Taxonomic revision of Drymoluber Amaral, 1930 (Serpentes: Colubridae)

Zootaxa 3716 (3): 349–394 www.mapress.com /zootaxa / Copyright © 2013 Magnolia Press

Article

ISSN 1175-5326 (print edition)

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http://dx.doi.org/10.11646/zootaxa.3716.3.3 http://zoobank.org/urn:lsid:zoobank.org:pub:71B98313-E0FC-427D-A06F-6917B64A64F8

Taxonomic revision of Drymoluber Amaral, 1930 (Serpentes: Colubridae) HENRIQUE CALDEIRA COSTA1,4, MÁRIO RIBEIRO MOURA2, 3 & RENATO NEVES FEIO1 1

Universidade Federal de Viçosa, Departamento de Biologia Animal, Museu de Zoologia João Moojen. CEP 36570-000. Viçosa, MG, Brazil. 2 Universidade Federal de Minas Gerais, Instituto de Ciências Biológicas, Departamento de Zoologia, Laboratório de Herpetologia. Avenida Antônio Carlos, 6627, Pampulha. CEP 31270-901. Belo Horizonte, MG, Brazil. 3 Ecos Biota Consultoria Ambiental, Rua Dr. Milton Bandeira 95/401. CEP 36570-000. Viçosa, MG, Brazil. 4 Corresponding author. E-mail: [email protected]

Abstract The present study is a taxonomic revision of the genus Drymoluber Amaral, 1930, using meristic and morphometric characters, aspects of external hemipenial morphology and body coloration. Sexual dimorphism occurs in D. dichrous and D. brazili but was not detected in D. apurimacensis. Morphological variation within D. dichrous is related to geographic distance between populations. Furthermore, variation in the number of ventrals and subcaudals in D. dichrous and D. brazili follows latitudinal and longitudinal clinal patterns. Drymoluber dichrous is diagnosed by the presence of 15-15-15 smooth dorsal scale rows with two apical pits, and 157–180 ventrals and 86–110 subcaudals; it occurs along the eastern versant of the Andes, in the Amazon forest, on the Guiana Shield, in the Atlantic forest, and its transitional areas with the Caatinga and Cerrado. Drymoluber brazili has 17-17-15 smooth dorsal scale rows with two apical pits, 182–202 ventrals and 109– 127 subcaudals, and ranges throughout the Caatinga, Cerrado, Atlantic forest and transitional areas between these last two domains. Drymoluber apurimacensis has 13-13-13 smooth dorsal scale rows without apical pits, 158–182 ventrals and 84–93 subcaudals, and occurs in the Apurímac Valley, south of the Apurímac and Pampas rivers in Peru. Key words: Snakes, South America, taxonomy, morphological variation, clinal variation

Resumo O presente estudo é uma revisão taxonômica do gênero Drymoluber Amaral, 1930, usando caracteres merísticos e morfométricos, aspectos da morfologia externa do hemipênis e de coloração. Dimorfismo sexual ocorre em D. dichrous e D. brazili, mas não foi detectado em D. apurimacensis. A variação morfológica dentro de D. dichrous tem relação com a distância geográfica entre as populações. Ademais, a variação no número de escamas ventrais e subcaudais em D. dichrous e D. brazili seguiu um padrão clinal com relação à latitude e longitude. Drymoluber dichrous se caracteriza pela presença de 15-15-15 fileiras de escamas dorsais lisas, com duas fossetas apicais, 157–180 ventrais e 86–110 subcaudais; ocorre na região oriental da Cordilheira dos Andes, Amazônia, Escudo das Guianas, Mata Atlântica e áreas de transição desta com a Caatinga e o Cerrado. Drymoluber brazili possui 17-17-15 fileiras de escamas dorsais lisas, com duas fossetas apicais, 182–202 ventrais e 109–127 subcaudais, distribuindo-se pela Caatinga, Cerrado, Mata Atlântica, e áreas de transição entre esses dois últimos domínios. Drymoluber apurimacensis possui 13-13-13 fileiras de escamas dorsais lisas, sem fossetas apicais, 158–182 ventrais e 84–93 subcaudais, e ocorre no Vale Apurímac ao sul dos rios Apurímac e Pampas, no Peru. Palavras-chave: Serpentes, América do Sul, taxonomia, variação morfológica, variação clinal

Introduction The genus Drymoluber Amaral, 1930 comprises three species of aglyphous, medium-sized, diurnal and terrestrial Colubrinae snakes (Gomes 1918; Amaral 1930; Martins & Oliveira 1998; Argôlo 2004a; Lehr et al. 2004) that are distributed in tropical South America east of the Andes (Peters & Orejas-Miranda 1970; Lehr et al. 2004; Cacciali et al. 2005; Vrcibradic 2007). Accepted by H. Zaher: 2 Jul. 2013; published: 24 Sept. 2013

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The taxonomic history of snakes currently allocated to Drymoluber began with the description of Herpetodryas dichroa by Peters (1863) who based the description on three specimens, two of them collected in Brazil by Georg Wilhelm Freyreiss, and the other “supposedly bought in Surinam.” Five years later, Cope (1868) and Günther (1868) described Spilotes piceus and Herpetodryas occipitalis respectively, based on specimens from the Amazonia region of Ecuador and Peru. Boulenger (1894) considered S. piceus and H. occipitalis junior synonyms of H. dichroa, which he transferred to the genus Coluber. Besides specimens from Peru (including the holotype of Herpetodryas occipitalis), Boulenger had access to one specimen from Bahia, in eastern Brazil, collected by Otto Wucherer. Griffin (1916) cited a specimen of “Elaphe dichroa” collected by J. D. Haseman. However, Amaral (1926), who had access to that specimen, concluded the snake was actually a young of Drymobius bifossatus Raddi, 1820 (currently in the genus Mastigodryas), an identification we confirmed after examining pictures sent to the senior author. Gomes (1918) mentioned a specimen of “Elaphe dichrous” from nothern Brazil, but did not cite Griffin (1916) in his list of synonyms nor explain the reasons that led him to transfer Coluber dichrous to the genus Elaphe. Gomes (1918) also described Drymobius brazili, based on six specimens, five of them from central Brazil and the other without locality data. Amaral (1923) described Drymobius rubriceps based on one young specimen from Penápolis, state of São Paulo, and close to the area where D. brazili was known to occur at that time. Later, Amaral (1929), emphasizing that the type-specimen of D. rubriceps might be anomalous with relation to labial and cephalic plates and coloration, transferred it to the synonymy of Drymobius boddaerti (Sentzen, 1796). Amaral (1930) stated that Coluber dichrous (the name Elaphe dichrous, presented by Gomes (1918) is not even mentioned) was not the most appropriate name for the specimens described by Peters (1863), and that they deserved to be allocated to a new genus. Based on dentary and hemipenial characters, Amaral (1930) considered it to be a taxon close to and intermediate between Coluber and Drymobius and erected the new generic name Drymoluber. When revising the taxonomic status of Drymobius, Stuart (1932) stated that Drymobius brazili should be allocated in Drymoluber, and that Drymobius rubriceps is a synonym of Drymoluber brazili, not of Eudryas (=Drymobius) boddaerti, as suggested by Amaral (1929) (Stuart 1933). For subsequent decades (Peters & Orejas-Miranda 1970), the taxonomy of Drymoluber remained unchanged, with the genus containing two species: Drymoluber dichrous (Peters, 1863) and Drymoluber brazili (Gomes, 1918). Recently, Lehr et al. (2004) described a new species, Drymoluber apurimacensis, based on six specimens from the region of the Apurímac Valley in the Peruvian Andes. In recent years, there has been a significant increase in the knowledge of the geographic distribution of D. dichrous and D. brazili from new specimens collected in the Atlantic Forest (e.g. Borges-Nojosa & Lima 2001; Passos & Brandão 2002; Santana et al. 2008), Amazonian savannas (França et al. 2006), Caatinga (Vanzolini 1981; Rodrigues 2003; Argôlo 2004b), Cerrado (Pavan & Dixo 2004; França & Araújo 2006) and transitional areas between the Atlantic Forest and the Cerrado (Cacciali et al. 2005). Although the work of Lehr et al. (2004) was published less than a decade ago, those authors did not examine specimens from the entire geographic range of Drymoluber (e.g. no specimens from the Atlantic Forest). Thus, an analysis of specimens of this genus with the broadest possible distribution is needed to evaluate the morphological variation throughout their range and to evaluate the possibility of unidentified new species. These were, therefore, the goals of this study.

Material and methods Specimens examined We examined 370 specimens of Drymoluber (286 D. dichrous, 83 D. brazili and one D. apurimacensis) including the type-series of Drymobius brazili, the holotype of Drymobius rubriceps and one paratype of Drymoluber apurimacensis (Appendices I and II). Photographs of 10 additional specimens were examined: the three syntypes of Herpetodryas dichroa, the holotypes of Herpetodryas occipitalis and Spilotes piceus, the holotype and three paratypes of Drymoluber apurimacensis and the only known specimen of D. brazili from outside of Brazil (Appendices I and II).

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Techniques used and characters analyzed The diagnostic characters used involve morphometry, maxillary dentition, coloration, pholidosis and hemipenial morphology. Sex was determined by the presence or absence of hemipenes. Terminology follows Peters (1964) and Vanzolini et al. (1980) for pholidosis, Lehr et al. (2004) for morphometric characters and Dowling & Savage (1960) for hemipenial characters. Hemipenes selected for morphological comparisons (n=24) were prepared following the methods proposed by Pesantes (1994) and Zaher & Prudente (2003). Hemipenes were manually everted by the senior author, with the exception of AMNH 54930, FMNH 40206 and FMNH 11259, which had been previously prepared. Morphometric measures were taken to the nearest 0.1 mm with a caliper, except for the snout-vent and tail lengths, which were measured with a flexible ruler to the nearest 1.0 mm. For specimens that were stiff due to fixation, a string was extended along the snake dorsum and later measured with a ruler. Maxillae were examined in situ, by making an incision between supralabial scales and the maxillary arch and lifting the soft tissue around the teeth. Later, the teeth and the empty sockets were counted. Because this procedure was invasive to preserved specimens, we performed it only to the left maxilla of each individual. Morphometric characters analyzed: 1) snout-vent length (SVL); 2) tail length (TL); 3) head length (HL) (measured from the point of the snout to the end of the retroarticular process); 4) head width between the posterior lateral margins of the supraocular plates (HWS); 5) internasal distance (head width between external nares) (ID); 6) eye diameter (ED) (measured horizontally); 7) eye-nostril distance (END) (from the anterior edge of the eye). Meristic characters analyzed: 1) number of anterior dorsal scale rows, counted one head length from the retroarticular process (AD); 2) number of midbody dorsal scale rows (MD); 3) number of posterior dorsal scale rows, counted one head length anterior to the cloacal shield (PD); 4) number of apical pits on the dorsal scales (AP); 5) number of ventrals (VE); 6) number of pre-ventrals (PV); 7) number of subcaudals (SC); 8) cloacal shield (CP) entire or divided; 9) rostral scale (RS) dorsally visible or not; 10) number of supralabials (SL); 11) number of supralabials contacting the orbit (SLO); 12) number of preoculars (PeO); 13) number of postoculars (PoO); 14) pattern of organization of the anterior temporals (AT); 15) pattern of organization of the posterior temporals (PT), see below; 16) nasal scale (NS) entire, semidivided or divided; 17) number of infralabials (IL); 18) pairs of infralabials contacting each other (ILC); 19) number of chinshields and their relative size (first chinshield longer, shorter or equal to the second chinshield) (CS); 20) infralabials contacting the first pair of chinshields (ILC1); 21) infralabials contacting the second pair of chinshields (ILC2); 22) infralabials contacting the gulars (ILG); 23) loreal (LO) as high as long, higher than long or longer than high; and 24) number of teeth in the left maxilla (T). Because of considerable variation in the arrangement of temporal plates in Drymoluber, we used a system to record the temporal formula that was different from that usually used (e.g. Peters 1964): W/X+Y/Z where W is the number of upper anterior temporals, X is the number of lower anterior temporals, Y is the number of upper posterior temporals and Z is the number of lower posterior temporals). Coloration characters analyzed: 1) dorsum of head; 2) gular region; 3) supralabials; 4) dorsum of the body; 5) venter; 6) dorsum of the tail; 7) subcaudals. For small specimens with coloration pattern different from that of the large specimens, the following measures were also taken: 1) number of dark dorsal crossbands along the body; 2) widths (measured by the number of scales in the vertebral and paravertebral rows) of the first (B1), fifth (B5), fifteenth (B15), fifth anterior to the cloacal shield (B5L) and the last (BL) dark dorsal crossband of the body; 3) widths of the light interspace after B1 and anterior to B5, B15, B5L and BL; 4) cephalic stripe immaculate, marked with ocelli/spots, or interrupted by large dark markings. Hemipenial characters analyzed: 1) lobe size in relation to the hemipenial body; 2) type of ornamentation of the lobe; 3) average number of papillae in lobe calyces; 4) ornamentation of the hemipenial body; 5) approximate number of spines on the hemipenial body; 6) ornamentation of the sulcate side of the hemipenis; 7) number of spines between the suculs spermaticus and the left basal hook; 8) ornamentation (papillae or spinules) of the walls of the sulcus spermaticus; 9) average number of spines bordering the walls of the sulcus spermaticus on each side; 10) presence of a lateral spine similar to or longer than the hooks; 11) side (right or left) of the most proximal hook; 12) ornamentation of the asulcate side of the hemipenis. Geographic coordinates The search for the geographical coordinates of localities was done using the following methods: 1) search in the catalogues of collections visited; 2) search for published works conducted in areas where the toponym is

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located; 3) search in the online Gazetteer Glosk (http://www.glosk.com), the software Google Earth® and the gazetteers by Paynter (1982; 1992; 1993; 1997) and Paynter & Taylor (1991); and 4) consultation with other researchers. Geographic coordinates are expressed in decimal degrees, the latitude followed by the longitude (e.g. 10.15, -59.45). Morphological groups With the aim of analyzing interpopulational variation in Drymoluber, the sample was divided in morphological groups (MG), following criteria of differences in the number of midbody dorsal scale rows (13, 15 or 17) and distribution assigned according to the region of occurrence. The defined groups were: MG1—13 midbody scale rows from the Andean region (Drymoluber apurimacensis; n=5); MG2—15 midbody scale rows from the Andean region (Drymoluber dichrous; n=19); MG3—15 midbody scale rows from the Amazonia north of the Amazon River (D. dichrous; n=61); MG4—15 midbody scale rows from the Amazonia south of the Amazon River (D. dichrous; n=136); MG5—15 midbody scale rows from the Atlantic Forest north of the Doce River (D. dichrous; n=54); MG6—15 midbody scale rows from the Atlantic Forest south of the Doce River (D. dichrous; n=9); MG7— 15 midbody scale rows from the Cerrado, Amazonian savannas and transitional areas between them and forest regions (D. dichrous; n=12); MG8—17 midbody scale rows from the Atlantic Forest south of the Doce River (D. brazili; n=5); MG9—17 midbody scale rows from the Cerrado and transitional areas with the Atlantic Forest (D. brazili; n=72); MG10—17 midbody scale rows from the Caatinga of northeastern Brazil (D. brazili; n=6). Specimens of Drymoluber dichrous of morphological group 2 (Andean region) were considered separately to evaluate whether they could have differentiated from the rest of the Amazonian sample because of their occurrence at high elevation sites (higher than 1000 meters above the sea level). Samples from north and south of the Amazon River (MG3 and MG4) were included in different morphological groups to evaluate whether this river could act as a natural barrier and contribute to diversification in Drymoluber. The division of specimens from the Atlantic Forest north and south of the Doce River (MG5 and MG6) took into account suggestions by climatic models that this river was at the border of forest refuges during the Pleistocene (Carnaval & Moritz 2008). Statistical analyses Analyses involving pholidosis data addressed both juvenile and adult specimens. In those analyses involving morphological data, only specimens considered adults were used. The definition of juveniles and adults was arbitrary, based on the dorsal coloration pattern, since there are no published works about the reproductive biology of Drymoluber that could have been helpful with this issue. Specimens presenting uniform dorsal coloration were considered adults, while those with coloration formed by dark and light crossbands were considered juveniles. However, seven specimens of D. dichrous (IB 46626, MNRJ 17069, MPEG 16921, MPEG 17799, MPEG 17820, MZUESC 3739, MZUSP 7681) with uniform dorsal coloration were considered juveniles because of their small size (SVL=285-391 mm). On the other hand, five females (AMNH 54541, AMNH 55635, MZUSP 11444, USNM 204126, USNM 332470; SVL=445–590 mm) and four males (AMNH 22491, AMNH 91812, IBSP 69567, MPEG 20331; SVL=425–485 mm) of D. dichrous and one male of D. brazili (IBSP 18309; SVL=544 mm) having some dorsal crossbands (sometimes faded) were considered adults, because of the presence of several other specimens of similar size but with dorsal coloration totally uniform. First, the normality and homocedasticity of the sampled variables were verified through the KolmogorovSmirnov and Levene tests. The variables that did not meet those assumptions were later analyzed with the nonparametrical tests of Mann-Whitney U and Krush-Wallis (Zar 1999). The presence of sexual dimorphism in the three species was evaluated separately through the univariate analysis of variance (ANOVA) for number of ventral plates and number of subcaudal plates. Multivariate analyses of variance (MANOVA) were also used based on the seven morphometric measures (SVL, TL, HL, HWS, ID, ED and END). All statistical analyses were done with the Statistica 7.0 software (StatSoft 2004). Principal component analyses (PCA) were done separately for males and females, evaluating the distribution of specimens in the multivariate space, aiming to highlight differences among groups not defined a priori (Johnson & Wichern 1998; Manly 2004). Some characters were not included in the PCA because they did no show sufficient variation. Thus, the following pholidosis characters were used: MD, VE, SC, SL, SLO, TA, TP, IL, ILC1, ILC2, ILG and LO. For SL, SLO, AT, PT, IL, ILC1, ILC2 and ILG, both right and left sides were analyzed, since the presence of specimens with different scale counts at each side of the head was not uncommon. The classification of

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variations in AT and PT (see results) follows an ordinal scale from 1 to 26 and 1 to 16, respectively. For the PCAs based on morphometry, measures (SVL, TL, HL, HWS, ID, ED and END) were logarithmized, due to the presence of very different scales of magnitude, for example from 2 mm to more than 1000 mm. The MG6 was not included in the PCAs for females because of the absence of specimens. Measures of missing characters were estimated for some specimens using the mean substitution function of Statistica 7.0. Specimens with absence of information for more than 30% of the analyzed characters were excluded from the analyses. For both PCAs (based on pholidosis and morphometry), the covariance matrix was calculated, from which the eigenvalues and eigenvectors that define the components were extracted. The variables that most contributed to the formation of the principal components were determined in descending order by the correlation between the variables and the factor coordinates in each of the three first components (James & McCulloch 1990). The contribution of each variable (factor coordinates) in the formation of principal components was projected in the reduced space of the most contributing principal components, as well as the individual scores of the analyzed specimens (Cavalcanti & Lopes 2003). Only descriptive observations were used to compare the variation in coloration pattern of young specimens, because half of the morphological groups (MG1, MG6, MG7, MG8, MG10) had three or fewer specimens, precluding statistical analyses.

FIGURE 1. Distribution of morphological groups of males (A) and females (B) of Drymoluber dichrous used to verify the relationship between geographical and morphological distances using the Mantel test. Each point has 60 Km of radius. Males: 1) Iquitos + Nauta + Rio Ucayali (Loreto, Peru); 2) Porto Velho + Alto Paraíso (Rondônia, Brazil); 3) UHE Balbina (Presidente Figueiredo, Amazonas, Brazil); 4) Carajás (Pará, Brazil); 5) Belém + Benevides + Castanhal + São Domingos do Capim (Pará, Brazil); 6) BR 316 (Maranhão, Brazil); 7) Aracruz + Linhares + Vitória (Espírito Santo, Brazil); 8) Arataca + Barro Preto + Buerarema + Camacan + Ilhéus + Una (Bahia, Brazil); 9) Barra do Rocha + Cairu + Itacaré + Ituberá + Nova Ibiá (Bahia, Brazil); 10) Maranguape + Pacoti + Pacatuba (Ceará, Brazil). Females: 1) Cabeceras del Rio Arajuno + Cabeceras del Rio Bobonaza + Macas + Riobamba + Río Liguino (Ecuador); 2) Iquitos (Loreto, Peru); 3) Porto Walter + Reserva Extrativista Riozinho da Liberdade (Tarauacá, Acre, Brazil). 4) Porto Velho (Rondônia, Brazil); 5) FLONA Caxiuanã (Melgaço, Pará, Brazil); 6) Carajás (Pará, Brazil); 7) Buerarema + Ilhéus + Itabuna + Una (Bahia, Brazil); 8) Camacan + Ibirataia + Ituberá + Nova Ibiá (Bahia, Brazil). Tropical and subtropical moist forests; Tropical and subtropical savannas; Deserts and xeric formations; Mangroves; Floodplains; Tropical and subtropical dry forests; Mediterranean forest of woods and shrubs; Mountain grasslands. Habitat types follow Olson et al. (2001).

To infer the relationship between geographical distribution and the variation in pholidosis characters, we selected localities with at least five cospecific specimens of the same sex, designating those localities as geographical groups. To increase the number of groups, areas with an arbitrary radius of 60 km were used instead of point localities. The analysis could be done only with Drymoluber dichrous, since the sample of D. brazili did not permit the formation of at least three groups of the same sex. Even if the sample of D. apurimacensis were sufficient, the analysis would not be necessary, since this species has a restricted distribution. Thus, ten

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geographical groups for males and eight for females of D. dichrous were used (Fig. 1), from which we calculated the dissimilarity matrix based on the distance D² of Mahalanobis and the matrix of geographical distance (in km) for males and females separately. The correlation between matrices was analyzed with the Mantel test (Urban 2003), to verify the relationship between geographical and morphological distances (de Queiroz & Good 1997; Passos et al. 2005). The Pearson coefficient of correlation (Zar 1999) was used to evaluate whether variations in the number of ventral and subcaudal plates in D. brazili and D. dichrous follow latitudinal and/or longitudinal clinal patterns.

Results and Discussion Sexual dimorphism Drymoluber dichrous—Females have more ventrals than males (F(1,285) = 252.11; p < 0.001), and the variation in the number of subcaudals was not significant. Males had higher values with respect to morphometric variables (Λ = 0.841; F(5,135) = 5.071; p < 0.001), with greater snout-vent length (F(1,139) = 16.235; p < 0.001), greater tail length (F(1,139) = 11.078; p < 0.001), greater head length (F(1,139) = 21.180; p < 0.001), greater head width in the supraocular region (U = 2729; p < 0.001), greater internasal distance (F(1,139) = 16.573; p < 0.001), greater eye diameter (U = 3165; p < 0.001) and greater eye-nostril distance (F(1,139) = 18.453; p < 0.001). Drymoluber brazili—Females of Drymoluber brazili also had a higher number of ventrals than males (F(1,81) = 27.081; p < 0.001), while the variation in the number of subcaudals was not significant. Morphometric data did not show sexual dimorphism. Drymoluber apurimacensis—Sexual dimorphism was not found in ventral and subcaudal counts, however this may be a reflection of the small sample size (n=5). Morphometrical measures could not be compared because only two of the five examined specimens were females, one of them a juvenile and the other without head. Principal Component Analyses With regard to the pholidosis characters, there was an overlap between the morphological groups 1 to 7 (Drymoluber apurimacensis and D. dicrhous) and 8 to 10 (D. brazili) for males and females (Figs. 2 and 3; Tables 1 and 2). The different contribuitions of the variables to the formation of the principal components indicate that they do not present similar importance during the process of component formation (Reis et al. 1988). The discrimination obtained along the axis of the first principal component is mainly correlated with the variables VE (r = 0.967; p < 0.001) and SC (r = 0.864, p < 0.001), the contributions of which differed in magnitude with relation to all other variables in males (Table 1), and all variables in females with the exception ATl and ATr (Table 2). The variables VE and SC were also important to the formation of the second and third components, although do not always acting in the same direction of variation. In the projections related to the morphometry of specimens (Figs. 4 and 5; Tables 3 and 4), we observed a great overlap between the morphological groups of males of Drymoluber (Figs. 4A and 4B). This overlap is reduced when females are analyzed (Figs. 4C and 4D). Although there is no clear morphometric differentiation between the species of Drymoluber, there is a tendency for females of D. brazili (MG8 to 10) to attain greater proportions than females of D. apurimacencis and D. dichrous (MG1 to 7). The first component can be considered as a size indicator, since it presented the same magnitude and variation signal for all coefficients (Tables 3 and 4). The other components can be considered as shape indicators, since their coefficients alternate between positive and negative values (Humphries et al. 1981). Geographical distance in Drymoluber dichrous There were positive correlations between the Mahalanobis D2 and the geographical distances for males (r = 0.337; p = 0.023) and females (r = 0.513; p = 0.005) of Drymoluber dichrous (Fig. 6). This result indicates the presence of an isolation-by-distance model, where relatively great genetic differences can develop between geographically distant populations within the same genetically continuous unit (de Queiroz & Good 1997), suggesting a pattern of clinal variation in Drymoluber dichrous. A similar situation was recently found for Dipsas albifrons in the coastal Atlantic Forest of Brazil (Passos et al. 2005).

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Clinal patterns in ventral and subcaudal plates The variation in the number of ventrals and subcaudals in Drymoluber brazili and D. dichrous present a clinal pattern in relation to the latitude and/or longitude. In D. dichrous, there is a decrease from south to north in the number of ventrals in males (r = -0.226, p = 0.003) and females (r = -0.195, p = 0.037), and in the number of subcaudals in females (r = -0.242, p = 0.022). With regard to longitude, significant clinal variation occurs only in the number of subcaudals, increasing from west to east in both sexes (males: r = 0.410, p < 0.001; females: r = 0.434, p < 0.001) (Figs. 7 and 8). TABLE 1. Standardized coefficients and correlation (r) between coefficients and variables (factor loadings) resulted from the principal component analysis using 20 pholidosis characters of male specimens of Drymoluber. PC1

PC2

PC3

(r) PC1

(r) PC2

VE

0.812

0.188

-0.511

0.967 ***

0.102

SC

0.557

-0.422

0.702

0.864 ***

-0.298 ***

ILGl ILGr

-0.084 -0.079

MD

0.063

ATl PTr

-0.023 -0.022

0.039 0.043

-0.721 *** -0.737 *** 0.898 ***

N.S

(r) PC3 -0.225 *** 0.403275

-0.090

N.S

0.125 *

-0.094

N.S

0.151 *

N.S

-0.055 N.S

0.015

-0.010

0.061

0.595

0.285

0.176 **

0.777 ***

0.302 ***

0.055

0.126

-0.072

0.304 ***

0.315 ***

-0.147 *

ATr

0.049

0.633

0.384

0.137 *

0.796 ***

0.393 ***

PTl

0.023

0.109

-0.075

0.137 *

0.287 ***

-0.159 *

0.100

SLOl

0.023

0.014

0.007

0.399 ***

0.116

SLOr

0.022

0.005

-0.005

0.378 ***

0.039 N.S

-0.033 N.S

ILC1r

0.016

-0.005

-0.004

0.396 ***

-0.058 N.S

-0.037 N.S

ILC1l

0.015

0.005

-0.007

0.350 ***

0.056 N.S

-0.063 N.S

LO

-0.007

-0.001

-0.008

-0.223 ***

-0.016 *

-0.090 N.S

ILC2r

0.005

-0.019

0.006

0.065 N.S

-0.115 N.S

0.032 N.S

ILr

0.004

0.005

0.006

0.149 *

0.084 N.S

0.078 N.S

ILl

0.003

0.008

0.0009

0.119 N.S

0.116 N.S

0.010 N.S

SLl

0.0009

0.005

0.006

0.048 N.S

0.125 N.S

0.118 N.S

ILC2l

-0.0006

0.002

-0.006

-0.007 N.S

0.013 N.S

-0.028 N.S

SLr

0.0005

0.0004

-0.003

0.025 N.S

0.010 N.S

-0.062 N.S

Eigenvalues

144.349

174.292

194.094

–

–

–

Cum. prop. (%)

66.899

80.776

89.953

–

–

–

N.S

0.049 N.S

NS = not significant; * = p < 0.05; ** = p < 0.01; *** = p < 0.001; Cum. prop. = cumulative proportion of eigenvalues in percentage (%) for the first three principal components. For variable symbols see Material and methods (lower case ‘l’ and ‘r’ means left and right side of head, respectively). TABLE 2. Standardized coefficients and correlation (r) between coefficients and variables (factor loadings) resulted from the principal component analysis using 20 pholidosis characters of female specimens of Drymoluber. PC1

PC2

PC3

(r) PC1

(r) PC2

(r) PC3

VE

0.746

-0.468

0.439

0.950 ***

-0.242 **

0.190 *

SC

0.626

0.720

-0.288

0.896 ***

0.419 ***

-0.140 N.S

ATr

0.132

-0.378

-0.724

0.392 ***

-0.454 ***

-0.728 ***

ATl

0.100

-0.270

-0.420

0.367 ***

-0.402 ***

-0.523 ***

ILGl

-0.083

0.060

0.009

-0.757 ***

0.223 **

0.029 N.S

ILGr

-0.072

0.070

0.013

-0.688 ***

0.272 ***

0.044 N.S

MD

0.065

-0.024

-0.001

0.904 ***

-0.137 N.S

-0.007 N.S

......continued on the next page

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TABLE 2. (Continued) PC1

PC2

PC3

(r) PC1

(r) PC2

(r) PC3

PTr

0.043

-0.124

-0.077

0.267 ***

-0.315 ***

-0.163 *

PTl

0.038

-0.135

-0.120

0.256 **

-0.366 ***

-0.272 ***

SLOl

0.032

-0.036

0.029

0.426 ***

-0.193 *

0.132 N.S

SLOr

0.031

-0.015

0.010

0.453 ***

-0.092 N.S

0.051 N.S

LO

-0.019

0.006

0.001

-0.451 ***

0.060

ILC1r

0.012

-0.005

0.0002

0.317 ***

-0.053 N.S

0.001 N.S

N.S

0.015 N.S

N.S

0.013 N.S

ILC1l

0.010

-0.001

0.001

0.252 **

-0.013

ILC2l

-0.009

0.013

0.003

-0.118 N.S

0.064 N.S

0.013 N.S

ILr

0.009

0.012

0.009

0.265 ***

0.142 N.S

0.087 N.S

N.S

0.154 N.S

ILl

0.005

0.007

0.013

0.197 *

ILC2r

-0.004

0.011

-0.003

-0.057 N.S

0.057 N.S

-0.013 N.S

SLr

0.001

0.0007

0.001

0.077 N.S

0.013 N.S

0.021 N.S

N.S

0.107

SLl

0.0004

-0.004

0.006

Eigenvalues

155.089

180.710

198.614

–

–

–

Cum. prop. (%)

71.353

83.141

91.378

–

–

–

0.017

-0.064

N.S

0.081 N.S

NS = not significant; * = p < 0.05; ** = p < 0.01; *** = p < 0.001; Cum. prop. = cumulative proportion of eigenvalues in percentage (%) for the first three principal components. For variable symbols see Material and methods (lower case ‘l’ and ‘r’ means left and right side of head, respectively).

FIGURE 2. Projections of the individual scores resulting from the principal component analysis (PCA) and confidence ellipses (p > 0.95) in the reduced space of the three first axes for males (A, B) and females (C, D) of Drymoluber, using 20 pholidosis characters. Standardized coefficients and factor loadings (r) are presented in Tables 1 and 2, respectively. MG1 = D. apurimacensis; MG2–7 = D. dichrous; MG8–10 = D. brazili.

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FIGURE 3. Projections of the factor coordinates resulting from the principal component analysis (PCA) for the 20 pholidosis characters of Drymoluber in the reduced space of the three first axes for male (A, B) and female (C, D). VE = number of ventrals; SC = number of subcaudals; ATr e ATl = pattern of anterior temporals on right and left sides of the head, respectively; PTr and PTl = pattern of posterior temporals on right and left sides of the head, respectively.

TABLE 3. Standardized coefficients and correlation (r) between coefficients and variables (factor loadings) resulted from the principal component analysis using seven morphometric characters of male specimens of Drymoluber PC1

PC2

PC3

(r) PC1

(r) PC2

(r) PC3

SVL

-0.508

0.277

0.199

-0.938 ***

0.198 *

0.102 N.S

HL

-0.445

-0.894

0.024

-0.788 ***

-0.614 ***

0.011 N.S

TL

-0.367

0.154

-0.910

-0.815 ***

0.133 N.S

-0.561 ***

END

-0.365

0.182

0.237

-0.912 ***

0.177 *

0.164 *

ID

-0.337

0.178

0.098

-0.864 ***

0.176 *

0.070 N.S

HWS

-0.307

0.149

0.178

-0.922 ***

0.174 *

0.148 N.S

ED

-0.258

0.105

0.183

-0.862 ***

0.136 N.S

0.170 *

Eigenvalues

0.034

0.039

0.042

–

–

–

Cum. prop. (%)

75.307

86.642

92.449

–

–

–

NS = not significant; * = p < 0.05; ** = p < 0.01; *** = p < 0.001; Cum. prop. = cumulative proportion of eigenvalues in percentage (%) for the first three principal components. For variable symbols see Fig. 5 or Materials and Methods. TAXONOMIC REVISION OF DRYMOLUBER

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FIGURE 4. Projections of the individual scores resulting from the principal component analysis (PCA) and confidence ellipses (p > 0.95) in the reduced space of the three first axes for male (A, B) and female (C, D) specimens of Drymoluber, using seven morphometric characters. Standardized coefficients and factor loadings (r) are presented in Tables 3 and 4, respectively. MG1 = D. apurimacensis; MG2–7 = D. dichrous; MG8–10 = D. brazili.

In Drymoluber brazili, clinal variation in the number of ventral and subcaudal plates was found only for males. From south to north, there is an increase in the number of subcaudals (r = 0.416, p = 0.027). From west to east, ventral plates in males increase in number (r = 0.311, p = 0.039, Figs. 9 and 10). Latitudinal and/or longitudinal clinal patterns related to pholidosis, morphometry and coloration have been described for several snake species (e.g. Hoge et al. 1977; Passos et al. 2005; Allsteadt et al. 2006; Passos & Fernandes 2008), and, in some cases, two or more taxa were synonymized when it is realized that they simply constitute distint parts of a clinal pattern previously masked by insufficient sampling (e.g. McDiarmid 1968; Gardner & Mendelson III 2004; Manier 2004). The causes of clinal variation in the scutelation of snakes, especially in the number of ventrals and subcaudals, are still uncertain, requiring further study. In most snake families, the number of ventrals corresponds to the number of vertebrae (Fox 1948; Alexander & Gans 1966), which could be affected during somite formation (Fox 1948). Laboratory tests indicated that the number of ventrals is influenced by the temperature of incubation of the clutch (Fox 1948; Osgood 1978), suggesting that environmental conditions could explain clinal variations (e.g. Hoge et al. 1977; Passos et al. 2005; Passos & Fernandes 2008). However, recent findings suggest that geographic differences in meristic counts would not be caused by direct effects of the environment during snake development (Arnold & Peterson 2002). In some cases, it can actually be the result of a past fragmentation event of the species range, instead of recent ecological effects (Grazziotin et al. 2006). Coloration of juvenile specimens The juveniles of Drymoluber present a color pattern composed of dark crossbands that cover the entire dorsum and extend to the lateral margins of the ventrals and are separated by light interspaces. This pattern is common in all species of the genus, and is present in other Neotropical Colubrinae, such as Dendrophidion dendrophis, Mastigodryas spp. and some Chironius. Usually, the crossbands tend to disappear in the posterior–anterior direction, with few specimens having crossbands throughout the whole dorsum. Different crossbands tend to have different widths, but each crossband tends to have the same width across all the rows of scales it spans. Sometimes there are extensions or compressions in the crossbands, giving them a zig-zag aspect.

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FIGURE 5. Projections of the factor coordinates resulting from the principal component analysis (PCA) for seven morphometric characters of Drymoluber specimens in the reduced space of the three first axes for male (A, B) and female (C, D). SVL = snout-vent length; TL = tail length; HL = head length; HWS = head width between the lateral margins of the supraocular plates; ID = internasal distance eye diameter; DOn = eye-naris distance.

TABLE 4. Standardized coefficients and correlation (r) between coefficients and variables (factor loadings) resulted from the principal component analysis using seven morphometric characters of female specimens of Drymoluber. PC1

PC2

PC3

(r) PC1

(r) PC2

SVL

-0.513

0.124

0.169

-0.963 ***

0.085

TL

-0.403

0.683

-0.539

-0.801 ***

0.498 ***

END

-0.392

-0.108

0.324

-0.909 ***

-0.092

ID

-0.347

-0.211

0.269

-0.855 ***

-0.191 *

0.195 *

HL

-0.342

0.028

0.279

-0.922 ***

0.028

N.S

0.221 *

HWS

-0.331

-0.676

-0.643

-0.727 ***

-0.544 ***

ED

-0.265

-0.034

0.103

-0.876 ***

-0.041

Eigenvalues

0.028

0.032

0.035

–

–

–

Cum. prop. (%)

75.3621

85.5310

92.0589

–

–

–

N.S

N.S

N.S

(r) PC3 0.093 N.S -0.315 ** 0.221 *

-0.414 *** 0.100 N.S

NS = not significant; * = p < 0.05; ** = p < 0.01; *** = p < 0.001; Cum. prop. = cumulative proportion of eigenvalues in percentage (%) for the first three principal components. For variable symbols see Fig. 5 or Material and methods.

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FIGURE 6. Projections of the Mahalanobis D² distance versus the geographical distance (Km) calculated from pholidosis characters in males (A) and females (B) of Drymoluber dichrous. Dotted lines correspond to the 95% confidence interval for the estimated relationship.

FIGURE 7. Linear correlation of the number of ventrals in function of the latitude and longitude for males (A) and females (B) of Drymoluber dichrous.

FIGURE 8. Linear correlation of the number of subcaudals in function of the latitude and longitude for males (A) and females (B) of Drymoluber dichrous.

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FIGURE 9. Linear correlation of the number of ventrals in function of the latitude and longitude for males (A) and females (B) of Drymoluber brazili.

FIGURE 10. Linear correlation of the number of subcaudals in function of the latitude and longitude for males (A) and females (B) of Drymoluber brazili.

Only Drymoluber apurimacensis is distinguishable based on the juvenile color pattern. It has dark crossbands that usually are narrower and light interspaces that are wider than those in D. dichrous and D. brazili. In D. brazili, the light interspaces are on average wider than in D. dichrous, but the existing variation does not make it a reliable character to distinguish them (see more on Taxonomy). Apparently, the most variable coloration character in juvenile specimens of Drymoluber is that of the head. A distinct and wide transverse light stripe in the parietal region (immaculate or maculate, usually with two large spots) is often present, but it also may be absent, giving way to a dark coloration covering most of the parietals, usually leaving only a light longitudinal mark in the middle of the parietal region (Fig. 11). Apparently, during ontogeny there is no expansion of the pre-existing dark coloration, but a continuous darkening of the light areas. Thus, the three variation types found here would not represent different ontogenetic stages. Considerable variation of head coloration was observed in Drymoluber dichrous from Andes and Amazonia and D. brazili from the Cerrado and Caatinga, where the light head stripe can be present or not in young specimens, even from the same locality (e.g., D. dichrous from Aripuanã (-10.15, -59.45), in southwestern Brazilian Amazonia). On the other hand, no juveniles of D. dichrous from the Atlantic Forest (n=11) has the light stripe, which is present in specimens of D. brazili from this region (n=2) and its transitional areas with the Cerrado (n=2). This, however, may be a bias resulting from a small sample. TAXONOMIC REVISION OF DRYMOLUBER

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Taxonomy The results of the present study do not indicate undescribed species within Drymoluber. Although Drymoluber apurimacensis did not show distinction from D. dichrous in the principal component analyses, it must be maintained as a valid species, since two charactes easily distinguish it from D. dichrous: the presence of 13 dorsal scale rows along the body, without apical pits (15 dorsal scale rows along the body, with two apical pits in D. dichrous). The municipality of Nova Ponte (-19.14, -47.68) in the state of Minas Gerais, Brazil, a region of ecotone between the Cerrado and the Atlantic Forest domains is the only known locality where D. dichrous and D. brazili occur in sympatry. However, there are cases of two nearby localities each with a record of one species, and maybe the absence of the other species could be due to the lack of sampling: Ouro Preto, Estação Ecológica do Tripuí (20.40, -43.58; record of D. dichrous) and Mariana (-20.38, -43.42; record of D. brazili), Minas Gerais state; Caratinga, RPPN Feliciano Miguel Abdala (-19.71, -41.81; record of D. dichrous) and Alvarenga (-19.41, -41.72; record of D. brazili), Minas Gerais state; Missão Velha, Santo Antônio water spring (-7.41, -39.21; record of D. dichrous) and Milagres (-7.31, -38.95; record of D. brazili), Ceará state.

FIGURE 11. Variation in color of the dorsum of the head in young specimens of Drymoluber. A) Presence of a light parietal stripe (MZUSP 8494, D. dichrous, Alto Paraíso, Rondônia, Brazil); B) Light stripe present, although marked with two large spots (IBSP 16499, D. brazili, Turiba do Sul, São Paulo, Brazil); C) Light stripe absent (MZUSP 14298, D. brazili, UHE Luís Eduardo Magalhães, Tocantins, Brazil). Photos: Henrique C. Costa.

Drymoluber Amaral, 1930 Drymoluber Amaral, 1930. Memórias do Instituto Butantan, 4, p. 335. Type species by monotypy: Herpetodryas dichroa Peters, 1863

Diagnosis: Drymoluber is distinguished from all other Neotropical Colubrinae by the following combination of characters: a) dorsal scales smooth, in 13, 15 or 17 midbody rows; b) cloacal shield entire (rarely divided); c) 157– 202 ventrals; d) 84–127 divided subcaudals; e) caudal pseudoautotomy; f) two pairs of chinshields, the first about half the length of the second; g) 8 (less commonly 7 or 9) supralabials; h) 8 or 9 (less commonly 7 and 10) infralabials; i) 1 (rarely 2) preocular; j) 2 (rarely 1 or 3) postoculars; k) 14–26 maxillary teeth; l) ontogenetic variation in the dorsal coloration of body and head (small specimens have dark and white/red colored macules on the head, and the body with dark crossbands separated by light interspaces, while large specimens have dorsal coloration uniformly green, brown or gray); m) hemipenes single, subcylindrical, not capitate, with the lobe about

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half the length of the organ, ornamented with papillate calyces gradually replaced by spinulate flounces and spines. The spines are arranged in more or less transverse rows, those bordering the sulcus spermaticus having a basal hook. Content: Three species: Drymoluber dichrous (Peters, 1863), Drymoluber brazili (Gomes, 1918) and Drymoluber apurimacensis Lehr, Carrillo & Hocking, 2004. Geographical distribution (Fig. 12): The genus Drymoluber is widely distributed in the South America east of the Andes, almost entirely north of the Tropic of Capricorn. It occurs in Amazonia, the Guiana Shield and along the eastern side of Andes, the Atlantic Forest from northeastern to southeastern Brazil, the Brazilian Cerrado and Caatinga, transitional areas between the Atlantic Forest and Cerrado in Brazil and Paraguay, and transitional areas between the Atlantic Forest and Caatinga. The change of the climate and ecosystems south of the Tropic of Capricorn (-23.45) from tropical to subtropical seems to be a decisive factor limiting the austral distribution of Drymoluber (Bérnils et al. 2007). The elevational distribution of Drymoluber varies from sea level at the Brazilian coast to about 3500 meters in the Andes.

FIGURE 12. Geographical distribution of Drymoluber Amaral, 1930. Black symbols represent localities with specimens examined, and white symbols represent literature records. Circles = Drymoluber dichrous (Peters, 1863); Triangles = Drymoluber brazili (Gomes, 1918); Lozenges = Drymoluber apurimacensis Lehr, Carrillo & Hocking, 2004. Tropical and subtropical moist forests; Tropical and subtropical savannas; Deserts and xeric formations; Mangroves; Floodplains; Tropical and subtropical dry forests; Mediterranean forest of woods and shrubs; Mountain grasslands. Habitat types follow Olson et al. (2001).

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Etymology: Amaral (1930) stated that the dentary and hemipenial characters of Herpetodryas dichroa Peters, 1863 suggested that was a taxon closely related and intermediate between Drymobius and Coluber. The generic name Drymoluber (an amalgam of the words Drymobius and Coluber), was proposed to simultaneously distinguish and show the close relationship between Herpetodryas dichroa (now Drymoluber dichrous) and those genera. The name Drymoluber is masculine in gender.

Drymoluber dichrous (Peters, 1863) Herpetodryas dichroa Peters, 1863. Monatsberichte der königlich Akademie der Wissenschaften zu Berlin, 29, p. 284. Syntypes: ZMB 1661, ZMB 1662, ZMB 2603. Herpetodryas occipitalis Günther, 1868. Annals and Magazine of Natural History, Fourth Series, 1, p. 420. Holotype: BMNH 1946.1.14.61, formerly 1867.9.17.28. Spilotes piceus Cope, 1868. Proceedings of the Academy of Natural Sciences of Philadelphia, 20, p. 105–106. Holotype: ANSP 3920. Coluber dichrous—Boulenger, 1894. Catalogue of the Snakes in the British Museum (Natural History). Volume II. British Museum of Natural History, London, p. 30–31. Elaphe dichrous—Gomes, 1918. Memórias do Instituto Butantan, 1, p. 67. Drymoluber dichrous—Amaral, 1930. Memórias do Instituto Butantan, 4, p. 337.

Lectotype (here designated in accordance with Article 74 of the International Code of Zoological Nomenclature): Museum für Naturkunde Berlin ZMB 1661, adult of undetermined sex (probably a male), SVL 585 mm, TL 242 mm, collected in Brazil during the first half of the 19th century by Georg Wilhelm Freyreiss. Specimen examined by photographs. Although we do not know specifically where specimen ZMB 1661 was collected, the itinerary of its collector is known, and this information leads us to designate it as the lectotype, instead of the syntype ZMB 2603, which was listed as “probably from Suriname” or ZMB 1662 which has a broken tail. Paralectotypes: Museum für Naturkunde Berlin ZMB 1662, adult of undetermined sex, SVL 568 mm, TL 237+N mm (broken tail), collected in Brazil during the first half of the 19th century by Georg Wilhelm Freyreiss; ZMB 2603, adult of undetermined sex (probably a female), SVL 623 mm, TL 225+N mm (broken tail), supposedly bought in Suriname. This same information about the collection site of ZMB 2603 is recorded in the catalogue of the Museum für Naturkunde Berlin and on the oldest label of the specimen. However, a newer label (with the name Drymoluber dichrous) indicates “Brasilien Becker”. The reason and source for the adjusted locality is unknown even to the current curator (M.O. Rödel, pers. com.). We examined the specimens from photographs. About the type locality: There is no information about the dates of collection, shipment to Europe or arrival at the Museum für Naturkunde Berlin of the type series of D. dichrous (M.O. Rödel, pers. com.). Thus, the type locality of D. dichrous was reported as “Brazil and Surinam” (e.g. Peters & Orejas-Miranda 1970). Since the lectotype designated above (ZMB 1661) is the new name-bearing type of D. dichrous, its place of collection is the type locality. Despite the lack of detailed information as to where the specimen was collected, we consider the type locality to be the area traveled by its collector, G.W. Freyreiss, in Brazil (Fig. 13 and text below). Freyreiss was born in Frankfurt on 12 July 1789. In 1813 he left St. Petersburg bound for Brazil, starting his expeditions in June 1814. He departed from the province (currently state) of Rio de Janeiro and travelled south to Minas Gerais, along the Caminho do Proença, a path of the ancient royal road to Vila Rica (currently the municipality of Ouro Preto) (Freyreiss 1907; Papavero 1971) (Fig. 13, localities 1–12). In September 1814, Freyreiss explored the region around the Abaeté, Indaiá and São Francisco Rivers, later returning to Vila Rica (Freyreiss 1907; Papavero 1971) (Fig. 13, localities 12–18–12). In December 1814 he began a second trip to study Indian tribes, travelling from Vila Rica to the vicinities of Presídio São João Batista (currently the municipality of Visconde do Rio Branco) (Fig. 13, localities 12–23). Later he returned to Vila Rica, and then (January 1815) to Rio de Janeiro, leaving no records of his path in this part of the voyage (Freyreiss 1907; Papavero 1971). In July 1815, Maximilian Alexander Philipp, prince of Wied-Neuwied (also known just as Wied) arrived in Brazil, and in August began a natural history trip together with Freyreiss and Friedrich Sellow (Wied 1989; Papavero 1971). From the city of Rio de Janeiro they headed north to Espírito Santo and Bahia provinces (Wied 1989; Papavero 1971). In February 1816, the naturalists were in the Mucuri River, southern Bahia (Fig. 13, localities 1, 24–37). There, Freyreiss decided to return to Espírito Santo, while Wied continued his travel northward (Wied 1989, p. 186). In Espírito Santo, Freyreiss visited again the localities of São Mateus and Linhares (Fig. 13, localities 35 and 36), and

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in May 1816 he travelled back to Mucuri (Fig. 13, locality 37) to visit Wied (Wied 1989, p. 162, 170, 201). While Freyreiss and Sellow stayed in Mucuri, Wied continued travelling toward the north. In Ocotber 1816, Wied returned to Mucuri to visit Freyreiss and Sellow, with whom he spent three weeks before heading north again (Wied 1989, p. 273). There is no itinerary of the voyages of Freyreiss after that, but it is known that in Bahia he also visited the localities of Caravelas, Canavieiras (Wied 1989, p. 330) and Salvador (Fig. 13, localities 39, 40 and 41), and contributed to the foundation of a German colony, Colônia Leopoldina, currently part of the municipality of Nova Viçosa (Papavero 1971) (Fig. 13, locality 38). There are some inconsistencies related to the place of death of Freyreiss, in 1825. Löfgren (1902) and Papavero (1971) wrote that the German naturalist died in Colônia Leopoldina, while Rocha (1972 “1973”) stated that he died in Europe. Wied described the species collected during his travels, and his collections are now in the American Museum of Natural History. Even though the type series of D. dichrous is in Germany, we believe that if Freyreiss collected the specimens of D. dichrous during the trip from Rio de Janeiro to Bahia, it probably was done at times when he was not with Wied’s expedition. It is known that from his trips through Espírito Santo and Bahia, Freyreiss sent three shipments of collected specimens to Europe (Papavero, 1971), but, as we have written above, there is no information about the dates when the type series of D. dichrous was collected, shipped to, or arrived in Europe. Additionally, we have record of a single shipment of specimens sent to Europe from his trip through Minas Gerais, when he stated that his collections were sent to Rio de Janeiro on 30 July 1814 (Freyreiss 1907, p. 167). As his travels continued, he certainly made other shipments, of which we have no information. Diagnosis: Drymoluber dichrous is distinguished from D. brazili and D. apurimacensis by the following combination of characters: a) 15-15-15 dorsal scale rows with two apical pits; b) 157–173 ventrals in males, 160– 180 in females; c) 87–110 subcaudals in males, 86–109 in females; d) 19–26 maxillary teeth. See Table 5. Comparisons: Drymoluber brazili has 17-17-15 dorsal scales rows, and D. apurimacensis has 13-13-13. Apical pits are absent in D. apurimacensis. Drymoluber brazili has 182–200 ventrals in males and 185–202 in females, 109–127 subcaudals in males and 109–126 in females. Drymoluber apurimacensis is not distinguishable from D. dichrous based on ventrals and subcaudals counts, having 158–164 ventrals in males and 166–182 in females, 84–93 subcaudals in males and 87–91 in females. Drymoluber apurimacensis has 14–16 maxillary teeth. Small specimens of D. dichrous have dark crossbands 1.5–7 scales wide (mean 3.6) and light interspaces 0.5– 2.5 scales wide (mean 0.8), while in D. apurimacensis the dark crossbands are 1–2 scales wide, and the pale interspaces are 2–3 scales wide. Juvenile specimens of D. brazili have dark crossbands of similar width to those of D. dichrous (2–6 scales; mean 3.6), but the pale interspaces are wider (0.5–5 scales; mean 1.6). The hemipenes of D. dichrous tend to have more calyces than D. brazili, smaller spinulated flounces, and no spines in the lobular region. The walls of the sulcus spermaticus tend to have more ornamentation, at least in the lobular region, with small jagged papillae. The spines of the asulcate face are generally larger than those of D. brazili, especially those most proximal. The hemipenial morphology of D. dichrous and D. apurimacensis is similar and of little value in differentiating these species. Description of the lectotype (Fig. 14): Snout-vent length 585 mm, and tail length 242 mm; head distinct from the body, 24.6 mm length (4.2% of the SVL); greatest width of head 12.9 mm (52% of its length); width of head at the supraoculars 9.3 mm; internasal distance 5.1 mm; eye diameter 4.45 mm; eye-nostril distance 4.5 mm. The morphometric measurements were taken by Christoph Kucharzewski, Museum für Naturkunde Berlin. Smooth dorsal scales in 15-15-15 rows, with two apical pits; 161 ventrals and 1 preventral (sensu Peters 1964); cloacal shield entire; tail intact, with 96 divided subcaudals and one terminal spine; rostral wider than high, visible from above; internasals and prefrontals slightly wider than long; each prefrontal contacting the frontal, supraocular, internasals, posterior nasal, preocular and loreal; frontal about 1.5 times longer than wide; supraoculars longer than wide; parietals about 1.5 times longer than wide; nasal divided above and below the naris, mainly in contact with the first supralabial, but also with the second; loreal slightly longer than high, contacting the second and third supralabials; one preocular; two subequal postoculars; three anterior temporals (one upper and two lower) and two posterior temporals (one upper and one lower) on the right side (1/2+1/1); four anterior temporals (two uppers and two lowers) and two posterior temporals (one upper and one lower) on the left side (2/2+1/1); eight supralabials, the fourth and the fifth contacting the eye; mental triangular, wider than long; nine infralabials, the first pair in contact behind the mental; first to fifth infralabials in contact with the first pair of chinshields; fifth and sixth infralabials in contact with the second pair of chinshields; sixth to ninth infralabials contacting the gulars; first pair of chinshields about the half of the length of the second.

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FIGURE 13. Itinerary of the travels made by Georg Wilhelm Freyreiss in Brazil, in the 19th century, when the lectotype (ZMB 1661) and one paralectotype (ZMB 1662) of Drymoluber dichrous were collected. The inset map shows South America, highlighting in gray the current states of Rio de Janeiro (RJ), Espírito Santo (ES), Minas Gerais (MG) and Bahia (BA), visited by Freyreiss. For easy viewing, only the main localities are represented in the main map. For complete lists of localities, see Freyreiss (1907) and Wied (1989), Bokermann (1957) and Papavero (1971). When the old and current names of a toponym are different, the later is written inside brackets. 1 = Praia dos Mineiros, Rio de Janeiro; 2 = Porto Estrela (Magé); 3 = Fazenda Mandioca (Magé); 4 = Fazenda do Padre Correia (Corrêas, Petrópolis); 5 = Fazenda das Sebollas (Inconfidência, Paraíba do Sul); 6 = Rio Paraibuna; 7 = Matias Barbosa; 8 = Juiz de Fora; 9 = Chapéu D’Uvas (Juiz de Fora); 10 = Barbacena; 11 = Congonhas do Campo (Congonhas); 12 = Villa Rica (Ouro Preto); 13 = Ponte das Almoreiras (Ponte das Almorreimas, Brumadinho); 14 = Fazenda São Joanico (Maravilhas); 15 = Pompeu; 16 = Rio São Francisco; 17 = Fazenda do Comandante de Indaiá (Quartel Geral); 18 = Quartel do Assunção (Córrego dos Tiros, Tiros); 19 = Mariana; 20 = Santana dos Ferros (Guaraciaba); 21 = Santa Rita (Viçosa); 22 = Serra de S. Beralde (Serra de São Geraldo, São Geraldo); 23 = Presídio de São João Batista (Visconde do Rio Branco); 24 = S. Gonzalves (São Gonçalo); 25 = Freguesia de Maricá (Maricá); 26 = Araruama; 27 = Cabo Frio; 28 = Villa de S. João de Macahé (Macaé); 29 = Villa de S. Salvador dos Campos dos Goytacazes (Campos dos Goytacazes); 30 = Itapemerim; 31 = Povoação de Piuma (Piúma); 32 = Nossa Senhora da Victoria (Vitória); 33 = Vila Nova do Almeida (Nova Almeida, Serra); 34 = Quartel do Riacho (Riacho, Aracruz); 35 = Linhares; 36 = Barra do São Mateus (São Mateus); 37 = Rio Mucuri (Mucuri); 38 = Colônia Leopoldina (Nova Viçosa); 39 = Caravelas; 40 = Canavieiras; 41 = Salvador. Dashed line = Travel from Rio de Janeiro to Quartel do Assunção (Tiros), in 1814 (Localities 1–12, 12–11, 11–18); Dotted line = Travel from Villa Rica (Ouro Preto) to the vicinities of Presídio São João Batista (Visconde do Rio Branco), from December 1814 to January 1815 (Localities 12–23); Dashed-dotted line = Travel from Rio de Janeiro to Mucuri, from July 1815 to 1816 (Localities 1, 24–38). Localities 39–41 are not linked by lines because of the lack of information about the itinerary took by Freyreiss when visiting them. Tropical and subtropical moist forests; Tropical and subtropical savannas; Deserts and xeric formations; Mangroves; Tropical and subtropical dry forests. Habitat types follow Olson et al. (2001).

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FIGURE 14. Lectotype of Drymoluber dichrous (Peters, 1863) (ZMB 1661). A) dorsal view of body; B) ventral view of body; C) right side of the head; D) left side of the head; E) dorsum of head; F) gular region. Photos: Mark-Oliver Rödel. TAXONOMIC REVISION OF DRYMOLUBER

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Coloration of the lectotype: Peters (1863) described the coloration of Herpetodryas dichroa as: “Dorsum olive-brown, the sides, including the lateral edges of ventral plates; subcaudals olive-green; the whole venter to the tip of the tail of yellow color; a black stripe bordered by the yellow (darker than that of the belly) supralabials on both sides of the head, becoming suddenly wider behind the eyes.”. After almost 200 years of preservation, the dorsum is uniformly brownish-blue. Supralabials, gular region and venter are uniformly creamish colored, the lateral edges of ventrals and subcaudals are the same color as the dorsum. The upper margins of some supralabials, especially the last two, have the color of the dorsum. The dark lateral stripe of the head is indistinct. Coloration of preserved adults: The dorsal coloration of adult specimens after fixation is usually darker than the lectotype, with bluish-gray or dark-blue colors. Faint crossbands that formed the juvenile color pattern are visible in smaller specimens. The venter of most specimens is immaculate cream, with lateral edges of ventral plates having the same color as the dorsal scales (n=203; 95%). In some cases (n=3; 1.5%) the dark dorsal coloration continues into the ventral region from the lateral edges of ventral plates. There are specimens with a creamish venter with small black marks (n=3; 1.5%). The venter of some specimens is the same color as dorsal scales or a little paler (n=3; 1.5%). One specimen (0.05%) has a yellow venter, with the lateral and the posterior edges of ventral plates darkened. The ventral part of the tail usually is creamish with lateral edges of subcaudals darkened (n=207; 97%), with little variation. The lateral edges may not be darkened (n=1; 0.05%), the subcaudals may have small black dots, in larger numbers in the posterior region (n=1; 0.05%), or be completely black along the whole tail (n=4; 2%). The gular region is pale and immaculate in most specimens (n=153; 72%). Some of them, however, have dark marks in the posterior edges of infralabials, and sometimes on the chinshields (n=60; 28%). The lateral edges of supralabials (especially the last ones) in most specimens are dark colored (n=166; 78%), sometimes (n=14; 6.5%), with strong and thick marks. Supralabials can also be completely dark (n=15; 7%) or totally cream, without marks, or with inconspicuous marks (n=18; 8.5%). Coloration of adults in life: Based on descriptions in the literature (e.g. Cunha & Nascimento 1978; Martins & Oliveira 1998; Bartlett & Bartlett 2003; Argôlo 2004a) and some photographs (Fig. 15), we noted some variation in the color pattern of adult specimens of D. dichrous in life—but unrelated to the geographic distribution. The dorsum varies between brown, olive-brown, green, dark-green and gray-bluish. In some snakes the dorsal color changes posterior to the first third or the half of the body. The dorsum of head is sometimes a little paler than the body. A barely distinguishable black stripe from the preocular to the end of the posterior temporal may be present. Supralabials, gular region and the venter vary between yellow and white, with the lateral edges of ventrals and subcaudals of the same color or little paler than the dorsals. The supralabials also may be dark along their upper and lateral edges. Coloration of preserved juveniles: The number of dark crossbands along the body varies between 31 and 52 (mean 40; SD=4.3; n=49; 63%). Specimens with indistinct bands on the posterior third of the body are common (n=29; 37%), and in seven small specimens (SVL 285–391 mm), the banded coloration has disappeared completely. The tail bands are rarely visible (only in some specimens with SVL < 385 mm). Dorsal crossbands vary from de 1.5–7 vertebral/paravertebral scales wide (mean 3.6; SD=0.87; n=306 crossbands). The last crossband anterior to the cloacal shield tends to be the narrowest (1.5–5 scales; mean 2.9; SD=0.79; n=48) and the fifteenth after the head the widest (2–7 scales; mean 3.6; SD=0.75; n=71). The light interspaces between dark crossbands vary between 0.5–2.5 scales (mean 0.8; SD=0.43; n=306 interspaces). The interspaces anterior to the last crossband and the fifth crossband anterior to the cloacal shield are the narrowest, with 0.5–1 scales (mean 0.5; SD=0.1; n=48; and mean 0.5; SD=0.09; n=49). The interspace posterior to the first crossband and the interspace anterior to the fifth crossband are the widest, with 0.5–2.5 scales (mean 1.17; SD=0.49; n=67; and mean 0.96; SD=0.46; n=71). The venter of juveniles usually has a creamish coloration as in adults (n=66; 85 %). It also may have black marks spreading ventromedially from the lateral edges of ventral plates (n=9; 11%), or even be completely black along the whole body (n=1; 1.5%), or only on the posterior half (n=2; 2.5%). The subcaudals in juveniles are the same pattern as those in adults (cream color with darkened lateral edges) (n=73; 93.5%), although in some specimens these plates are completely black (n=5; 6.5%). The head pattern consists of light internasals; light prefrontals with darkened posterior edge; and dark frontal and supraoculars with light anterior edge. A light stripe in the parietal region may be present and immaculate (n=19; 26.1%), maculate (n=5; 6.8%), or absent (n=49; 67.1%) (Fig. 13).

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Like in the adults, the gular region of the juveniles usually is cream colored (n=48; 61.5%), but black marks may occur on the infralabials and chinshields (n=30; 38.5%). The supralabials have distinct dark marks on their lateral edges (n=51; 65.4%), but sometimes these marks are pale, as in adults (n=8; 10.25%), or even barely visible (n=19; 24.35%). Coloration of juveniles in life: The light colored regions of preserved juveniles may vary from shades of white, cream, light-brown and orange in life. The dark colored regions of preserved specimens vary in shades of brown, orange-brown, reddish-brown and grayish-brown (Fig. 16).

FIGURE 15. Color in life of some adult specimens of Drymoluber dichrous (Peters, 1863). A) Barra do Choça, Bahia, Brazil; B) Reserva Extrativista Riozinho da Liberdade, Tarauacá, Acre, Brazil; C) Reserva Ducke, Manaus, Amazonas, Brazil; D) Nouragues Station, French Guiana; E) Parque Estadual Guajará-Mirim, Rondônia, Brazil; F) Marabá, Pará, Brazil. Photos: Marco Antônio de Freitas (A), Paulo Sérgio Bernarde (B), Rafael de Fraga (C), Maël Dewynter (D), Laurie J. Vitt (E) and Pedro L. V. Peloso (F).

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TABLE 5. Summary of geographic distribution and morphological variation of the three species of Drymoluber Amaral, 1930. Most common (over 50% of frequency) character states are represented in bold.

Geographic distribution

Drymoluber apurimacensis

Drymoluber brazili

Drymoluber dichrous

South of the Apurimác and

Cerrado, Caatinga, Atlantic Forests of the lower Doce

Eastern Andes, Amazonia, Guiana Shield,

Pampas Rivers, in the Peruvian

River, and Atlantic Forest–Cerrado transitional areas.

Brazilian Atlantic Forest, Atlantic Forest–

Serranía Esteparia ecoregion.

Caatinga and Atlantic Forest–Cerrado transitional areas.

Maximum snout-vent length

670 mm (MHNSM 18647)

1178 mm (IBSP 34369)

1050 mm (MPEG 17235)

Number of dorsal scale rows

13–13–13

17–17–15

15–15–15 (n=289); 15–15–17 (n=1); 15–15–13 ( n=1)

Dorsal apical pits

0

2

2

Cloacal shield

Entire

Entire (n=82); Divided (n=1)

Entire (n=290); Divided (n=1)

Number of pre-ventrals

1 (n=1); 2 (n=2)

1 (n=74); 2 (n=8); 3 (n=1)

0 (n=12); 1 (n=277); 2 (n=2)

Number of ventrals in males

158–164* (n=2)

182–200 (mean 189; SD=3.61; n=46)

157–173 (mean 165; SD=3.35; n=172)

Number of ventrals in females

166–182 (n=2)

185–202 (mean 193; SD=3,68; n=37)

160–180 (mean 172; SD=3.72; n=114)

Number of subcaudals in males

84–93 (mean 88; SD=4.58; n=3)

109–127 (mean 118; SD=4.37; n=29)

87–110 (mean 97; SD=4.20; n=122)

Number of subcaudals in females

87–91 (mean 89; SD=2.08; n=3)

109–126 (mean 119; SD=4.45; n=26)

86–109 (mean 98; SD=4.79; n=89)

Loreal

longer than high

longer than high

longer than high (n=193); as high as long (n=92); higher than long (n=5)

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Preoculars

1

1

1 (n=572 sides); 2 (n=10 sides)

Postoculars

2

2 (n=159 sides); 3 (n=5 sides)

1 (n=1 side); 2 (n=551 sides); 3 (n=28) sides

Temporal formula

1+1/1

1+1/1 (n=9 sides), 1+2/3 (n=1 side), 1/1+1 (n=1 side),

1+1 (1 side), 1+1/1 (n=38 sides), 1/1+1 (n=2

1/1+1/1 (n=90 sides), 1/1+1/2 (n=8 sides), 1/1+2/1 (n=10

sides), 1/1+1/1 (n=480 sides), 1/1+1/2 (n=7

sides), 1/1+2/2 (n=1 side), 1/1+3/1 (n=1 side), 1/2+1/1

sides), 1/1+2/1 (n=8 sides), 1/1+2/2 (n=1

(n=19 sides), 1/2+1/2 (n=5 sides), 1/2+2/1 (n=5 sides),

side), 1/2+1/1 (n=20 sides), 2/1+1/1 (n=14

1/2+2/1/2 (n=1 side), 1/2+3/1 (n=1 side), 1/2+4/1 (n=1

sides), 2/1+1/2 (n=1 side) or 2/2+1/1 (n=4

side), 2/1+1/1 (n=5 sides), 2/1+1/2 (n=3 sides), 2/2+2/2

sides)

(n=1 side), 2/2+4/2 (n=1 side), 2/3+1/2 (n=1 side)

……continued on the next page

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TABLE 5. (Continued) Drymoluber apurimacensis

Drymoluber brazili

Drymoluber dichrous

Number of supralabials

8

7 (n=3 sides); 8 (n=143 sides); 9 (n=18 sides)

7 (n=2 sides); 8 (n=540 sides); 9 (n=37 sides)

Supralabials contacting the eye

iv–v

iii–v (n=1 side); iv–v (n=152 sides); iv–vi (n=2 sides);

iii–iv (n=1 side); iii–v (n=454 sides);

v–vi (n=9 sides)

iii–vi (n=2 sides); iv–v (n=88 sides);

orbit

iv–vi (n=34 sides) Number of infralabials

7 (n=2 sides); 8 (n=4 sides);

8 (n=8 sides); 9 (n=144 sides); 10 (n=11 sides)

9 (n=2 sides) Infralabials contacting the

st

8 (n=84 sides); 9 (n=489 sides); 10 (n=7 sides)

st

1 pair

1 pair

1st pair

i–iv (n=4 sides); i–v (n=2 sides)

i–iii (n=1 side); i–iv (n=21 sides);

i–iv (n=334 sides); i–v (n=244 sides);

i–v (n=140 sides); i–vi (n=2 sides)

i–vi (n=2 sides)

iii (n=1 side); iv (n=1 side); iv–v (n=20 sides);

iv–v (n=12 sides); iv–vi (n=322 sides);

v (n=134 sides); v–vi (n=8 sides)

v–vi (n=240 sides); v–vii (n=4 sides);

mental Infralabials contacting the first pair of chinshields Infralabials contacting the

iv–v (n=4 sides); v (n=2 sides)

second pair of chinshields

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vi–vii (n=2 sides) Infralabials contacting the gulars

v–vii (n=2 sides);

iii–viii (n=1 side); iv–viii (n=1 side);

v–viii (n=4 sides); v–ix (n=7 sides);

v–viii (n=4 sides) **

v–viii (n=4 sides); v–ix (n=143 sides);

vi–viii (n=75 sides); vi–ix (n=485 sides);

v–x (n=6 sides); vi–ix (n=4 sides);

vi–x (n=2 sides); vii–x (n=6 sides)

vi–x (n=4 sides) Maxillary teeth

14–16 (mean =15; SD=1)

19–25 (mean =23; SD=1.05)

19–26 (mean =23; SD=1.21)

* Lehr et al. (2004) cited 182 ventrals for MTKD 44669 (male), but this specimen is in poor condition (Fig. 23B), with at least three regions where the scale count is impaired; thus, we do not consider the countings of Lehr et al. (2004) in the current work. ** We have no information concerning the contact between infralabials, chinshields and gulars for MHNSM 18647, which, according to Lehr et al. (2004) has nine infralabials.

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FIGURE 16. Color in life of some juvenile specimens of Drymoluber dichrous (Peters, 1863). A) Reserva Ducke, Manaus, Amazonas, Brazil; B) Reserva Ducke, Manaus, Amazonas, Brazil; C) Barra do Choça, Bahia, Brazil; D) Ubajara, Ceará, Brazil. Photos: Rafael de Fraga (A), William Quatman (B), Marco Antônio de Freitas (C) and Daniel Loebmann (D).

Hemipenial morphology (n=17) (Fig. 17): Hemipenis single, subcylindrical, not capitate. Sulcus spermaticus single and centrolinear. Lobe about half of the hemipenis length, with papillate calyces (5–10 triangular papillae per calyx). Proximally, the calyces are gradually replaced by spinulate flounces and spines. Body covered by spines arranged in more or less transverse rows (about 60–70 spines in total). Walls of the sulcus spermaticus ornamented at least in the lobular region by jagged papillae (sometimes in small number), and some spinules. These walls are also bordered on both sides by a longitudinal row of 6–10 spines (mean 7; SD=1.22) increasing in size toward the proximal region. The spines from the left side may increase in size up to the middle of the row, and then decrease in size toward the proximal region. A hook is present at the end of each row of spines bordering the sulcus. Both left (n=11) and right (n=2) hooks can be located more proximally than the other hook, or reach the same position in the hemipenis base (n=4). None (n=8), or three to four small spines (n=7) or spinules (n=2) may be present between the left hook and the wall of the sulcus spermaticus. One (n=10) or two (n=2) lateral spines, larger than the hooks, may be present to the left of the sulcus spermaticus, or a single large lateral spine may be present on both sides of the sulcus (n=2). In some cases (n=3), lateral spines are smaller than the hooks. The asulcate face is formed by spines arranged in 5–7 more or less transverse rows (counted from proximal to distal region), with largest spines in the median rows. The base of the hemipenis is smooth, with some grooves and several sparse spinules. Variation: Largest male with SVL 1050 mm, TL 340+N mm (MPEG 17235); largest female with SVL 801 mm, TL 89+N mm (IBSP 2198). Seventy seven (77) examined specimens (26.5%) had broken tail. The tail of 54 of those specimens (18.5% of the total examined) was healed, suggesting that the breakage did not occur during collection or preservation, and is evidence of the presence of pseudoautotomy in this species, a defensive behavior

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already recorded in several other Neotropical Colubrinae, including Dendrophidion, Drymobius, Mastigodryas and Scaphiodontophis (Mendelson III 1992; Slowinski & Savage 1995; Prudente et al. 2007; Leite et al. 2009). Tail length of specimens with complete tail is 25.9–48.4% of the SVL in males (mean 40.6%; SD=3.18; n=122) and 35.4–65.3% in females (mean 41%; SD=3.75; n=89). For variation in meristic characters, see Table 5.

FIGURE 17. Hemipenis of Drymoluber dichrous (FMNH 40206, La Pampa, Madre de Dios, Peru). A) Sulcate face; B) Asulcate face. Scale bar = 1 cm. Photos: Henrique C. Costa.

Geographic distribution: Drymoluber dichrous inhabits the eastern portion of the Andean mountain range (Peru and Ecuador), the Amazonia and Guiana Shield (Bolivia, Peru, Ecuador, Colombia, Venezuela, Brazil, Guyana, Suriname and French Guiana), and the Atlantic Forest and its transitional areas with the Caatinga (brejos nordestinos) and Cerrado domains (Fig. 12; Appendices II and III). The Andes of Colombia and the Venezuelan Llanos may limit the northward expansion of D. dichrous, while the transition from tropical to subtropical climate below the Tropic of Capricorn may act as a southern barrier. The Andes may also limit its western distribution, while in the east the species reaches the coast. The elevational distribution of D. dichrous is extremely wide, varying from sea level to about 3500 meters. The absence of morphological characters distinguishing the Amazonian and Atlantic Forest populations of Drymoluber dichrous could be due to genetic continuity maintained until the Pleistocene, when both forested regions were in contact; today this past area of contact is located in the Caatinga (Vanzolini 1981; Rodrigues 1990; Costa 2003) and part of the Cerrado (Ledru 1993; Costa 2003). Future phylogenetic analyses can help solving the issue concerning where was the bridge between the Amazonia and the Atlanctic Forest that maintained the genetic flux of D. dichrous in the past. Natural history: Drymoluber dichrous inhabits mainly forests with additional records from natural savannas TAXONOMIC REVISION OF DRYMOLUBER

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and environments with a relative degree of anthropization (Cunha & Nascimento 1978; Martins & Oliveira 1998; Argôlo 2004a; França et al. 2006). Drymoluber dichrous is diurnal, forages on the ground and rests on the vegetation at night (Cunha & Nascimento 1978; Martins & Oliveira 1998). Its diet seems to be represented mainly by terrestrial anurans and lizards (including their eggs), but also snakes (even cospecifics). The following taxa were recorded as prey of D. dichrous: Adelophryne maranguapensis, Alobates aff. marchesianus, Anomaloglossus (=Colostethus) sp., Ischnocnema cf. ramagii, Leptodactylus sp., L. didymus and Physalaemus gr. cuvieri (Anurans); Ameiva ameiva, Anolis fuscoauratus, Arthrosaura reticulata, Cercosaura eigenmanni, Ce. ocellata, Coleodactylus meridionalis, Colobosauroides cearensis, Gonatodes hasemani, Go. humeralis, Iphisa elegans, Kentropyx calcarata, Leposoma percarinatum, Le. baturitensis, Neusticurus ecpleopus and Placosoma sp. (Lizards); Oxybelis sp. and Thamnodynastes hypoconia (Snakes) (Cunha & Nascimento 1978; Duellman 1978, 2005; Dixon & Soini 1986; Avila-Pires 1995; Martins & Oliveira 1998; Vitt et al. 2000; Borges-Nojosa & Lima 2001; Pinto 2006; Bernarde & Abe 2010; Veríssimo et al. 2012). During the present study, three newborns of an opossum species (Didelphidae) and the remains of an amphisbaenian (Amphisbaenidae) where found in the stomach of a previously dissected specimen (MPEG 2670, male, 920 mm SVL). These food items constitute new registers for the species’ diet. Information about reproduction in D. dichrous is scarce and restricted to Amazonian specimens. Drymoluber dichrous is oviparous and there are reports of 2–6 eggs per clutch (Fitch 1970; Martins & Oliveira 1998). Three females examined in this study had four (MPEG 10832, MPEG 16551, MPEG 19007) and one (MPEG 20330) had five eggs. This information, together with other literature records, indicates an average of four eggs per clutch. The reproductive season in Amazonia seems to be prolonged throughout the year, according data from Fitch (1970) and Martins & Oliveira (1998). Drymoluber dichrous has several defensive behaviors. When handled, individuals will rotate their body, vibrate their tail and occasionally bite (Martins & Oliveira 1998). Martins et al. (2008) observed three types of defensive behaviors directed to visually oriented predators: crypsis, head elevation and neck S-coil. Brodie III & Brodie Jr. (2004) suggested that young cross-banded specimens might mimic coralsnakes (Elapidae). As mentioned above, the long tail of D. dichrous (mean 40.6% of the SVL in males and 41% in females), with a tail breakage ratio of 18.5% (considering only those specimens where breakage occurred before collection) is evidence of the defensive behavior known as pseudoautotomy (Slowinski & Savage 1995). This defensive behavior may explain the long tail in this primarily terrestrial species. The presence of sexual dimorphism in D. dichrous, with males larger than females, was already observed by Fitch (1981). The longer tail of males of D. dichrous (mean 277 mm) related to females (mean 253 mm) is a common character in snakes (Shine et al. 1999; Pizzatto et al. 2007). Future research on the reproductive biology of the species would provide more information and a better understanding of these topics. Etymology: Peters (1863) did not comment on the reasons that led him to choose the name dichroa for the species he described. The words dichroa / dichrous comes from the ancient Greek, meaning “two-colored” or “two-skins”, and we suggest that the name may be an allusion to the contrasting color of adult specimens, which dorsum is dark colored, while the venter is light colored.

Drymoluber brazili (Gomes, 1918) Drymobius brazili Gomes, 1918. Memórias do Instituto Butantan, 1, p. 81. Holotype: IBSP 696 (probably destroyed during the fire of May 2010). Drymobius rubriceps Amaral, 1923. Proceedings of the New England Zoological Club, 8, p. 85. Holotype: IBSP 1844 (probably destroyed during the fire of May 2010). Drymobius boddaerti (partim.)—Amaral 1929. Memórias do Instituto Butantan, 4, p. 11. Drymoluber brazili—Stuart 1932. Occasional Papers of the Museum of Zoology, University of Michigan, 236, p. 4.

The types of Drymoluber brazili and its junior synonym, Drymobius rubriceps, were personally examined by the senior author in 2009. On May 2010, the collections of Instituto Butantan in São Paulo, Brazil, where those and most known specimens of D. brazili were maintained were consumed in a fire (Warrell et al. 2010; Franco 2012). It is estimated that 80% of the snake collection was destroyed, but there is still no information if the types of Drymoluber brazili were lost (F. Franco, pers. comm.).

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Holotype: Instituto Butantan, São Paulo, IBSP 696, adult male, SVL 1090 mm, TL 473 mm, collected on September 1914 in the Estação Ferroviária de Engenheiro Lisboa (currently inactive), near the municipality of Uberaba (-19.80, -47.60), Minas Gerais, Brazil. Specimen personally examined by the senior author. Paratypes: Instituto Butantan, São Paulo, IBSP 383, adult male, SVL 863 mm, TL 394 mm, collected in February 1913, in the Estação Ferroviária Santa Eudóxia (currently inactive), municipality of São Carlos (-21.77, -47.78), São Paulo, Brazil; IBSP 573, adult female, SVL 854 mm, TL 422 mm, collected in February 1914, in the Estação Ferroviária de Sampaio Vidal (currently inactive), municipality of Ribeirão Bonito (-22.07, -48.18), São Paulo, Brazil; IBSP 574, adult male, SVL 862 mm, TL 268+N mm (broken tail), without locality data; IBSP 741, adult male, SVL 1120 mm, TL 162+N mm (broken tail), collected in December 1914, in the Estação Ferroviária Java (currently inactive), municipality of Boa Esperança do Sul (-21.99, -48.39), São Paulo, Brazil; IBSP 1286, adult female, SVL 898 mm, TL 402 mm, collected in May 1917, in the Estação Ferroviária Pedregulho (currently inactive), municipality of Pedregulho (-20.26, -47.48), São Paulo, Brazil. All specimens personally examined by the senior author. About the type locality: In the 20th century the Instituto Butantan often received snakes from several Brazilian localities, sent by collectors to the institute via the railroads that crossed some regions of the country at that time. Thus, the accuracy of localities of specimens from railroads (like the type series of D. brazili) must be treated with caution (Pereira et al. 2007). In the specific case of D. brazili, the Estação Ferroviária Engenheiro Lisboa (type locality), was located at km 555 of the Tronco-Catalão railroad, which, at the time of the collection of the holotype (September 1914) went from Campinas, São Paulo (-22.09, -47.05) to Ipameri, Goiás (-17.72, -48.15) (Giesbrecht 2009; Cavalcanti 2010). Although it is hard to identify the exact locality where the holotype of D. brazili was collected, we believe it really was in the proximities of the Tronco-Catalão railroad, since the road is located in areas with other confirmed records of the species (see map in Fig. 12). Similar locality problems exist with the paratypes that purportedly also are from rail stations. The Santa Eudóxia, Sampaio Vidal and Java stations, were part of a wide railroad network belonging to the Companhia Paulista de Estradas de Ferro that crossed the state of São Paulo from the southeast to the north and northwest (Giesbrecht 2009; Cavalcanti 2010). On the other hand, the Estação Ferroviária Pedregulho was located at km 455 on the Tronco-Catalão railroad. Diagnosis: Drymoluber brazili can be distinguished from D. apurimacensis and D. dichrous by the following combination of characters: a) 17-17-15 dorsal scale rows with two apical pits; b) 182–200 ventrals in males, 185– 202 in females; c) 109–127 subcaudals in males, 109–126 in females; d) 19–25 maxillary teeth. See Table 5. Comparisons: Drymoluber apurimacensis has 13-13-13 dorsal scales rows, and D. dichrous has 15-15-15. Apical pits are absent in D. apurimacensis. Drymoluber apurimacensis has 158–164 ventrals in males and 166–182 in females, 84–93 subcaudals in males and 87–91 in females. Drymoluber dichrous has 157–173 ventrals in males and 160–180 in females, 87–110 subcaudals in males and 86–109 in females. Drymoluber apurimacensis has 14– 16 maxillary teeth. Small specimens of Drymoluber brazili have dark crossbands 2–6 scales wide (mean 3.6) and light interspaces 0.5–5 scales wide (mean 1.6), while in a single specimen of D. apurimacensis the dark crossbands are 1–2 scales wide and the light interspaces 2–3 scales wide. Young specimens of D. dichrous have dark crossbands with similar width to those of D. brazili (1.5–7 scales, mean 3.6), but the pale interspaces are on average narrower (0.5–2.5 scales, mean 0.8). The hemipenes of Drymoluber brazili tend to have fewer calyces than that of D. dichrous and D. apurimacensis, larger spinulate flounces, and spines in the lobular region. The walls of the sulcus spermaticus are less ornamented. The spines of the asulcate face in general are smaller than those of D. dichrous and D. apurimacensis, especially those most proximal. Redescription of the holotype (Fig. 18): Snout-vent length 1090 mm and tail length 473 mm; head distinct from the body, 35.1 mm long (3.2% of the SVL); greatest width of head 15.20 mm (43% of head length); width of head at the supraoculars 11.3 mm; internasal distance 6.1 mm; eye diameter 6.0 mm; eye-nostril distance 5.9 mm. Smooth dorsal scales in 17-17-15 rows with two apical pits; 190 ventrals and 1 preventral (sensu Peters 1964); cloacal shield entire; tail intact with 117 divided subcaudals and one terminal spine; rostral wider than high, visible from above; internasals and prefrontals slightly wider than long; each prefrontal contacting the frontal, supraocular, internasals, posterior nasal, preocular and loreal; frontal about 1.5 times longer than wide; supraoculars longer than wide; parietals about 1.5 times longer than wide; nasal divided above and below the naris, mainly in contact with

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the first supralabial, but also with the second; loreal slightly longer than high, (vaguely resembling a parallelogram), contacting the second and third supralabials; one preocular; two postoculars, the upper one larger than the lower; two anterior temporals (one upper and one lower) and two posterior temporals (one upper and one lower) (1/1+1/1) on both sides of the head; eight supralabials, the fourth and the fifth contacting the eye; mental triangular, wider than long; nine infralabials, the first pair in contact behind the mental; first to fifth infralabials in contact with the first pair of chinshields; fifth infralabial in contact with the second pair of chinshields; fifth to ninth infralabials contacting the gulars; first pair of chinshields about the half the length of the second; 23 maxillary teeth, increasing in size posteriorly. There are some small differences between the redescription of the holotype presented here (inside brackets) and the original description. Gomes (1918) decribed the holotype as having 22 maxillary teeth, internasals as wide as long, 10 infralabials, 191 ventrals, SVL 1110 mm and TL =480 mm. The difference in the number of infralabials may be explained on the basis of the method used in both studies. In the present work, the posterior boundary of the infralabial row is defined as the last scale completely covered by the last supralabial (Peters 1964) (Fig. 18C), which is not the definition seemingly used by Gomes (1918). The differences in SVL and TL may be due to a small error in the original measuring, or a reduction in size caused by fixation and preservation over time time (Vervust et al. 2009; Guimarães et al. 2010). This situation is also applicable to the paratypes of D. brazili. Coloration of the holotype: Gomes (1918) described the holotype in life as having olive-green color in the head and on the anterior part of the body, becoming reddish-brown in the posterior part of body and on the tail. The venter was yellowish-white, while the lateral edges of the ventral plates were the same color as the dorsum. When the senior author examined the holotype, it was grayish-brown on the dorsum of head and anterior half of the body, becoming yellowish-brown on the posterior half of the body and tail. Supralabials, gular region and venter were yellowish-brown and similar to or slightly paler than the dorsum. Coloration of preserved adults: The dorsal coloration of most of the examined adults (n=49; 77.8%) is uniform bluish-gray, bluish-brown or yellowish-brown throughout. A few specimens (n=13; 20.6%) are grayishbrown or bluish-brown on the anterior half of the body and yellowish-brown on the posterior half. One specimen has faint crossbands as are present in the juvenile color pattern. The venter of most specimens (n=49; 77.8%) is immaculate, yellowish-cream, and the lateral edges of ventrals have the same color as the dorsal scales. In one specimen (1.6%) the dorsal coloration extends into the ventral region; in other specimens (n=13; 20.6%) the venter is yellowish-colored with some dark spots, and the lateral edges of ventrals are the same color as the dorsal scales. Dorsally, the tail is the same color as the body, but shows some variations in the subcaudal coloration. In 24 specimens (38.1%) the subcaudals are yellowish-cream, immaculate, and with their lateral edges the same color as the dorsum. In some snakes, the lateral edges are inconspicuous and the same color as subcaudals (n=5; 7.9%). In a few, the subcaudals have more than half their area dark like the dorsals (n=1; 1.6%), or have black spots throughout their area (n=1; 1.6%). In others, black spots are numerous only in the posterior region of the tail (n=24; 38.1%), or the posterior part of tail is completely black (n=8; 12.7%). In all specimens the head is uniform dorsally and the same color as the dorsum of body. The gular region is immaculate, yellowish-cream. The supralabials of most adult specimens have dark lateral edges (mainly in the last scales (n=59; 93.6%), but in some specimens visible markings are absent (n=4; 6.4%). Coloration of adults in life: Unlike Drymoluber dichrous, there is little information about the coloration of D. brazili in life. Gomes (1918) described the the type series in life as follows: two specimens have the dorsum completely olive-green, while in the other three, and the holotype, the posterior part of the dorsum is reddishbrown. This same pattern is seen in the illustration (Fig. 19A) of one specimen from Visconde de Parnaíba, São Paulo (IBSP 4812) published by Amaral (1978). In some specimens, the dorsal color may consist of grayish tones (Fig. 19B). Coloration of preserved juveniles: The color pattern of preserved juveniles of Drymoluber brazili is similar to that of the other species of the genus, consisting of dark crossbands separated by pale interspaces. Only two specimens (10%) have crossbands (47 and 54) visible throughout the body (MZUSP 9596 and IBSP 29221). Crossbands on the tail were not visible in any specimen. The dark crossbands varied in width between 2–6 vertebral/paravertebral scales (mean 3.6; SD=0.89; n=62 crossbands). The last crossband anterior to the cloaca is the narrowest and two scales wide (n=2; 10%), while the first crossband on the body is the widest, 2–6 scales wide (mean 4; SD=0.95; n=20; 100%). The widths of the pale interspaces vary between 0.5–5 scales (mean 1.6;

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SD=0.97; n=62 interespaces). The interspace anterior to the fifth crossband anterior to the cloaca is the narrowest, with 0.5 or 1.5 scales (n=2; 10%). The interspace posterior to the first crossband is the widest, with 1–5 scales (mean 2.32; SD=0.90; n=20; 100%). The venter of young specimens is always yellowish-cream and immaculate, and the lateral edges of ventral plates are the same color as the dorsals.

FIGURE 18. Holotype of Drymoluber brazili (Gomes, 1918) (IBSP 696). A) dorsal view of body; B) ventral view of body; C) right side of the head; D) left side of the head; E) dorsum of head; F) gular region. Photos: Henrique C. Costa (A–D, F) and Ana Bárbara Barros (E). TAXONOMIC REVISION OF DRYMOLUBER

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FIGURE 19. Color in life of some adult specimens of Drymoluber brazili (Gomes, 1918). A) Visconde de Parnaíba, São Paulo, Brazil (Illustration by R. K. Leyer presented in Amaral [1978]); B) UHE Peixe-Angical, Tocantins, Brazil. Photo: Pedro Henrique Bernardo.

FIGURE 20. Color in life of juvenile specimens of Drymoluber brazili (Gomes, 1918). A) Parque Nacional das Emas, Goiás, Brazil (IBSP 62682); B) Unknown locality data. Photos: Paula H. Valdujo (A) and Eduardo Santos (B).

The subcaudals are cream colored, with the lateral edges darkened (n=9; 45%), cream (n=6; 30%), or cream with only the posterior edges darkened (n=5; 25%). The head has light internasals, light pre-frontals with darkened posterior edges, and a dark frontal and supraoculars with light anterior edges. A light stripe in the parietal region can be present and immaculate (n=9; 45%) or maculate (n=4; 20%). In the remaining specimens (n=7; 35%) the parietals are dark (Fig. 11). As occurs in other species of the genus, lighter shades on the head darken as the specimens grow. As in adults, the gular region of the juveniles is pale and immaculate. The supralabials have faint dark markings on their lateral edges. Coloration of juveniles in life: In life, the light colored regions in preserved juveniles are shades of white, cream and light-brown, except in the head and neck, where an orange-red color prevails. The dark colored regions (including crossbands) vary in shades of brown, olive-green and dark-gray (Fig. 20; Amaral 1923). Apparently, only D. brazili juveniles have this reddish coloration on the head. However, since those colors tend to become white in preserved specimens, and because photographs and information about the coloration of live juveniles of Drymoluber are uncommon, it is difficult to know whether this character is diagnostic for D. brazili.

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Hemipenial morphology (n=6) (Fig. 21): Hemipenis single, subcylindrical, not capitate. Sulcus spermaticus single and centrolinear. Lobe about half of the hemipenis length, with papillate calyces (5–10 papillae per calyx, most calyces deep) and spines bordering the sulcus spermaticus. The calyces are gradually replaced by spinulate flounces and spines proximally. The hemipenial body is covered by spines arranged in more or less transverse rows (about 60 or a few more spines in total). Walls of the sulcus spermaticus weakly ornamented, usually with few papillae and spinules. These walls are also bordered on both sides by a longitudinal row of 8–13 spines (mean 8; SD=1.97). The spines from the right side tend to increase in size toward the proximal region, while the spines from the left side increase in size up to the middle of the row, then decrease in size toward the proximal region (n=4). In the rest of the examined specimens (n=2), spines from both sides increase toward the proximal region of the hemipenis. A basal hook is present at the end of each row of spines bordering the sulcus, with the left hook more proximally than the right hook. None (n=2), or three to four small spines (n=2) or spinules (n=2) may be present between the left hook and the wall of the sulcus spermaticus. One (n=2) or two (n=2) lateral spines, larger than the hooks, may be present to the left of the sulcus spermaticus, or two large lateral spines may be present on both sides of the sulcus (n=1). The asulcate face is formed by spines arranged in 6–7 more or less transverse rows (counted from proximal to distal region), with largest spines in the median rows, and proximal spines smaller than distal spines. In two specimens (MCNR 1736 and UFMT 6970), those spines are very small. The base of the hemipenis is smooth, with some grooves and several sparse spinules Variation: Largest male with SVL 1178 mm, TL 410+N mm (IBSP 34369); largest female with SVL 1035 mm, TL 400+N mm (IBSP 17019). Twenty eight (28) specimens examined (33.7%) had broken tail. The tail of 24 of those specimens (28.9% of the total examined) was healed, suggesting the presence of pseudoautotomy in this species. Tail length of specimens with complete tail is 38.1–62.7% of the SVL in males (mean 44.2%; SD=4.23; n=29) and 41.11–56% (mean 45.7%; SD=3.5; n=26) in females. For variation in meristic characters, see Table 5. Geographic distribution: Drymoluber brazili is distributed mainly in Brazil, with a single record from Paraguay (Canindeyú province). The species inhabits the Cerrado domain, with additional records from the Caatinga, Atlantic Forest and its transitional areas with the Cerrado (Fig. 12; Appendices II and III). França et al. (2006) reported the species in Vilhena, Rondônia, Brazil (-12.74, -60.15), in Amazonian savannas. However, that specimen (CHUNB 12791) is actually a D. dichrous. There is a record from Aripuanã, Mato Grosso, Brazil (10.15, -59.45) in the MPEG collection, but the specimen (MPEG 10419) could not be found, and we do not consider this record further. Therefore, the presence of D. brazili in the Amazonian region is unconfirmed. In their material examined section, Lehr et al. (2004) listed three specimens of Drymoluber brazili deposited in MZUSP that were misidentified: MZUSP 5329 (from Morro Branco, Ceará, Brazil) actually is a young Drymarchon corais; MZUSP 11263 (from Gaúcha do Norte, Mato Grosso, Brazil) and MZUSP 11442 (from Vila Rica, Mato Grosso, Brazil) are specimens of the genus Mastigodryas. The records of Drymoluber brazili from the Atlantic Forest, in the municipalities of Colatina (-19.54, -40.64) and Baixo Guandú (-19.52, -41.02) state of Espírito Santo, and in Aimorés (-19.50, -41.06), state of Minas Gerais, suggest relictual populations. The lower Doce River region is relatively arid with notable occurrences of rocky outcrops that resemble Caatinga areas, different from the Atlantic Forest of the nearest regions (Jackson 1978; Ribon 1995). Modelings of the Atlantic Forest range under past climatic scenarios of 6000 and 21,000 years ago suggest that much of the area south of the Doce River was not predicted to retain a large, stable forest refuge, leading to an eastward expansion of the Cerrado (Carnaval & Moritz 2008). Thus, it is possible that in one of these times and benefited by the increasing of non-forested areas in southeastern Brazil, D. brazili expanded its distribution range. Natural history: Drymoluber brazili inhabits open areas (Rodrigues 1996; França & Araújo 2006; França et al. 2008), and seems to be absent from altered habitats (França & Araújo 2006). Moreira et al. (2009) recorded the species inhabiting termite mounds, a common behavior in Cerrado snakes. Drymoluber brazili has diurnal activity, and is probably terrestrial (França & Araújo 2006; França et al. 2008). Gomes (1918), however, observed one of the specimens from the type series climbing in arboreal structures in captivity. Amaral (1978) and Pavan & Dixo (2004) cited D. brazili as arboriculous, but they did not state whether their data were obtained from previous published references or represent field observations.

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FIGURE 21. Hemipenis of Drymoluber brazili. IBSP 33668, Goiânia, Goiás, Brazil, A) Sulcate face; B) Asulcate face. MCNR 1736, Usina Hidrelétrica de Irapé, Minas Gerais, Brazil, C) Sulcate face; D) Asulcate face. Scale bar = 1 cm. Photos: Henrique C. Costa.

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There is a report of an unidentified Gymnophthalmidae as prey of Drymoluber brazili (França et al. 2008). Additionally, Pavan & Dixo (2004) cited anurans as the main prey of this snake, probably based on uncited literature references, since they collected only one specimen of D. brazili. França & Araújo (2006) recorded less than five eggs per clutch of Drymoluber brazili. One specimen examined in the present study (MZUFV 780) has four eggs. Gomes (1918) recorded some observations of the defensive behavior of Drymoluber brazili: “When handled, the specimens I had examined did not try to bite; however, when bothered by repeated light hits on the dorsum, they assumed a strike posture similar to D. bifossatus and other species of close related genera (Coluber, Spilotes, Herpetodryas), also rapidly shaking the tail.” (translation by the authors). Brodie III & Brodie Jr. (2004) suggested that young cross-banded specimens might mimic coralsnakes (Elapidae). The long tail of Drymoluber brazili (mean 44.2% of the SVL in males and 45.7% in females), with a tail beakage ratio of 28.9% (considering only those specimens where the breakage certainly occurred before the collection) is ample evidence of pseudoautotomy (Slowinski & Savage 1995), as in D. dichrous. Although morphometric data did not show the presence of sexual size dimorphism in D. brazili, examined males has on average longer SVL than females (mean 870 mm versus mean 856 mm). Etymology: The specific name brazili honors Vital Brazil Mineiro da Campanha, well known as the discoverer of the species-specificity of antivenoms, founder of the Instituto Butantan and its director when Gomes (1918) described the species. Vital Brazil developed a system by which the Butantan sent antivenom kits to farmers and ranchers in exchange for live snakes that were freely shipped to the institute by the railroad lines (Adler 2007). This system provided in a few years a snake collection of thousand of specimens to Butantan including the type series of Drymoluber brazili. One disadvantage was the fact that the specific locality of collected specimens could not be certified, and they were labeled as being from the railway stations from where they were sent to the institute (see information about the holotype and paratypes above).

Drymoluber apurimacensis Lehr, Carrillo & Hocking, 2004 Drymoluber apurimacensis Lehr, Carrillo & Hocking, 2004. Copeia, 2004, p. 47.

Holotype: Museu de Historia Natural Universidad Nacional Mayor de San Marcos, MHNSM 20672, juvenile female, SVL 206 mm, TL 73 mm, collected by P. Hocking on 15 January 2001, in Abancay (-13.64, -72.88), Abancay Province, Apurimác Department, Peru, 2500 m above sea level (Lehr et al. 2004). Specimen examined by photographs (Fig. 22). Paratypes: Field Museum of Natural History, FMNH 81542, male, SVL 460 mm, TL 192 mm, collected by C. Kalinowski in September 1953 in Hacienda Palmira (-13.65, -73.38), Huancarama district, Andahuaylas Province, Apurimác Department, Peru, 3300 m above sea level (specimen examined by the senior author; Fig. 23A); Museum für Tierkunde, Dresden, MTKD 44669, adult female, SVL 653 mm, TL 253 mm, killed by a local farmer in May 2002, in Cconoc (-13.06, -72.64), Abancay Province, Apurimác Department, Peru, 1925 m above sea level (Lehr et al. 2004) (specimen examined by photographs; Fig. 23B); MHNSM 18647, female, SVL 670 mm, TL 274 mm (Lehr et al. 2004; not examined), MTKD 45192, male, SVL 480 mm, TL 165 mm (Lehr et al. 2004; examined by photographs; Fig. 23C), MTKD 45193, female, without head, TL 216 mm (Lehr et al. 2004; examined by photographs; Fig. 23D), all collected by an inhabitant of Cconoc, Abancay Province, Apurimác Department, Peru, 1925 m above sea level (Lehr et al. 2004). Diagnosis: Drymoluber apurimacensis is distinguished from D. brazili and D. dichrous by the following combination of characters: a) 13-13-13 dorsal scale rows without apical pits; b) 158–164 ventrals in males, 166– 182 in females; c) 84–93 subcaudals in males, 87–91 in females; d) 14–16 maxillary teeth. Comparisons: Drymoluber brazili has 17-17-15 dorsal scales rows, and D. dichrous has 15-15-15. Apical pits are present in D. brazili and D. dichrous. Drymoluber brazili has 182–200 ventrals in males and 185–202 in females, 109–127 subcaudals in males and 109–126 in females. Drymoluber dichrous is not distinguishable from D. apurimacensis based on ventrals and subcaudals counts, having 157–173 ventrals in males and 160–180 in females, 87–110 subcaudals in males and 86–109 in females. Drymoluber brazili has 19–15 maxillary teeth, and D. dichrous has 19–16.

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FIGURE 22. Holotype of Drymoluber apurimacensis Lehr, Carrillo & Hocking, 2004 (MHNSM 20672). Photo: Claudia P. Torres Gastello.

The only known juvenile of Drymoluber apurimacensis (holotype) has dark crossbands 1–2 scales wide, and pale interspaces 2–3 scales wide. Juveniles of D. brazili have dark crossbands 2–6 scales wide (mean 3.6) and pale interspaces 0.5–5 scales wide (mean 1.6). Juveniles of D. dichrous have dark crossbands 1.5–7 scales wide (mean 3.6) and pale interspaces 0.5–2.5 scales wide (mean 0.8). The hemipenis of Drymoluber apurimacensis has more calyces than those of D. brazili, smaller spinulate flounces, and no spines in the lobular region. The walls of the sulcus spermaticus tend to be more ornamented, at least in the lobular region, with small jagged papillae. The spines on the asulcate face, mainly the most proximal, are larger than those in some specimens of D. brazili. The hemipenial morphology of D. apurimacensis and D. dichrous is similiar, making inaccurate the differentiation of these species by hemipenial characters. According to the description by Lehr et al. (2004), Drymoluber apurimacensis has dorsal scales with two apical pits. However, a reexamination of the holotype and four paratypes did not reveal apical pits (H.C. Costa pers. obs.; M. Auer, pers. comm.; C.P. Torres-Castello, pers. comm.). Lehr et al. (2004) also suggested that D. apurimacensis could de distinguished from its congeners based on the temporal formula of 1+2 (1+1/1 in the present study), while D. brazili and D. dichrous have temporal formulae of 2+2 (1/1+1/1 in the present study). As previously discussed, marked polymorphism of temporal plates characterizes Drymoluber, and the temporal formula 1+1/1 also occurs in D. dichrous and D. brazili, although the 1/1+1/1 formula is more frequent.

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Coloration of preserved adults: Three of the four paratypes examined (MTKD 44669, 45192, 45193) have the dorsum covered with old scales of olivre-brown color. The inner layer of scales is light bluish-gray. The labials, gular region and the venter are cream colored. The venter becomes bluish-gray with some cream marks on the posterior 2/3 of the body. Subcaudals are bluish-gray and cream colored.

FIGURE 23. Four of the five paratypes of Drymoluber apurimacensis Lehr, Carrillo & Hocking, 2004. A) FMNH 81542; B) MTKD 44669; C) MTKD 45192; D) MTKD 45193. Photos: Henrique C. Costa (A) and Barbara Bastian (B–D).

The coloration of FMNH 81542 is darker; its dorsum is dark-gray, and the venter is cream on the anterior third, becoming gray with cream marks posteriorly. The supralabials are cream with dark upper and lateral edges (the last four scales are almost completely black), and the gular region is cream. There is no information about the coloration of living adult specimens of D. apurimacensis. Coloration of preserved juveniles: The only known specimen of D. apurimacensis with juvenile coloration is the holotype (MHNSM 20672). Its dorsum has more than 40 dark crossbands of uniform size (about 1–2 scales wide), which become less visible in the posterior third of the body. The light interspaces are 2–3 scales wide. The gular region and venter are cream to light-gray color. The dark crossbands reach the lateral edges of ventrals. The head has the same color of the juveniles of D. dichrous and D. brazili with a transversal white stripe crossing the parietals. The supralabials are pale colored, with slightly darkened lateral edges. Coloration of juveniles in life: The holotype had black crossbands, and the interspaces are brown. The venter was light-gray and the head brown with black marks (Lehr et al. 2004). Hemipenial morphology (n=1) (Fig. 24): Hemipenis single, subcylindrical, not capitate. Sulcus spermaticus single and centrolinear. Lobe with just less than half of the hemipenis length, with papillate calyces (5–10 triangular papillae per calyx). Proximally, the calyces are gradually replaced by spinulate flounces and spines. Body covered by spines arranged in more or less transverse rows (about 60–70 spines in total). Walls of the sulcus spermaticus ornamented at least in the lobular region by jagged papillae and some spinules. These walls are also bordered on both sides by a longitudinal row of eight spines that tend to increase in size toward the proximal region. A hook is present at the end of each of row bordering the sulcus; the right hook is located more proximally

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than the left. Neither spines nor spinules occur between the left hook and the sulcus spermaticus. There are two lateral spines larger than the hooks left of the sulcus. The asulcate face of the hemipenis is formed by spines arranged in about seven more or less transverse rows (counted from proximal to distal region), with largest spines in the medial rows. The base of the hemipenis is smooth, with some grooves and several sparse spinules. Variation: When available, data of the paratype MHNSM 18647 given by Lehr et al. (2004) were used. Largest male has SVL 643 mm, TL 253 mm (MTKD 44669); largest female has SVL 670 mm, TL 274 mm (MHNSM 18647; not examined). The tail is 34.4–41.7% of the SVL in males (mean 38.5%; SD=3.75; n=3) and 35.4–40.9% (n=2) in females. For variation in meristic characters, see Table 5. Geographic distribution: Drymoluber apurimacensis is known from three localities between 1920 and 3300 meters above sea level south of the Apurimác and Pampas Rivers, department of Apurimác, in the Serranía Esteparia ecoregion. The area is characterized by matorral seco and matorral sub-húmedo vegetation (Lehr et al. 2004). Drymoluber apurimacensis probably is endemic to the deep valley of the Apurímac River, which seems to be a significant geographical barrier to the North-south distribution of some Andean taxa (Lehr et al. 2004). Amphibians of the genus Bryophryne (Strabomantidae), the bird Synallaxis courseni (Furnariidae) and the plant Solanum anomalostemon (Solanaceae) are some species endemic to the Apurímac region (Lehr & Catenazzi 2008; BirdLife International 2009; Knapp & Nee 2009). Natural history: The holotype was collected under a rock at midday, and the paratype MTKD 45193 had a lizard Euspondylus sp. (Gymnophthalmidae) in its stomach (Lehr et al. 2004). Drymoluber apurimacensis is sympatric with the snakes Dipsas peruana, Leptotyphlops sp., Oxyrhopus melanogenys and Tachymenis peruviana (Lehr et al. 2004).

FIGURE 24. Hemipenis of Drymoluber apurimacensis, FMNH 81542, Hacienda Palmira, Huancarama district, Andahuaylas Province, Apurimác Department, Peru. A) Sulcate face; B) Asulcate face. Scale bar = 0.5 cm. Photos: Henrique C. Costa.

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Etymology: The specific name apurimacensis means “from Apurimác”. Apurimác, the name of the region where the species is known to occur is a Quechuan (an indiginous language from the Andes) word meaning “the mountain spirit that speaks” (Lehr et al. 2004).

Key to the species of Drymoluber Amaral, 1930 1A. 1B. 2A. 2B.

Seventeen (17) midbody scale rows, with reduction to fifteen (15) rows anterior to the cloacal shield; 182–202 ventrals; 109– 127 subcaudals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. brazili Thirteen (13) or fifteen (15) midbody scale rows, without posterior reduction; 157–182 ventrals; 84–110 subcaudals . . . . . . . 2 Thirteen (13) dorsal scale rows at midbody, without apical pits; 158–182 ventrals; 84–93 subcaudals; 14–16 maxillary teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. apurimacensis Fifteen (15) dorsal scale rows at midbody, with two apical pits; 157–180 ventrals; 86–110 subcaudals; 19–26 maxillary teeth.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. dichrous

Acknowledgements We are indebt to several persons and institutions: the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) and the Smithsonian Institution for schorlarships to the senior author; Guarino R. Colli, Marcela A. Brasil and Mariana C.M. Viana (CHUNB), Paulo C.A. Garcia and Patrícia S. Santos (CHUFMG), Jacques H.C. Delabie (CZGB), Francisco Luís Franco and Valdir José Germano (IBSP), Maria Rita S. Pires and Bruna Imai (LZVUFOP), Luisa Sarmento and Fabiano Lanschi (MBML), Gláucia Maria F. Pontes (MCP), Luciana B. Nascimento (MCN), Hussam Zaher and Carolina Mello (MZUSP), Antônio J.S. Argôlo (MZUESC), Ronaldo Fernandes and Marcelo Gomes (MNRJ), Ana Lúcia C. Prudente, João Fabrício M. Sarmento, Alessandra Travassos and Paula C.R. Almeida (MPEG), Paulo S. Bernarde (UFACF), Diva Maria Borges-Nojosa and Daniel C. Lima (UFC), Marcos André Carvalho (UFMT), Darrel Frost and David Kizirian (AMNH), Alan Resetar and Kathleen M. Kelly (FMNH), Ronald Heyer, Kevin de Queiroz, Roy McDiarmid, George Zug, Addison Wynn, Kenneth Tighe, Jeremy F. Jacobs and Robert Wilson (USNM) for allowing the access to specimens under their care; Edgar Lehr, Markus Auer and Barbara Bastian (MTD), Mark-Oliver Rödel and Christoph Kucharzewski (MNB), Pier Cacciali (MNHNP), Cesar Aguilar and Claudia P. Torres Gastello (MHNSM), Colin McCarthy (BMNH), Stephen P. Rogers (CMNH) for data and pictures of specimens under their care; Giovanna G. Montingelli, for pictures of the holotype of Spilotes piceus; Daniel Loebmann, Eduardo Santos, Laurie J. Vitt, Maël Dewynter, Marco Antônio de Freitas, Paula H. Valdujo, Paulo S. Bernarde, Pedro Henrique Bernardo, Rafael de Fraga and William Quatman, for kindly allowing the use of their pictures in this work; Valdir José Germano, for teaching the technique of hemipenes preparation; Peter Uetz, for copies of the original descriptions of Herpetodryas dichroa, H. occipitalis and Spilotes piceus; Daniel Burnier and Peter Uetz for the translation of the descrpition of H. dichroa to English; Addison Wynn, Alan Resetar, Burger Family, Débora C. Rodrigues, Fabiano Godoy, Fiona Wilkinson, Jeremy F. Jacobs, Kathleen M. Kelly, Kenneth Tighe, Larry Matheson, Marcelo J. Sturaro, Márcio M. Morais Jr., Pedro L.V. Peloso, Pedro Nunes, Robert Wilson, Ronald and Miriam Heyer and Roy McDiarmid, for their contribution before and during the stay of the senior author in the USA; Francisco L. Franco, Jorge A. Dergam, Renato S. Bérnils, Rômulo Ribon, and two anonymous reviewers, for valuable corrections and suggestions in early drafts of the manuscript; Roy McDiarmid for English review through the Society for the Study of Amphisbians and Reptiles (SSAR) editorial assistance program; Hussam Zaher for editorship.

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Appendix I Collections from which specimens of Drymoluber were personally examined for this study. BRAZIL: Coleção Herpetológica da Universidade de Brasília (CHUNB), Brasília, DF; Coleção Herpetológica da Universidade Federal de Minas Gerais (CHUFMG), Belo Horizonte, MG; Coleção Zoológica Gregório Bondar (CZGB), Ilhéus, BA; Instituto Butantan (IBSP), São Paulo, SP; Laboratório de Zoologia de Vertebrados, Universidade Federal de Ouro Preto (LZV), Ouro Preto, MG; Museu de Biologia Mello Leitão (MBML), Santa Teresa, ES; Museu de Ciências e Tecnologia, PUC-RS (MCP), Porto Alegre, RS; Museu de Ciências Naturais, PUC-MG (MCNR), Belo Horizonte, MG; Museu de Zoologia da Universidade de São Paulo (MZUSP), São Paulo, SP; Museu de Zoologia da Universidade Estadual de Santa Cruz (MZUESC), Ilhéus, BA; Museu de Zoologia João Moojen, Universidade Federal de Viçosa (MZUFV), Viçosa, MG; Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ), Rio de Janeiro, RJ; Museu Paraense Emílio Goeldi (MPEG), Belém, PA; Universidade Federal do Acre, Campus Floresta (UFACF), Cruzeiro do Sul, AC; Universidade Federal do Ceará (UFC), Fortaleza, CE; Universidade Federal do Mato Grosso (UFMT), Cuiabá, MT; UNITED STATES OF AMERICA: American Museum of Natural History (AMNH), New York, NY; Field Museum of Natural History (FMNH), Chicago, IL; National Museum of Natural History, Smithsonian Institution (USNM), Washington, DC. Collections from which photographs of specimens of Drymoluber were examined for this study. GERMANY: Museum für Naturkunde Berlin (ZMB), Berlin; Museum für Tierkunde (MTKD), Dresden; PARAGUAY: Museo Nacional de Historia Natural Del Paraguay (MNHNP), Asuncíon; PERU: Museo de Historia Natural Universidad Nacional Mayor de San Marcos (MHNSM), Lima; UNITED KINGDOM: National Museum of Natural History (BMNH), London; UNITED STATES OF AMERICA: Academy of Natural Sciences of Philadelphia (ANSP), Philadelphia, PA.

Appendix II Material examined. Toponyms are cited in descending order (COUNTRY, STATE / PROVINCE / DEPARTMENT, City / Municipality, Locality), with geographic coordinates inside brackets; * indicates specimens examined only by photographs; † indicates specimens which hemipenis was also examined. Geographic coordinates expressed in decimal degrees as “latitude, longitude” (e.g. 2.60, -72.60 mean 2.60° N, 72.60° W). Drymoluber apurimacensis PERU: APURÍMAC: Abancay, Abancay (MHNSM 20672* [Holotype]) [-13.64, -72.88], Cconoc (MTKD 44669*, 45192*, 45193* [Paratypes]) [-13.06, -72.64]; Andahuaylas, Huancarama, Hacienda Palmira (FMNH 81542 † [Paratype]) [-13.65, -73.38]. Drymoluber brazili BRASIL: Without precise locality (IBSP 574 [Paratype of Drymobius brazili]). BAHIA: Without precise locality

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(IBSP 2717); Gentio do Ouro, Santo Inácio (MZUSP 9596) [-11.10, -42.73]; Jacobina, Caatinga do Moura (MZUSP 7544) [-10.98, -40.77]; João Dourado, Gruta dos Brejões (MZUESC 3815) [-11.00, -41.25]; Morro do Chapéu (MZUSP 7807) [-11.55, -41.16]. CEARÁ: Milagres (IBSP 76968) [-7.31, -38.95]. DISTRITO FEDERAL: Brasília (CHUNB 37345, CHUNB 3747, CHUNB 3748, CHUNB 6054) [-15.78, -47.93]. ESPÍRITO SANTO: Baixo Guandú (IBSP 8312, IBSP 8836 †) [-19.52, -41.02]; Colatina (IBSP 37413) [-19.54, -40.63]. GOIÁS: Alto Paraíso de Goiás (CHUNB 3839) [-14.13, -47.51]; Goiânia (IBSP 33668 †) [-16.68, -49.25]; Mineiros, Parque Nacional das Emas (IBSP 62682) [-18.08, -52.92]; São Domingos, Parque Estadual Terra Ronca (IBSP 62650) [-13.65, -46.35]; Uruaçu, Cana Brava (IBSP 26716, IBSP 9144) [-14.53, -49.14]; Serra da Mesa (ponto 2) (MZUSP 11021) [-13.90, -48.31]. MATO GROSSO: Chapada dos Guimarães (UFMT 6970 †, UFMT 728) [-15.45, -55.74]. MATO GROSSO DO SUL: Campo Grande (IBSP 10465 †, IBSP 31692) [-20.44, -54.65]; Miranda (IBSP 1845) [-20.24, -56.38]; Ponta Porã (IBSP 25379 †) [-22.54, -55.73]; Terenos (IBSP 10018, IBSP 9829) [-20.44, -54.86]; Três Lagoas (IBSP 51700) [-20.75, -51.68], Jupiá (IBSP 27199) [-20.75, 51.68]. MINAS GERAIS: Aimorés (MCNR 1538) [-19.50, -41.06]; Alvarenga (CHUFMG 2843) [-19.41, -41.72]; Frutal (IBSP 52290) [-20.03, -48.94]; Grão Mogol (MZUSP 7988) [-16.71, -42.59]; Nova Ponte (MZUFV 780) [-19.14, 47.68]; Sabará (IBSP 40475) [-19.89, -43.81]; Santana do Riacho (CHUFMG 417) [-19.11, -43.68], Serra do Cipó (MZUSP 7692) [-19.23, -43.55]; Uberaba (IBSP 1707); Uberaba, Engenheiro Lisboa (IBSP 696 [Holotype of Drymobius brazili]) [-19.80, -47.60]; UHE Irapé (MCNR 1736 †) [-16.74, -42.58]. PARAÍBA: Teixeira, Serra do Teixeira (MZUSP 7562) [-7.20, -37.25]. SÃO PAULO: Andradina (IBSP 40779) [-20.90, -51.38]; Araçatuba (IBSP 26688) [21.21, -50.43]; Araraquara (IBSP 23693, IBSP 33120, IBSP 8103) [-21.79, -48.18]; Barretos (IBSP 7675) [-20.56, 48.57]; Birigui (IBSP 21928, IBSP 22415) [-21.29, -50.34]; Boa Esperança do Sul, Estação Ferroviária Java (IBSP 741 [Paratype of Drymobius brazili]) [-21.99, -48.39]; Castilho (IBSP 24422) [-20.87, -51.49]; Coroados (IBSP 17019) [21.35, -50.28]; Dourado (IBSP 18309) [-22.10, -48.32]; Dracena (IBSP 25187) [-21.48, -51.53]; Guaraçaí (IBSP 32897) [-21.03, -51.21]; Guatapará, Estação Ferroviária de Monteiros (IBSP 12692) [-21.50, -48.04]; Itápolis (IBSP 18053) [21.60, -48.81]; Moreira Melo (IBSP 16595) [not found]; Novo Horizonte, Estação Ferroviária Porto Ferrão (IBSP 22502, IBSP 22515) [-21.47, -49.22]; Oswaldo Cruz (IBSP 32624, IBSP 32657) [-21.80, -50.89]; Pedregulho (IBSP 1286 [Paratype of Drymobius brazili]) [-20.26, -47.48]; Penápolis (IBSP 1844 [Holotype of Drymobius rubriceps]) [21.42, -50.08]; Ribeirão Bonito (IBSP 46996) [-21.18, -47.81], Estação Ferroviária Sampaio Vidal (IBSP 573 [Paratype of Drymobius brazili]) [-22.07, -48.18]; Ribeirão Preto (IBSP 18992) [-21.18, -47.81]; Santa Lúcia, Fazenda Santa Izabel (IBSP 34369) [-21.69, -48.08]; São Carlos, Estação Ferroviária Santa Eudóxia (IBSP 383 [Paratype of Drymobius brazili]) [-21.77, -47.78]; São Joaquim da Barra, Estação Ferroviária Jussara (IBSP 16231) [-20.58, -47.86]; São José do Rio Preto, Estação Ferroviária Rio Preto (IBSP 10188, IBSP 8925) [-20.82, -49.38]; São Simão, Fazenda Vale da Saúde (IBSP 33660) [-21.48, -47.55]; Turiba do Sul (IBSP 10538, IBSP 16499) [-23.75, -49.28]; Urânia (IBSP 17224) [-20.25, -50.64]; Valparaíso (IBSP 29221) [-21.23, -50.87]. TOCANTINS: UHE Luís Eduado Magalhães (MZUSP 14298) [-9.75, -48.40]; UHE Peixe Angical (MZUSP 15506, MZUSP 15507) [-12.40, -48.25]. PARAGUAI: CANINDEYÚ: Reserva Natural del Bosque Mbaracayú (MNHNP 11025*) [-24.14, -55.32]. Drymoluber dichrous BOLIVIA: BENI: Riberalta (AMNH 22491) [-10.98, -66.10]; Rurrenabaque (AMNH 119926) [-14.44, -67.53]; Vaca Diez, Tumi Chucua (USNM 280382) [-11.15, -66.17]. BRASIL: Without precise locality (ZMB 1661* [Syntype of Herpetodryas dichroa], ZMB 1662* [Syntype of Herpetodryas dichroa]). Without precise locality, boundary with Peru (AMNH 52191); ACRE: Porto Walter (MZUSP 7377, MZUSP 7378) [-9.33, -70.42], Rio Juruá (MPEG 20384 †) [-8.27, -72.74]; Rio Branco (IBSP 69567) [-9.97, -67.80]; Santa Rosa do Purus, Alto Purus (MZUSP 2497) [-9.33, -70.42]; Tarauacá, Reserva Extrativista Riozinho da Liberdade (UFACF 1333, UFACF 663, UFACF 720, UFACF 741, UFACF 801) [-8.00, -72.00]. AMAPÁ: Macapá, Zebrão (MNRJ 9240) [0.04, -51.06]; Serra do Navio (IBSP 25411) [0.90, -52.00]; Tartarugalzinho, Reserva DNERu, Rio Tracajatuba (MPEG 421) [1.00, -51.00]; Rio Maracá (MZUSP 7681) [-0.43, 51.43]. AMAZONAS: Without precise locality (IBSP 7706); Arajatuba, Rio Negro (IBSP 2198) [-3.10, -60.49]; Barcelos, Rio Negro MZUSP 5467 [-0.97, -62.93]; Benjamin Constant, Próximo a igarapé (MPEG 18240 †) [-4.47, -70.03]; Madeirera SCHEFFER, Rio Ituxi (MPEG 20330, MPEG 20331) [-8.20, -65.42]; Presidente Figueiredo, UHE Balbina (IBSP 51703, IBSP 52003, IBSP 52140 †, MPEG 17437, MPEG 17443, MPEG 17446, MPEG 17481, MPEG 17494, MPEG 17566, MZUSP 9644, MZUSP 9645) [-1.54, -59.84]; Reserva INPA-WWF (MZUSP 7609) [-2.42, -59.72]; Serra do Tapirapecó (MZUSP 14284, MZUSP 14285, MZUSP 14286) [1.20, -64.80]. BAHIA: Without precise locality (MZUESC 5801); Arataca (CZGB 396) [-15.26, -39.41], Fazenda Alto Rocha (MZUESC 1528) [-15.15, -39.30]; Barra do Choça (CZGB 7981, CZGB 7396) [-14.86, -40.56], Fazenda Cangussu (MZUESC 2540) [-14.96, -40.65]; Barra do Rocha, Fazenda Pedra Preta (MZUESC 1993) [-14.06, -39.60]; Barro Preto (CZGB 822) [-14.77, -39.40]; Buerarema (CZGB 339, CZGB 4441) [-14.95, -39.30]; Caatiba (CZGB 7059) [-14.98, -40.41]; Cairu, Fazenda Ilha do Barro (MZUESC 636) [-13.55, -39.06]; Camacan (CZGB 846) [-15.42, -39.50], Fazenda Uraiçu (MZUESC 5162) [-15.39, 39.56]; Camamu (CZGB 5180) [-13.95, -39.10]; Dario Meira (CZGB 3164, CZGB 3342) [-14.44, -39.91]; Ibirataia (CZGB 7397) [-14.07, -39.64]; Ilhéus (CZGB 1222, CZGB 1269, CZGB 3176, CZGB 666 †) [-14.78, -39.11; Itabuna (CZGB 324) [-14.79, -39.28]; Itacaré (CZGB 2101) [-14.28, -39.00]; Itagi (MZUESC 5697 †) [-14.07, -39.95; Itagiba (CZGB 5430) [-14.28, -39.84; Ituberá (CZGB 5609, CZGB 6825, CZGB 6826) [-13.73, -39.15]; Mutuípe (CZGB 7717 TAXONOMIC REVISION OF DRYMOLUBER

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†) [-13.23, -39.51]; Nova Ibiá (CZGB 6809, CZGB 7385) [-13.83, -39.59]; Poções, Fazenda Inveja (MZUESC 1383) [14.63, -40.27]; Una (CZGB 331) [-15.27, -39.07], Fazenda Araraúna (MZUESC 3739) [-15.27, -39.07]. CEARÁ: Maranguape, Serra de Maranguape (UFC 2101, UFC 2201, UFC 2204, UFC 2205, UFC 2211, UFC 2220, UFC 2712, UFC 2730) [-3.90, -38.72], Trilha do Pico da Rajada, Serra de Maranguape (UFC 2081, UFC 2083 †, UFC 2104, UFC 2128) [-3.90, -38.72]; Pacatuba, Serra do Aratanha (UFC 2233) [-3.97, -38.62]; Pacoti, Serra do Baturité (UFC 2221) [4.23, -38.92]; Ubajara, Planalto da Ibiapaba, Fazenda Buriti-INCRA (UFC 2177 †) [-3.84, -40.90]. ESPÍRITO SANTO: Aracruz (MBML 483) [-19.82, -40.27]; Linhares (CZGB 2121 †, MNRJ 10661, MNRJ 10663) [-19.39, -40.07]; Vitória (MNRJ 4858) [-20.32, -40.34]. MARANHÃO: BR 316, 25 Km distante do Rio Gurupi, Nova Vida (MPEG 11099, MPEG 11155, MPEG 12106, MPEG 12228 †, MPEG 12229, MPEG 12230, MPEG 12257, MPEG 14466, MPEG 15321, MPEG 16207) [-2.00, -45.99]; Buriticupu (MPEG 17713) [-4.35, -46.40]; Santa Luzia do Paruá, BR 316 (MPEG 10832) [-2.63, -45.77]. MATO GROSSO: Alta Floresta (IBSP 41469) [-9.88, -56.09]; Aripuanã (UFMT 5184, UFMT 5474) [10.15, -59.45], Ilha do Salto Dardanelos (UFMT 2772) [-10.15, -59.45]; Juruena (MZUSP 11303) [-10.32, -58.36]; Rio Teles Pires, Paraná do Cristóvão (MZUSP 10983) [-7.55, -57.97]; Vale do São Domingos (UFMT 1997 †, UFMT 919) [-15.29, -59.06]; Vila Rica (MZUSP 11443, MZUSP 11444) [-10.01, -51.12]. MINAS GERAIS: Almenara, Fazenda Limoeiro (MZUFV 1208 †) [-16.01, -40.50]; Caratinga, RPPN Feliciano Miguel Abdala (Estação Biológica de Caratinga) (CHUFMG 1397, CHUFMG 1398) [-19.71, -41.81]; Nova Ponte (MZUFV 779 †) [-19.14, -47.68]; Ouro Preto, Estação Ecológica do Tripuí (LZV-S 373) [-20.40, -43.58]; Parque Estadual do Rio Doce (MNRJ 9297, MZUFV 1422) [-19.43, -42.45]. PARÁ: Almerim, Reserva Itapioara (MPEG 20108) [-1.53, -52.56]; Belém (IBSP 3035, IBSP 3122 †, IBSP 3141, IBSP 5089) [-1.46, -48.50]; Benevides, Santa Bárbara (MPEG 2647) [-1.36, -48.25], Pratinha, Estrada de Genipauba, antiga estrada do Açucareiro (MPEG 8400, MPEG 8607) [-1.36, -48.25]; Canindé, Rio Gurupi (MZUSP 4237) [-2.55, -46.52]; Castanhal, Boa Vista (MPEG 2652, MPEG 2670, MPEG 4041) [-1.36, -47.99]; Conceição do Araguaia (IBSP 41522) [-8.26, -49.26]; Juruá, Rio Xingu (MZUSP 9351, MZUSP 9352) [-3.40, -51.88]; Marabá (CHUNB 30419, MPEG 23235) [-5.37, -49.12], Serra Norte, Carajás (MPEG 17233, MPEG 17235) [-5.85, 50.20], Serra Norte, Carajás, Área do Caldeirão (MPEG 16921) [-5.85, -50.20], Serra Norte, Carajás, Campo rupestre do N1 (MPEG 16551, MPEG 16982) [-5.85, -50.20], Serra Norte, Carajás, Próximo as margens do Igarapé Azul, Estrada N1, Caldeirão (MPEG 17057) [-5.85, -50.20], Serra Norte, Carajás, Área do Pojuca (MPEG 17085) [-5.85, 50.20]; Melgaço, FLONA Caxiuanã (MPEG 19750, MPEG 20044, MPEG 20146, MPEG 20298, MPEG 20800, MPEG 21833, MPEG 21834) [-1.80, -51.50]; Monte Cristo, Rio Tapajós (MZUSP 1264) [-4.07, -55.65]; Novo Progresso (CHUNB 35062) [-7.15, -55.38]; Ourilândia do Norte, Acampamento Onça (MPEG 20793) [-7.15, -51.00]; Paragominas, Fazenda Agroeste (MPEG 20000) [-3.00, -47.35]; Peixe-Boi (MPEG 1838) [-1.20, -47.32]; Projeto Igarapé Bahia, Carajás (MPEG 20762, MPEG 20765, MPEG 20766) [-6.05, -50.10]; Reserva Biologica Rio Trombetas, Igarape Jacare above Lago Jacaré (USNM 289091) [-1.19, -56.67]; Santarém (MCP 7617, MCP 7618, MCP 7906) [2.44, -54.71]; São Domingos do Capim, Km 16 da estrada do Acará (MPEG 10811, MPEG 11698, MPEG 15189, MPEG 8688) [-1.68, -47.77]; Serra de Kukoinhokren (MZUSP 10763, MZUSP 11488) [-7.77, -51.95]; Tucuruí (IBSP 46626) [-3.83, -49.69]. PERNAMBUCO: Cabo de Santo Agostinho, Mata do Cutio, Reserva Florestal do sistema Gursaú (MNRJ 17069) [-8.27, -35.06]. RONDÔNIA: Alto Paraíso (MZUSP 8494) [-9.62, -63.45]; Jaru, Santa Cruz da Serra (MZUSP 8500) [-10.67, -62.57]; Ministro Andreazza, Nova Brasília (MZUSP 8747) [-11.15, -61.57]; Monte Negro (IBSP 67619) [-10.24, -63.29]; Porto Velho, UHE Samuel (IBSP 52828, IBSP 53256) [-8.86, -63.43], Margem esquerda do Rio Jamarí. Área de inundação da UHE Samuel (MPEG 17779, MPEG 17808, MPEG 17820, MPEG 17829, MPEG 17841, MPEG 17849) [-8.86, -63.43]; Margem do Rio Jamarí. Área de inundação da UHE Samuel (MPEG 17897, MPEG 17915, MPEG 17932, MPEG 17995, MPEG 17996, MPEG 18759) [-8.86, -63.43]; Vilhena (CHUNB 12791) [12.74, -60.15]. RORAIMA: Amajari, Ilha de Maracá (MZUSP 8806) [3.42, -61.67]; BR 210, 4Km do Rio Ajarani (MPEG 19007) [1.92, -61.17]; Caracaraí, Missão Catrimani (MZUSP 10298, MZUSP 10422, MZUSP 10423, MZUSP 10424) [1.68, -62.28]; Mucajaí, Apiaú (MZUSP 9777) [2.63, -61.25]; Cachoeira do Cujubim, Rio Catrimani (MZUSP 6388, MZUSP 6389, MZUSP 6971) [1.75, -62.28]. COLOMBIA: AMAZONAS: Leticia (AMNH 91812) [-4.17, -69.95]. GUAVIARE: Caño Agua Bonita (FMNH 75688) [2.60, -72.60]. META: Bellavista, acima de Villavicencio (MZUSP 6098) [4.25, -73.67]; Villavicencio (IBSP 6208, IBSP 7221) [4.16, -73.64], Finca El Buque (MZUSP 8077) [4.15, 73.62]. ECUADOR: Without precise locality, Napo or Maranon (ANSP 3920* [Holotype of Spilotes piceus]). CHIMBORAZO: Riobamba (AMNH 23248) [-1.67, -78.63]. MORONA-SANTIAGO: Arapicos, Río Llhushin (USNM 204133) [-1.85, -77.95]; Macas (USNM 65475; AMNH 28830) [-2.32, -78.12]; Sucua (USNM 283952) [-2.47, -78.17]; Chiguaza + Macuma (USNM 204132) [-2.10, -77.87]; Riobamba, Chamala, Normandia (in the vicinities of Macas, according to Savage, 1960. Misc. Pub. Mus. Zool. Univ. Mich. 112) (AMNH 35927) [-2.31, -78.12]. NAPO: Cabeceras del Rio Arajuno, tributary of Rio Napo (USNM 204126) [-1.40, -77.88]; Loreto (USNM 204125) [-0.67, -77.33]. PASTAZA: Andoas, Rio Pastaza (AMNH 49074) [-2.57, -76.63]; Rio Conambo (FMNH 206028) [-2.12, -76.05]; Río Liguino (USNM 204127) [-1.48, -77.37]; Cabeceras Del Río Bobonaza (USNM 204128) [-1.47, -77.88]; Río Bufeo, tributary of Río Bobonaza (USNM 204129) [-2.33, -76.67]; Río Conambo, near mouth of Río Romarizo (USNM 204130) [-1.87, -76.78]; Paracachi, Rio Curaray (USNM 204131) [-1.60, -76.35. GUYANA: CUYUNI-MAZARUNI: Kartabo (AMNH 18160) [-6.35, -58.68]; East bank of Waruma River, Camp 3 (USNM 549327) [5.34, -60.77]. EAST BERBICE: ca. 18 mi. SW Kwakwani, ca. 2 mi. downriver from confluence of Berbice River and Kurudini River, Berbice River camp

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(USNM 566261) [5.09, -58.24]. PERU: AMAZONAS: Kayamas, Río Cenepa (USNM 560425) [-4.47, -78.17]; San Antonio, Río Cenepa (USNM 316601) [4.50, -78.17]; Huampami, across the Rio Cenepa from, Yusa Patagkamu (USNM 316602) [-4.47, -78.17]. HUÁNUCO: Tingo Maria (USNM 193808, USNM 193813) [-9.30, -75.98], Universidad Agraria de la Selva, Rio Huallaga (USNM 193807, USNM 193812) [-9.30, -75.98]. JUNIN: Perene (AMNH 23354, AMNH 23374) [-10.95, -75.23]; Tarma, Chanchamayo (FMNH 45593) [-11.40, -75.70]. LORETO: Contamana, Suhuaya Isla (AMNH 52295) [-7.34, -75.06]; Iquitos (AMNH 52318, AMNH 52644, USNM 197267) [-3.76, -73.26], Río Itaya (AMNH 54100, AMNH 54168, AMNH 54260, AMNH 54392, AMNH 54479, AMNH 54507, AMNH 54518, AMNH 54541, AMNH 54672, AMNH 54699, AMNH 54803, AMNH 54919, AMNH 54930 †, AMNH 55193, AMNH 55276) [-3.77, -73.25]; Nauta, Rio Samiyia, Santa Elena (FMNH 109807) [-4.53, -73.45]; Pampa Hermosa, Río Cushabatay, Río Ucayali (AMNH 55359) [-7.21, -75.26], Río Cushabatay (AMNH 55963) [-7.23, -75.30]; Pebas (BMNH 1946.1.14.61, formerly 1867.9.17.28* [Holotype of Herpetodryas occipitalis]) [-3.18, -71.78]; Requena, Monte Carmelo (AMNH 55609, AMNH 55628, AMNH 55635) [-4.99, -73.96]; Yagua, Indian Village, Río Loretoyacu (AMNH 114706) [-3.82, 70.43]; Yurimagua (FMNH 11259 †) [-5.90, -76.11]; Estirón, Rio Ampiyacu (MZUSP 4390, MZUSP 4391) [-3.12, 71.92]; Río Samiria and Parinari Cañon (AMNH 57258) [-4.70, -74.30]; Upper Ucayali River (AMNH 71115) [-4.50, 73.45]; Upper Amazon (FMNH 11177) [-6.34, -75.17]; Yarinacocha (FMNH 45592) [-5.52, -74.37]. MADRE DE DIOS: ca. 30 Km Puerto Maldonado Tambopata Reserve, Explorer's Inn (USNM 247501, USNM 247500, USNM 247694) [12.83, -69.28]; La Pampa (FMNH 40206 †) [Toponym not found]; Maldonado (AMNH 56148) [-12.60, -69.18]; West bank of Río Tambopata, Zona Reservada Tambopata-Candamo, Colpa de Guacamayo (USNM 332470) [-13.14, -69.61]. SURINAME: Without precise locality data (ZMB 2603* [syntype of Herpetodryas dichroa]). BROKOPONDO: Brownsberg, Brownsberg Nature Park, Mazaroni Plateau (AMNH 119431) [4.95, -55.17]. COMMEWIJNE: Plantation Ma Retraite (AMNH 130505) [5.74, -54.87]. MAROWIJNE: Moengo (USNM 64634) [5.62, -54.40]; Anapaike Village (MZUSP 4207) [3.33, -54.07]. SIPALIWINI: Temomairem, Sierra Tumuc Humac (AMNH 104610) [3.00, -55.38]. VENEZUELA: AMAZONAS: Cerro Ya-Pacana (USNM 83946) [3.75, -66.80]; Mount Duida region (AMNH 36610) [3.48, -65.58]; Middle camp, Mount Duida region (AMNH 36614) [3.38, -65.94]; Esmeralda, 56 km NW of Rio Cunucunuma, Belen (USNM 217185) [3.65, -65.77].

Appendix III Localities without material examined, but with literature records or personal observation. Toponyms are cited in descending order (COUNTRY, STATE / PROVINCE / DEPARTMENT, City / Municipality, Locality), with geographic coordinates and the source of record inside brackets. Drymoluber brazili BRASIL: MINAS GERAIS: Mariana [-20.38, -43.42; Silveira et al. 2010 “2011”]. GOIÁS: Iporá/Arenópolis, Pequena Central Hidrelétrica de Mosquetão [-16.35, -51.43; Silva-Jr. et al. 2007]. PIAUÍ: Parque Nacional da Serra das Confusões [-9.45, -43.86; Silva et al. 2007]; São Gonçalo do Gurguéia [-10.04, -45.29; Freitas et a. 2012]. Drymoluber dichrous BOLÍVIA: BENI: Parque Nacional y Territorio Indígena Isiboro Sécure [-16.38, -66.04; Embert 2007]; Reserva de la Biósfera Estación Biológica del Beni [-14.66, -66.33; Embert 2007]; Reserva de la Biósfera y Tierra Comunitaria de Origen Pilón Lajas [-15.00, -67.31; Embert 2007]. COCHABAMBA: Parque Nacional Carrasco [-17.50, -65.00; Embert 2007]; Santuario de Vida Silvestre Cavernas del Repechón [-17.38, -66.06; Embert 2007]; Villa Tunari [-16.97, -65.43; Lehr et al. 2004]. LA PAZ: Área Natural de Manejo Integrado Apolobamba [-15.00, -69.00; Embert 2007]; Parque Nacional y Área Natural de Manejo Integrado Cotapata [-16.17, -68.00; Embert 2007]; Parque Nacional y Área Natural de Manejo Integrado Madidi [-13.85, -68.57; Embert 2007]. SANTA CRUZ: Guarayos, Campamento Río San Martin [15.05, -61.93; Montero et al. 1995]; Parque Nacional Noel Kempff Mercado [-14.80, -60.64; Harvey 1998; Embert 2007] ; Parque Nacional y Área Natural de Manejo Integrado Amboró [-17.84, -64.08; Embert 2007]. PANDO: Reserva Nacional de Vida Silvestre Amazónica Manuripi [-11.69, -67.61; Embert 2007]. BRASIL: ALAGOAS: Maceió [-9.67, 35.72; Freire 2000]; Rio Largo [-9.50, -35.83; Freire 2000]. AMAZONAS: Manaus, Reserva Adolpho Ducke [-2.98, 59.93; Martins & Oliveira 1998]. CEARÁ: Missão Velha, Santo Antônio water spring [-7.41, -39.21; Veríssimo et al. 2012]. PARÁ: Agropecuária Treviso [-3.15, -54.83; Ávila-Pires et al. 2009]; Bragança, Parada Bom Jesus [-1.07, -46.79; Cunha & Nascimento 1978]; Colônia Nova, Km 264, BR 316 [-1.80, -46.48; Cunha & Nascimento 1978]; Curuçá, Vila Marauá [-0.75, -47.87; Cunha & Nascimento 1978]; Maracanã, Km 23 da estrada de Maracanã [-0.78, -47.46; Cunha & Nascimento 1978]; Ourém, Limão Grande [-1.55, -47.11; Cunha & Nascimento 1978]; Santo Antônio do Tauá, PA-140, estrada da Vigia [-1.11, -48.13; Cunha & Nascimento 1978]; Vigia, Santa Rosa, PA-140, estrada da Vigia [-0.97, -48.08; Cunha & Nascimento 1978]; Viseu, Bela Vista [-1.21, -46.19; Cunha & Nascimento 1978], Fazenda Real [-1.27, -46.22; Cunha & Nascimento 1978]. PARAÍBA: João Pessoa, Área de Preservação Permanente Mata do Buraquinho [-7.15, 34.87; Santana et al. 2008]. RIO DE JANEIRO: Campos dos Goytacazes [-21.84, -41.67; photograph of a juvenile specimen, taken by Carlos Henrique Oliveira Nogueira]. Rondônia: Espigão D’Oeste [-11.39, -60.74; Bernarde & Abe TAXONOMIC REVISION OF DRYMOLUBER

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2006]; Parque Estadual Guajará-Mirim [-10.32, -64.55; Avila-Pires et al. 2009]. COLÔMBIA: BOYACÁ: 8 Km SSE Corocito [5.53, -73.06; Lehr et al. 2004]. EQUADOR: AZUAY: San José Cuchipamba [-3.02, -78.75; Peracca 1897]. MORONA-SANTIAGO: Gualaquiza [-3.40, -78.58; Peracca 1897]. NAPO: Aliñahui [-1.07, -77.60; Lehr et al. 2004]. ORELLANA: Estacíon de Biodiversidad Tiputini [-0.62, -76.17; Cisneros-Heria 2003]. PASTAZA: Shell Mera [-1.51, 78.06; Lehr et al. 2004]. SUCUMBIOS: Santa Cecilia [0.05, -75.96; Duellman 1978; Lehr et al. 2004]; Puerto Libre, Río Aguarico [0.05, -77.48; Duellman 1978; Lehr et al. 2004]. FRENCH GUIANA: CAYENNE: Nouragues Rserve [4.07, 52.73; Gaucher et al. 2008]. PERU: CUSCO: Cashiriari-2, Lower Urubamba Region [-11.86, -72.78; Icochea et al. 2001]. HUÁNACO: Ambo [-10.07, -77.24; Lehr et al. 2004]; Huancapallac [-9.90, -76.39; Lehr et al. 2004]; Rio Llullapichis, Panguana [-9.62, -74.93; Lehr et al. 2004]. JUNÍN: Chanchamayo [-11.04, -75.32; Lehr et al. 2004]. LORETO: Iquitos, Centro Unión [-3.78, -73.12; Dixon & Soini 1986], Mishana [-4.01, -73.58; Dixon & Soini 1986], Moropon [-3.72, -73.33; Dixon & Soini 1986], Río Maniti [-3.54, -72.70; Dixon & Soini 1986]; Sarayacu [-6.73, -75.10; Boulenger 1894]; Yurimagua, Rio Huallaga [-5.91, -76.09; Boulenger 1894]. PASCO: Pozuzo [-10.15, -75.56; Lehr et al., 2004]; Iscozacin (Bosque de Villa America) [Toponym not found; Lehr et al. 2004]. SAN MARTIN: Moyobamba [6.05, -76.94; Boulenger 1894]. UCAYALI: Yarinacocha [-8.33, -74.62; Lehr et al. 2004].

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