than
third; fifth and sixth supralabials with similar height,
seventh taller (1.4 mm) and longer (2.3 mm) than anterior
supralabials; symphysial subtriangular, twice as wide (1.6
mm) as long (0.7 mm); seven infralabials, first three pairs
contacting chinshields; first pair of infralabials in contact
behind symphysial, avoiding symphysial–chinshields con-
tact; chinshields more than twice as long (2.5 mm) as wide (1
mm); dorsal scale rows smooth, in 17/17/17 series; four rows
of gular scales between last supralabial and preventrals; four
preventrals; 163 ventrals; 26/25 (left/right) subcaudals.
Maxillary bone arched anteriorly in dorsal view and flattened
on the middle toward its end; maxilla with ten teeth
arranged linearly; teeth angular in cross section, robust at
the base, narrower at the apex, curved 45–608posteriorly;
first tooth anteriorly projected and more spaced from the
other first five teeth; four subsequent teeth similar in size and
slightly spaced; sixth to tenth teeth gradually decreasing in
size but increasing the spacing among teeth; no apparent
diastema: jaw with a lateral process strongly developed (Fig.
6).
After 17 months of storage in 70% ethanol after fixation in
10% formalin solution, dorsum of head dark brown with pale
brown spots covering medial-posterior portion of internasals,
prefrontals, supraoculars, and lateral-posterior area of parie-
tals; lateral surface of head dark brown to dorsal edges of
supralabials, except irregular cream spots on the prenasal and
postnasal; rostral with dark irregular spots dorsally; dark
Fig. 4. Dorsal and ventral views of
recently dead specimens (before
fixation in formalin solution) of Atrac-
tus marthae observed in the type-
locality.
Fig. 5. Sulcate (left) and asulcate (right) sides of the hemipenis of
Atractus marthae (UIS-R 3110). Scale bar ¼5 mm.
Fig. 6. Morphology of the maxillary arch of Atractus marthae (UIS-R
3061) in labial (top), ventral (middle), and lateral (bottom) views. Scale
bar ¼1 mm.
254 Copeia 107, No. 2, 2019
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brown descending postocular stripe covering two postocu-
lars, anterior-ventral portion of anterior temporal, posterior
half of fifth supralabial to anteriormost border of seventh
supralabial; remaining supralabials mostly cream, with upper
edges adjacent to orbit dark brown pigmented; cream
temporal region adjacent to descending postocular stripe;
temporal area scattered with diffuse brown dots, extending
from mid-posterior portion of anterior temporal to upper and
lower posterior temporals including immaculate seventh
supralabial; occipital region uniformly dark brown connected
dorsally to nuchal band (two to three scales long); gular
region cream with few dispersed brown dots covering first
pair of infralabials near chinshields suture, lateral portions of
second and third pairs of supralabials, and anterior chin-
shields; preventrals almost uniformly cream; ventral ground
color anteriorly (to the level of 17
th
) mostly cream with few
irregular black spots; from the 18
th
ventral belly mostly black
with a few square cream spots diffuse along body, except for
cream lateral portion of ventrals; cream spots concentrated
on the posterior region of body near the cloacal region;
ventral surface of tail mostly cream with lateral-anterior areas
of subcaudals pigmented with black, leaving the mid-ventral
suture region uniformly cream colored until 17
th
subcaudal;
posterior portion of tail cream with irregular black spots;
dorsal ground color of body light brown with dark brown
nuchal band connected to three dark brown dorsal-lateral
continuous stripes extending to tip of tail; vertebral line (one
scale wide) and two dorsal-lateral narrow stripe pigmenting
the suture between fourth and fifth scale rows (Fig. 2).
Color pattern variation in life.—(n¼112) Dorsum of head and
body with green, yellow, orange, red, or reddish brown
ground color; background coloration with continuous black
longitudinal stripes, constituted by vertebral (one to four
scales wide) and two dorsal-lateral lines (half-scale to two
scales wide); sometimes black stripes tenuously bordered
with yellow pigment; paraventral region (first three scales
rows) frequently with contrasting (lighter, Fig. 3A–B or
darker, Fig. 3C, F) background with respect to paravertebral
area; lower part of supralabials, gular region, and ventral
portion of head cream to yellow; belly frequently mostly
black scattered with square creamish yellow, orange, or red
spots or blotches; sometimes belly predominantly cream,
yellow, or red with few black spots or blotches; ventral marks
(lighter or darker) restricted to a single or a series of ventrals
or even to lateral region of each scale; usually anterior third
of body predominantly lighter (cream, yellow, or red) with
few black spots laterally concentrated; ventral surface of tail
almost entirely lighter (cream, yellow, or red) with a few
black dispersed spots to entirely black (Figs. 3–4).
Chromatic variability of the adult specimens.—(n¼92) A total
of 92 specimens (100%) were analyzed and grouped into
color categories as follows: Pattern A: back with thin, black
vertebral band (1–2 scales wide) and background greenish to
almost yellow (Fig. 3C, E, n¼36, frequency ¼39.1%); Pattern
B: back with wide, black vertebral band (3–4 scales wide) and
greenish to almost yellow background (Fig. 3F, n¼15,
frequency ¼16.3%); Pattern C: back with thin, black
vertebral band (1–2 scales wide) and orange-red background
(Fig. 3D, n¼29, frequency ¼31.5%); Pattern D: back with
wide, black vertebral band (3–4 scales wide) and orange-red
background (Fig. 3A, B, n¼12, frequency ¼13%). We
performed a discriminant analysis for each pattern and the
results for males and females do not show any structuration
with respect to a priori labels.
Sexual dimorphism.—We found significant differences in
body size (t¼–2.76, df ¼30, P,0.01), with females
presenting larger SVL (mean ¼276.7630.2, n¼37) than
males (mean ¼265.6 624.7, n¼32), and males presenting
larger tail size (t¼10.34; df ¼31; P,0.0001; mean ¼
31.463.2, n¼32) than females (mean ¼24.462.6, n¼32).
Similarly, in the meristic data we found that the males had a
lower number of ventral scales than the females (U
–6.7
¼33.5,
P,0.0001), while females have lower numbers of subcaudal
scales than males (U
–6.9
¼16.5, P,0.0001).
Hemipenial morphology.—(n¼7) Organ in situ (fully retracted)
extends and bifurcates at level of seventh to eighth
subcaudals; fully everted and maximally expanded hemi-
penis renders a moderately bilobed, semicapitate, and semi-
calyculate organ; lobes clavate with flattened tips; lobes with
similar size and moderately distinct from capitulum; lobes
with higher concentration of non-calcified calyces arranged
in transverse series; capitulum located at the level of
spermatic sulcus bifurcation; capitular groove well defined
on the asulcate side and indistinct on the sulcate side of
hemipenis; capitulum of equal size of hemipenial body on
both sides of organ; sulcus spermaticus bifurcates on the
middle part of organ with each branch centrifugally oriented,
running to tip of lobes; sulcus spermaticus margins well
defined and narrow, bordered by spinules from its base to
apices of lobes; hemipenial body uniformly covered by
Table 1. Meristic and morphometric variation from the type-series of Atractus marthae. Abbreviations are as follow: SVL ¼snout–vent length, TL ¼
tail length, BD ¼body diameter, HL ¼head length, HW ¼head width, and ED ¼eye diameter.
Character
Juvenile males Juvenile females Adult males Adult females
(n¼7) (n¼12) (n¼32) (n¼37)
Mean Range Mean Range Mean Range Mean Range
SVL 115.5 103–137 136.2 100–216 264.8 197–307 276.7 216–346
TL 13.4 11–15 13 10–16 31.4 22–35 24.2 19–29
TL/SVL x 100 11.6 9.5–14.3 9.7 7.8–14 11.2 8.1–14.4 8.8 7.9–10.2
BD 3.9 3.0–5.2 4.2 3.4–4.9 6.4 5.1–7.7 6.37 4.6–7.9
HL 6.3 5.8–6.7 6.4 6.0–6.9 8.0 7.0–9.0 7.9 6.3–9.3
HW 4.2 3.5–4.7 4.2 3.7–4.8 5.4 4.1–6.3 5.4 4.2–6.8
ED 1.0 0.8–1.1 0.9 0.8–1.0 1.2 0.9–1.4 1.2 0.9–1.4
Ventrals 160.1 157–162 164.4 157–171 160.4 153–169 169.3 166–174
Subcaudals 27.6 26–29 22.4 20–28 26.6 23–30 21.7 20–27
Meneses-Pelayo and Passos—New polychromatic Atractus from Colombia 255
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calcified hooked spines; large spines restricted to lateral sides
of organ; basal portion with longitudinal plicae and dispersed
spinules (Fig. 5).
Meristic and morphometric variation.—(n¼69) Largest male
307 mm SVL, 35 mm TL; largest female 346 mm SVL, 29 mm
TL; adult midbody diameter 2.5–2.6% (mean ¼2.55, SD ¼
0.69, n¼30) SVL in males, 2.2–2.3% (mean ¼2.25, SD ¼0.48,
n¼36) SVL in females; tail 8.1–14.4% (mean ¼11.9, SD ¼
1.12, n¼30) SVL in males, 7.9–10.2% (mean ¼8.7, SD ¼0.6,
n¼36) SVL in females; ventrals 153–169 (mean ¼160.4, SD ¼
5.43, n¼32) in males, 166–174 (mean ¼169.3, SD ¼2.99, n¼
37) in females; subcaudals 23–30 (mean ¼26.6, SD ¼1.4, n¼
32) in males, 20–26 (mean ¼21.7, SD ¼2.04, n¼37) in
females; preventrals 3–5 (mean ¼4, SD ¼0.37, n¼69); rows
of gular scales 3–5 (mean ¼3.9, SD ¼0.31, n¼69);
supralabials 6 (n¼3), 7 (n¼65), or 8 (n¼1); infralabials 6
(n¼1) or 7 (n¼68); infralabials contacting chinshields 3 (n¼
68) or 4 (n¼1); rows of dorsal scales at the level of the second
subcaudal 7–10 (mean ¼8.4, SD ¼0.8, n¼62); midbody
diameter 4.6–7.9 mm (mean ¼6.4, SD ¼0.8; n¼69);
maxillary teeth 6 (n¼2), 7 (n¼3), 8 (n¼25), 9 (n¼27), or 10
(n¼12; Table 1).
Etymology.—The new species is named herein in honor of
Martha Patricia Ram´
ırez-Pinilla for her invaluable contribu-
tion to the knowledge of the biology of Colombian
amphibians and reptiles. Furthermore, we would like to
acknowledge her dedicated vocation as a professor at the
Universidad Industrial de Santander, during which she has
contributed to the formation of many generations of
herpetologists.
Distribution and natural history.—Atractus marthae is known
from three localities in the western slope of Cordillera
Oriental, department of Santander, Colombia. These regions
are covered by cloud forest between 1990–2400 m asl (Fig.
7). We found specimens under rocks in wide areas of
paddocksbothinthetypelocalityandinthesouthernmost
locality in the path Potrero Grande (Fig. 8A), municipality
of Guaca. One specimen was collected near the Cerro de la
Jud´
ıa, Regional Natural Park in the municipality of Florida-
blanca. Many individuals at the type locality were found
crossing trails of well-conserved forested areas dominated by
Quercus humboldtii (Fig. 8B), in pastures of short grasses
(savannas) with high density of rocks, very humid soils, and
little arborization (Fig. 8C), as well as in areas of blackberry
crops.
Atractus marthae was found active during crepuscular
periods between 1700–1830 hrs, generally associated with
non-compacted soils with high humidity and with the
presence of abundant earthworms that could constitute
their main diet, as reported for other species of Atractus
(Balestrin et al., 2007). In periods of inactivity, the
specimens are found generally under rocks and under-
ground until 30 cm of depth; in such microhabitats were
also found litters of 3–5 eggs (Fig. 8D–E). These eggs were
collected and monitored in the laboratory under controlled
conditions until their hatching (temperature between 15–
188C, relative humidity between 80–100%). Neonates did
not vary in coloration with respect to adults (n¼12), and
within the same litter there were neonates with different
coloration patterns, displaying A–D morphotypes. We
observed reproductive behaviors in April with four males
and five females found under a rock forming a ‘‘mating
ball’’; when the rock was lifted, the snakes freed from the
ball and hid themselves underground. Males attained
maturity at a smaller size (137 mm SVL, n¼7) than females
(216 mm SVL, n¼12; Gualdr´
on-Dura
´netal.,inpress),as
reported for other congeners (Balestrin et al., 2005; Resende
and Nascimento, 2015; Passos et al., 2016a).
DISCUSSION
Of the 140 currently recognized species of Atractus, almost
ahalf(~70) occur in the northern Andes (Ecuador,
Colombia, and Venezuela), this being a major diversifica-
tion zone for the genus (Passos et al., 2009b). Of this great
Andean diversity and species richness, approximately 14%
of the species (10 spp.) are restricted (¼endemic) to the
Cordillera Oriental of Colombia from the south of the
Sumapaz highland (038520N, 748250W) north to the
municipality of Oca ˜
na (088140N, 73821 0W), department of
Norte Santander, Colombia. Atractus marthae occur sym-
patrically or parapatrically with A. pamplonensis, A. varie-
gatus, and A. wagleri. However, the new species is easily
distinguished from these congeners by having a very
distinctive coloration consisting of dorsum of body with
three continuous and conspicuous dorsal-lateral stripes
from the neck to the tip of the tail (vs. variable color
pattern but lacking longitudinal stripes in A. pamplonensis,
A. variegatus,andA. wagleri).
The systematized study of polychromatism has been
elusive for the genus Atractus and directed to species with
large series available in collections mostly from the cis-
Andean lowlands, such as: Amazonia (Passos and Prudente,
2012; Almeida et al., 2014; Passos et al., 2016b), Atlantic
Forest (Passos et al., 2010c), Cerrado (Passos et al., in press),
or Caatinga (Passos et al., 2016a). In many instances, the
available samples for Andean taxa do not allow us to access
reasonably the interspecific variation, making it difficult to
infer species boundaries among many hypothesized taxa (cf.
Esqueda and Lamarca, 2005). In contrast, the huge variability
displayed by several congeners has greatly impacted the
taxonomy of Atractus, with recognition of several junior
synonyms based essentially on color morphs or ontogenetic
phases inside of a given species (see Passos et al., 2009d
‘‘2008’’, 2018a, 2018c; Passos and Prudente, 2012).
Recently, Passos et al. (2016a) carried out an exhaustive
analysis of the variation in different phenotypic axes for
Atractus postchi and discussed some scenarios for the
evolution of the chromatic polymorphism in the genus. In
the case of the A. marthae, all color morphs occur in the same
population sampled from the type locality and such
variability does not show any sexual or ontogenetic basis.
These patterns are characterized mainly by continuous
longitudinal lines that vary in thickness, the patterns with
thin longitudinal lines being the most common (Pattern A ¼
39.1% and Pattern C ¼31.5%; see Results), followed by those
with broad stripes (Pattern B ¼16.3% and Pattern D ¼13%).
Despite the selective pressure involved in the maintenance of
polychromatism among several species of Atractus and the
paucity of detailed studies on this topic, it is clear that color
variance must be taken seriously into account before any
taxonomic decisions are made. As a rule for the genus, all
species with large series show great color variation, and the
studies based on a few specimens should compare them in
detail at least with all morphologically similar and sympatric
species.
256 Copeia 107, No. 2, 2019
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MATERIAL EXAMINED
Atractus acheronius: (n¼1) Venezuela: Zulia: Machiques de
Perija
´:R
´
ıo Negro Valley: MHNLS 398 (holotype of A.
acheronius).
Atractus crassicaudatus: (n¼455) Colombia: without locality:
IBSP 2443, ICN 8505, 8508–25, 8922–25, MLS 139, 152,
156, 293, 2640, MUJ 92, 355; Boyaca
´: Badohondo: ICN
10693, Bel´
en: ICN 10709*, Chiquinquira
´: MLS 2577,
Coper: MLS 2578–79, Duitama: ICN 10700–07, Garagoa:
ICN 10627, MUJ 315–22, 398–99, 509, Guayata
´: IAvH 864–
65, Pajarito: IAvH 1059, ICN 2608–11, 2831–33, Pesca:
IAvH 1880, R´
ıo Tectino: IAvH 799, Sogomoso: MLS 282,
2751–52, Tunja: MUJ 04, Ventaquemada: MLS 2243, Villa
de Leyva: IAvH 2172–73, 3039, 3189, 4788, 4811–20, 4852,
4878, 4889, 4892–93, 4912, 4960, 4976, ICN 2792*, 8332–
33, 9016–19, 9027, MLS 2021, 2564–65, 2918–20, Zipa-
quira
´: MUJ 05; Cundinamarca: without locality: MUJ 482;
Aguadita: MLS 169, Alba
´n: IAvH 4749, ICN 10626, Bogota
´:
IAvH 129, 204, 2478, ICN 1394–426*, 1455, 1460–61,
2588, 2623, 2633, 2641, 3377, 4217, 4240, 6209, 6236,
6340, 6449, 6490–91, 6504–05, 6509, 7100, 7102, 8260,
10806*, IBSP 226, 7216–17, 10164–67, 42945, MLS 153,
164–65, 167, 172, 178, 2546, 2607–09, 2614–15, 2617,
2644–45, MUJ 03, 07, 09–10, 17, 22, 24, 151, 180, 206–09,
211, 400, 609–10, 692, Arrachal: MLS 265, 2805–13, Cerro
de Suba: La Conejera: ICN 6336, 6577–79, 6580–81, 10692,
Codazzi: MLS 2386, San Joaqu´
ın: MLS 2964–65, Santana:
IAvH 4964, Cajica
´: IAvH 500, Chia: ICN 7101, MLS 2373–
77, 2382–83, 2600, 2622–23, 2830–93, 2900–08, 2935–36,
MLS not catalogued, MUJ 18, 477, Choconta
´: MLS 174
(holotype of A. colombianus), 155 (paratype of A. colombia-
nus), 159, 2620, Cogua: MLS 163, 185, Cota: MUJ 164,
Facatativa: MUJ 264, 461–62, Fontinb ´
on: MUJ 25, Fuquen´
e:
MUJ 16, 20–21, Fusagasuga: MLS 2634, MUJ 92, Guachan-
cipa
´: ICN 8261, Guacheta
´: MLS 2263, Guaduas: MUJ 01,
Guasca: MLS 2626, MUJ 203–05, 215, La Calera: MUJ 298,
La Union: MLS 157, Macheta
´: MLS 2568–70, 2653, 2921–
22, 2927, 2931, Mosquera: ICN 1453–54, 1456, 1458–59,
Laguna Herrera: IAvH 3815, ICN 859, 1277, 1457, Nem-
oc´
on: ICN 7041, Pacho: MLS 170 (holotype of A. long-
imaculatus), 154, 2611–12, 2616, 2923–30, MUJ 550, Pasca:
ICN 485–86, MLS 2602–04, Quetame: between Quetame
and Guayabetal: ICN 4477, Sisga dam: IAvH 08, Reserva
Carpanta: MLS 26, San Antonio del Tequendama: IAvH
3038–39, MLS 150–51, 200, Sesquila
´: MLS 2571, Sibat´
e:
MLS 175–76, 295, Sopo: MLS 2624, Suesca: MUJ 214, 649,
Sumapaz: MLS 168, Sutatenza: MLS 283–84, 288, 292,
1860–63, 2493–94, Tabio: MLS 1898, Tausa: MUJ 142, Tena:
MUJ 12, 19, Une: MLS 160, 177, 2709–10, Usaqu´
en: MLS
2378–79, 2381, 2894–99, MUJ 13, Villapinz ´
on: ICN 2816,
MLS 299, Villeta: IAvH 1587; Meta: Ca˜
non La Curia: MLS
06, Lomalinda: IAvH 967; Santander: without locality: MUJ
212, Bol´
ıvar: MLS 162, Jes ´
us Maria: MLS 2246–48, Puente
Nacional: MLS 2629, Santa Rita: MLS 2630. Localities likely
incorrectly labeled: Meta: Puerto Lopez: ICN 6500, MUJ 15;
Santander: Encino: MUJ 267.
Atractus elaps: (n¼89) Brazil: without locality: ZMH-R 4421
(holotype of Rabdosoma brevifrenum), IBSP 20314; Amazonas:
Borba: MNRJ 1523. Colombia: without locality: MLS 182;
Amazonas: Parque Natural Nacional Amacayacu: IAvH 3211;
Boyaca
´: Macanal: MLS 2637; Caqueta
´: without locality: MLS
183, Florencia: MLS 185, 187, 195, 197, 1316–18, 1322–23,
1326–27, 1739, 2730, 2733–39; Cauca: Puerto Bello: ICN
8240*, Santa Rosa: El Carmen: IAvH 4410; Cundinamarca:
Guaicarano: IBSP 5314 (holotype of A. elaps tetrazonus),
Paratebueno: MLS 188, Medina: MLS 192, Sasaima: MLS
2527; Meta: Acacias: MLS 191, Cubarral: ICN 7266, Pi˜
nalito:
ICN 7099, San Juan de Arama: IAvH 929, Villaviciencio: ICN
8313, MLS 179, 189, 193, 196, 266, 1396, 2054–55; R´
ıo Ocoa,
S Villavicencio: MLS 190; Putumayo: without locality: MLS
180. Ecuador: E Ecuador, without locality: EPN 6892, EPN not
catalogued; NAPO: upper R´
ıo Napo: EPN 6856, 8686,
Archidona: QCAZ 2101; R´
ıo Huataracu: EPN 8687; Orellana:
Balsayacu: Parque Sumaco: QCAZ 6502, Fuerte: EPN 7324;
Loreto: El Tena
´: EPN 8688, Parque Nacional Yasun´
ı: EPN
Fig. 7. Geographic distribution of
Atractus marthae. Type locality ¼
red triangle and additional records
¼blue circles.
Meneses-Pelayo and Passos—New polychromatic Atractus from Colombia 257
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2536, QCAZ 3249, 3959, R´
ıo Coca: QCAZ 440; Pastaza: Mera:
EPN 1175, Montalvo: Andoas: EPN 758, Nueva Vida: Misi ´
on
Agua Santa: QCAZ 3450, Puyo: QCAZ 1277, R´
ıo Bobonaza:
EPN 8678–83, R´
ıo Tall´
ın: upper R´
ıo Bobonaza: EPN 8675–77,
Sarayacu-Pucayacu: EPN 8685; Sucumb´
ıos: Lagartococha:
EPN 8689, Lago A
´grio: EPN 5781, Shushufindi: QCAZ 3303.
Peru: Amazonas: Bagua: MHNSM 2447; Hua
´nuco: Poncio
Prado: MHNSM 2082; Loreto: Maynas: MHNSM 2513; San
Mart´
ın: San Mart´
ın: MHNSM 3133, 3337, Tarapoto: MHNSM
3278. Localities likely incorrectly labeled: Pichincha: Oci-
dente: EPN 8692; El Oro: Santa Rosa: EPN 8690–91.
Atractus emigdioi: (n¼4) Venezuela: Lara: Moran: La Palma:
Pa
´ramo El Ja
´bon: MHNLS 9299; Trujillo: Bocon´
o: Valera-
Fig. 8. Landscape views of the type locality of Atractus marthae along the Vereda Esparta, municipality of Santa Ba
´rbara, department of Santander,
Colombia. General habitat (A), microhabitats (B, C), oviposition site (D), egg orientation in situ (E).
258 Copeia 107, No. 2, 2019
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Trujillo road: ULABG 3791, Parque Nacional Guaramacal: N
versant: MHNLS 16209, Trujillo: ULABG 4473.
Atractus eriki: (n¼4) Venezuela: Tujillo: Escuche: ULABG
6710 (paratype of A. eriki), Trujillo: ULABG 6694 paratype of
A. eriki), Valera: ULABG 6693 (holotype of A. eriki), CV-ULA
one specimen not catalogued.
Atractus erythromelas: (n¼31) Venezuela: M´
erida: Jaji: CV-
ULA two specimens not catalogued, La Mucuy: CV-ULA two
specimens not catalogued, La Vega: CV-ULA one specimen
not catalogued, Libertador: MHNLS 902, CBA one specimen
not catalogued, Los Guaimaros: CV-ULA one specimen not
catalogued, Manzano Alto: CV-ULA nine specimens not
catalogued, M´
erida: BMNH 1.716–17 (paratypes of A.
erythromelas), CV-ULA four specimens not catalogued, CSJ
519, Mucuruba
´: MHNLS 276–78*, 630, 902, Santo Domingo:
CV-ULA one specimen not catalogued, Timotes: CV-ULA one
specimen not catalogued; Ta
´chira: Uribante: CV-ULA one
specimen not catalogued.
Atractus indistinctus: (n¼10) Colombia: Cesar: R´
ıo de Oro:
ICN 11487; Norte de Santander: Oca˜
na: MLS 166 (holotype
of A. indistinctus), 261–62, 264, 2695–96. Venezuela: M´
erida:
La Azulita: El Hato: CV-ULA one specimen not catalogued;
Ta
´chira: CV-ULA 6117; Zulia: Sierra de Perija
´: Cerro de Las
Tetas: MBUCV not catalogued*.
Atractus meridensis: (n¼10) Venezuela: M´
erida: Libertador:
ULABG 4090–91, 4408, 4941 (paratypes of A. meridensis),
M´
erida: ULABG 4154 (paratype of A. meridensis), Las Piedras:
R´
ıo Pueblo Llano: ULABG 4341 (holotype of A. meridensis),
Parque Chorros: ULABG 2533 (paratype of A. meridensis),
Santo Domingo: ULABG 4694–96 (paratypes of A. meridensis).
Atractus mijaresi: (n¼11) Venezuela: M´
erida: La Carvonera:
CBA 01–10, Mucuruba
´: Rangel: ULABG 4697 holotype of A.
mijaresi).
Atractus multidentatus: (n¼1) Venezuela: M´
erida: La Vega: El
Parasiso: CV-ULA 7080 (holotype of A. multidentatus).
Atractus ochrosetrus: (n¼2) Venezuela: M´
erida: Tovar: ULABG
4698 (holotype of A. ochrosetrus);Tovar-Guaraque road:
ULABG 4696 (paratype of A. ochrosetrus).
Atractus pamplonensis: (n¼94) Colombia: Norte de Santander:
Bochalema: MHNLS not catalogued, Chinacota
´: MLS 2338–
39; Chitaga
´: Chucarima: MLS 273–74, 248, 287, 300; Cutilla:
MHUA 14163–64; El Diamante: MLS 1920; Labateca: ICN
10715–18*; La Donjuana: MLS 248: Oca˜
na: MLS 277;
Pamplona: IBSP 9192 (holotype of A. pamplonensis), 9190–
91, 9040, 9021 (paratypes of A. pamplonensis), MLS 241–44,
247, 250–52, 276, 2001–02, 2364, 2369–71, 2458–60, 2688–
94, 2711–15, 2753–69, MLS not catalogued, ICN 10 speci-
mens not catalogued; Alto de la Lej´
ıa: ICN 10719–24*;
Toledo: MLS 249, 253, 2700–03.
Atractus tamaensis: (n¼6) Venezuela: Ta
´chira: Junin: Betania:
MHNLS 8307 (holotype of A. tamaensis), 8301, 8303–06*
(paratypes of A. tamaensis).
Atractus taphorni: (n¼5) Venezuela: without locality: IBSP
25785; M´
erida: without locality: CV-ULA not catalogued, El
Chorotal: La Azulita road: CV-ULA 1838, La Carbonera: CV-
ULA 6417, Libertador: ULABG 3909.
Atractus turikensis: (n¼1) Venezuela: Zulia: Sierra de Perija
´:
Mesa Turik base: MBLUZ 302 (paratype of A. turikensis).
Atractus variegatus: (n¼16) Colombia: Boyaca
´: Boavita: MLS
2484–85, La Uvita: MLS 260 (holotype of A. variegatus), 217,
259, 267, 272, 278, 281, 2266, 2268–69, 2271–73, 2697.
Atractus ventrimaculatus: (n¼15) Venezuela: M´
erida: Betania:
ULABG 2409, Jaji: CV-ULA one specimen not catalogued, La
Princesa: ULABG 6701–02, Libertador: El Valle: MHNLS 897–
901*; Manzano Alto: CV-ULA two specimens not catalogued,
M´
erida: BMNH 1946. 1.5.15 (holotype of A. ventrimaculatus),
La Mucuy: Parque Nacional Sierra Nevada: MBUCV 2016,
Pico Humbo: EBRG 4052, Tapa Azul: CV-ULA one specimen
not catalogued.
Atractus vertebrolineatus: (n¼1) Colombia: Norte de
Santander: Oca˜
na: MLS 184 (holotype of A. vertebrolineatus).
Atractus wagleri: (n¼7) Colombia: Santander: Floridablanca:
UIS-R 71, Los Santos: Mesa de los Santos, El Roble Farm,
Vereda El Carrizal: UIS-R 1767, Piedecuesta: Conjunto
Residencial Pinares de Granada: UIS-R 1716, Guatigura
´:
Vereda Viricute: UIS-R 281, Mesa de los Santos, El Roble
Farm, Vereda El Carrizal: UIS-R 1488–89, San Vicente de
Chucuri: MHUA 14504*, Surata
´: near Nueva San Luis school:
Vereda San Luis: UIS-R 1725.
Atractus werneri: (n¼39) Colombia: without locality: MLS 144,
289, 483; Cundinamarca: El Col´
egio: IAvH 4327, Fusagasuga
´:
ICN 2727, MLS 2329, 2334, 2345–44, 2427, 2514, 2518, 2523,
2563, 2914–16, 2932–34, MUJ 92, La Mesa: MLS 161, La Vega:
IAvH 2068, San Francisco: ICN 5738, 10696*, Santandercito:
IAvH 3014, ICN 11207, MLS 1915–16, 2118, 2020, Sasaima:
ICN 2612, MLS 236, 238, Silvania: IAvH 145, 823–24, ICN
7268, Vereda Santa Rita: IAvH 17.
ACKNOWLEDGMENTS
We are grateful to the following curators and staff for
permission and facilities to examine specimens under their
care: J. Lynch and M. Calder ´
on (ICN), M. Ramirez-Pinilla
(UIS), R. Casallas and A. Rodrigues (MLS), A. Acosta (MUJ), D.
Per´
ıco (IAvH), V. Paez and J. Daza (MHUA), F. Bisbal (EBRG),
C. Ferreira (MBUCV), C. Se˜
naris, G. Rivas, and F. Rojas
(MHNLS), E. La Marca (ULABG), A. Pascual (VC-ULA), F.
Franco and V. Germano (IBSP), and C. McCarthy (BMNH).
Financial support for PP was provided by Conselho Nacional
de Pesquisa e Desenvolvimento Tecnol´
ogico e Cient´
ıfico
(#439375/2016-9, #306227/2015-0, and #309560/2018-7)
and Funda¸ca
˜o Carlos Chagas Filho de Apoio a
`Pesquisa do
Estado do Rio de Janeiro (#E-26/202.737/2018). EMP thanks
Don Prudencio, Do˜
na Arelis, and Don Robinson, inhabitants
of the Vereda Esparta, who kindly granted access to their
farms and received us cordially. EMP especially thanks V.
Serrano and M. Ram´
ırez-Pinilla (UIS) because they provided
substantial logistical support throughout the study.
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