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

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright copy 2013 Magnolia Press

Zootaxa 3716 (3) 349ndash394

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Taxonomic revision of Drymoluber Amaral 1930 (Serpentes Colubridae)

HENRIQUE CALDEIRA COSTA14 MAacuteRIO RIBEIRO MOURA2 3 amp RENATO NEVES FEIO1

1 Universidade Federal de Viccedilosa Departamento de Biologia Animal Museu de Zoologia Joatildeo Moojen CEP 36570-000 Viccedilosa MG

Brazil2 Universidade Federal de Minas Gerais Instituto de Ciecircncias Bioloacutegicas Departamento de Zoologia Laboratoacuterio de Herpetologia

Avenida Antocircnio Carlos 6627 Pampulha CEP 31270-901 Belo Horizonte MG Brazil3 Ecos Biota Consultoria Ambiental Rua Dr Milton Bandeira 95401 CEP 36570-000 Viccedilosa MG Brazil4 Corresponding author E-mail ccostahyahoocombr

Abstract

The present study is a taxonomic revision of the genus Drymoluber Amaral 1930 using meristic and morphometric char-acters 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 dis-tance 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 157ndash180 ventrals and 86ndash110 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 182ndash202 ventrals and 109ndash127 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 158ndash182 ventrals and 84ndash93 subcaudals and occurs in the Apuriacutemac Valley south of the Apuriacutemac and Pampas rivers in Peru

Key words Snakes South America taxonomy morphological variation clinal variation

Resumo

O presente estudo eacute uma revisatildeo taxonocircmica do gecircnero Drymoluber Amaral 1930 usando caracteres meriacutesticos e mor-fomeacutetricos aspectos da morfologia externa do hemipecircnis e de coloraccedilatildeo Dimorfismo sexual ocorre em D dichrous e D brazili mas natildeo foi detectado em D apurimacensis A variaccedilatildeo morfoloacutegica dentro de D dichrous tem relaccedilatildeo com a distacircncia geograacutefica entre as populaccedilotildees Ademais a variaccedilatildeo no nuacutemero de escamas ventrais e subcaudais em D dichrous e D brazili seguiu um padratildeo clinal com relaccedilatildeo agrave latitude e longitude Drymoluber dichrous se caracteriza pela presenccedila de 15-15-15 fileiras de escamas dorsais lisas com duas fossetas apicais 157ndash180 ventrais e 86ndash110 subcaudais ocorre na regiatildeo oriental da Cordilheira dos Andes Amazocircnia Escudo das Guianas Mata Atlacircntica e aacutereas de transiccedilatildeo desta com a Caatinga e o Cerrado Drymoluber brazili possui 17-17-15 fileiras de escamas dorsais lisas com duas fossetas apicais 182ndash202 ventrais e 109ndash127 subcaudais distribuindo-se pela Caatinga Cerrado Mata Atlacircntica e aacutereas de transiccedilatildeo entre esses dois uacuteltimos domiacutenios Drymoluber apurimacensis possui 13-13-13 fileiras de escamas dorsais lisas sem fossetas apicais 158ndash182 ventrais e 84ndash93 subcaudais e ocorre no Vale Apuriacutemac ao sul dos rios Apuriacutemac e Pampas no Peru

Palavras-chave Serpentes Ameacuterica do Sul taxonomia variaccedilatildeo morfoloacutegica variaccedilatildeo clinal

Introduction

The genus Drymoluber Amaral 1930 comprises three species of aglyphous medium-sized diurnal and terrestrial

Colubrinae snakes (Gomes 1918 Amaral 1930 Martins amp Oliveira 1998 Argocirclo 2004a Lehr et al 2004) that are

distributed in tropical South America east of the Andes (Peters amp Orejas-Miranda 1970 Lehr et al 2004 Cacciali

et al 2005 Vrcibradic 2007)

Accepted by H Zaher 2 Jul 2013 published 24 Sept 2013 349

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 ldquosupposedly bought in Surinamrdquo Five years later Cope (1868)

and Guumlnther (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 ldquoElaphe dichroardquo 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 ldquoElaphe dichrousrdquo 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 Penaacutepolis state of Satildeo

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 amp 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 Apuriacutemac 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 (eg Borges-Nojosa amp Lima 2001

Passos amp Brandatildeo 2002 Santana et al 2008) Amazonian savannas (Franccedila et al 2006) Caatinga (Vanzolini 1981

Rodrigues 2003 Argocirclo 2004b) Cerrado (Pavan amp Dixo 2004 Franccedila amp Arauacutejo 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 (eg 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)

COSTA ET AL350 middot Zootaxa 3716 (3) copy 2013 Magnolia Press

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 amp

Savage (1960) for hemipenial characters Hemipenes selected for morphological comparisons (n=24) were

prepared following the methods proposed by Pesantes (1994) and Zaher amp 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 01 mm with a caliper except for the snout-vent and tail

lengths which were measured with a flexible ruler to the nearest 10 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 (eg Peters 1964) WX+YZ 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 ocellispots 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

Zootaxa 3716 (3) copy 2013 Magnolia Press middot 351TAXONOMIC REVISION OF DRYMOLUBER

located 3) search in the online Gazetteer Glosk (httpwwwgloskcom) the software Google Earthreg and the

gazetteers by Paynter (1982 1992 1993 1997) and Paynter amp Taylor (1991) and 4) consultation with other

researchers Geographic coordinates are expressed in decimal degrees the latitude followed by the longitude (eg -

1015 -5945)

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 MG1mdash13 midbody scale

rows from the Andean region (Drymoluber apurimacensis n=5) MG2mdash15 midbody scale rows from the Andean

region (Drymoluber dichrous n=19) MG3mdash15 midbody scale rows from the Amazonia north of the Amazon

River (D dichrous n=61) MG4mdash15 midbody scale rows from the Amazonia south of the Amazon River (D

dichrous n=136) MG5mdash15 midbody scale rows from the Atlantic Forest north of the Doce River (D dichrous

n=54) MG6mdash15 midbody scale rows from the Atlantic Forest south of the Doce River (D dichrous n=9) MG7mdash

15 midbody scale rows from the Cerrado Amazonian savannas and transitional areas between them and forest

regions (D dichrous n=12) MG8mdash17 midbody scale rows from the Atlantic Forest south of the Doce River (D

brazili n=5) MG9mdash17 midbody scale rows from the Cerrado and transitional areas with the Atlantic Forest (D

brazili n=72) MG10mdash17 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 amp 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=445ndash590 mm) and four males (AMNH 22491 AMNH 91812 IBSP 69567 MPEG

20331 SVL=425ndash485 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 Kolmogorov-

Smirnov 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 70 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

amp 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


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 70 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 amp 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 amp 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 Paraiacuteso (Rondocircnia Brazil) 3) UHE Balbina (Presidente Figueiredo Amazonas Brazil) 4) Carajaacutes (Paraacute Brazil) 5) Beleacutem + Benevides + Castanhal + Satildeo Domingos do Capim (Paraacute Brazil) 6) BR 316 (Maranhatildeo Brazil) 7) Aracruz + Linhares + Vitoacuteria (Espiacuterito Santo Brazil) 8) Arataca + Barro Preto + Buerarema + Camacan + Ilheacuteus + Una (Bahia Brazil) 9) Barra do Rocha + Cairu + Itacareacute + Ituberaacute + Nova Ibiaacute (Bahia Brazil) 10) Maranguape + Pacoti + Pacatuba (Cearaacute Brazil) Females 1) Cabeceras del Rio Arajuno + Cabeceras del Rio Bobonaza + Macas + Riobamba + Riacuteo Liguino (Ecuador) 2) Iquitos (Loreto Peru) 3) Porto Walter + Reserva Extrativista Riozinho da Liberdade (Tarauacaacute Acre Brazil) 4) Porto Velho (Rondocircnia Brazil) 5) FLONA Caxiuanatilde (Melgaccedilo Paraacute Brazil) 6) Carajaacutes (Paraacute Brazil) 7) Buerarema + Ilheacuteus + Itabuna + Una (Bahia Brazil) 8) Camacan + Ibirataia + Ituberaacute + Nova Ibiaacute (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


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 Dsup2 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 amp 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 andor longitudinal clinal patterns

Results and Discussion

Sexual dimorphism

Drymoluber dichrousmdashFemales have more ventrals than males (F(1285) = 25211 p lt 0001) and the variation

in the number of subcaudals was not significant Males had higher values with respect to morphometric variables

(Λ = 0841 F(5135) = 5071 p lt 0001) with greater snout-vent length (F(1139) = 16235 p lt 0001) greater tail length

(F(1139) = 11078 p lt 0001) greater head length (F(1139) = 21180 p lt 0001) greater head width in the supraocular

region (U = 2729 p lt 0001) greater internasal distance (F(1139) = 16573 p lt 0001) greater eye diameter (U =

3165 p lt 0001) and greater eye-nostril distance (F(1139) = 18453 p lt 0001)

Drymoluber brazilimdashFemales of Drymoluber brazili also had a higher number of ventrals than males (F(181) =

27081 p lt 0001) while the variation in the number of subcaudals was not significant Morphometric data did not

show sexual dimorphism

Drymoluber apurimacensismdashSexual 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

= 0967 p lt 0001) and SC (r = 0864 p lt 0001) 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 =

0337 p = 0023) and females (r = 0513 p = 0005) 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 amp 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)


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 andor longitude In D dichrous there is a decrease from south to north in the

number of ventrals in males (r = -0226 p = 0003) and females (r = -0195 p = 0037) and in the number of

subcaudals in females (r = -0242 p = 0022) With regard to longitude significant clinal variation occurs only in

the number of subcaudals increasing from west to east in both sexes (males r = 0410 p lt 0001 females r =

0434 p lt 0001) (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

NS = not significant = p lt 005 = p lt 001 = p lt 0001 Cum prop = cumulative proportion of eigenvalues in percentage () for the first three principal components For variable symbols see Material and methods (lower case lsquolrsquo and lsquorrsquo 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 0812 0188 -0511 0967 0102 NS -0225

SC 0557 -0422 0702 0864 -0298 0403275

ILGl -0084 -0023 0039 -0721 -0090 NS 0125

ILGr -0079 -0022 0043 -0737 -0094 NS 0151

MD 0063 0015 -0010 0898 0100 NS -0055 NS

ATl 0061 0595 0285 0176 0777 0302

PTr 0055 0126 -0072 0304 0315 -0147

ATr 0049 0633 0384 0137 0796 0393

PTl 0023 0109 -0075 0137 0287 -0159

SLOl 0023 0014 0007 0399 0116 NS 0049 NS

SLOr 0022 0005 -0005 0378 0039 NS -0033 NS

ILC1r 0016 -0005 -0004 0396 -0058 NS -0037 NS

ILC1l 0015 0005 -0007 0350 0056 NS -0063 NS

LO -0007 -0001 -0008 -0223 -0016 -0090 NS

ILC2r 0005 -0019 0006 0065 NS -0115 NS 0032 NS

ILr 0004 0005 0006 0149 0084 NS 0078 NS

ILl 0003 0008 00009 0119 NS 0116 NS 0010 NS

SLl 00009 0005 0006 0048 NS 0125 NS 0118 NS

ILC2l -00006 0002 -0006 -0007 NS 0013 NS -0028 NS

SLr 00005 00004 -0003 0025 NS 0010 NS -0062 NS

Eigenvalues 144349 174292 194094 ndash ndash ndash

Cum prop () 66899 80776 89953 ndash ndash ndash


VE 0746 -0468 0439 0950 -0242 0190

SC 0626 0720 -0288 0896 0419 -0140 NS

ATr 0132 -0378 -0724 0392 -0454 -0728

ATl 0100 -0270 -0420 0367 -0402 -0523

ILGl -0083 0060 0009 -0757 0223 0029 NS

ILGr -0072 0070 0013 -0688 0272 0044 NS

MD 0065 -0024 -0001 0904 -0137 NS -0007 NS

continued on the next page



FIGURE 2 Projections of the individual scores resulting from the principal component analysis (PCA) and confidence ellipses (p gt 095) 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 MG2ndash7 = D dichrous MG8ndash10 = D brazili

TABLE 2 (Continued)


PTr 0043 -0124 -0077 0267 -0315 -0163

PTl 0038 -0135 -0120 0256 -0366 -0272

SLOl 0032 -0036 0029 0426 -0193 0132 NS

SLOr 0031 -0015 0010 0453 -0092 NS 0051 NS

LO -0019 0006 0001 -0451 0060 NS 0013 NS

ILC1r 0012 -0005 00002 0317 -0053 NS 0001 NS

ILC1l 0010 -0001 0001 0252 -0013 NS 0015 NS

ILC2l -0009 0013 0003 -0118 NS 0064 NS 0013 NS

ILr 0009 0012 0009 0265 0142 NS 0087 NS

ILl 0005 0007 0013 0197 0107 NS 0154 NS

ILC2r -0004 0011 -0003 -0057 NS 0057 NS -0013 NS

SLr 0001 00007 0001 0077 NS 0013 NS 0021 NS

SLl 00004 -0004 0006 0017 NS -0064 NS 0081 NS




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

NS = not significant = p lt 005 = p lt 001 = p lt 0001 Cum prop = cumulative proportion of eigenvalues in percentage () for the first three principal components For variable symbols see Fig 5 or Materials and Methods


SVL -0508 0277 0199 -0938 0198 0102 NS

HL -0445 -0894 0024 -0788 -0614 0011 NS

TL -0367 0154 -0910 -0815 0133 NS -0561

END -0365 0182 0237 -0912 0177 0164

ID -0337 0178 0098 -0864 0176 0070 NS

HWS -0307 0149 0178 -0922 0174 0148 NS

ED -0258 0105 0183 -0862 0136 NS 0170




FIGURE 4 Projections of the individual scores resulting from the principal component analysis (PCA) and confidence ellipses (p gt 095) 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 MG2ndash7 = D dichrous MG8ndash10 = 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 = 0416 p = 0027) From west to east

ventral plates in males increase in number (r = 0311 p = 0039 Figs 9 and 10)

Latitudinal andor longitudinal clinal patterns related to pholidosis morphometry and coloration have been

described for several snake species (eg Hoge et al 1977 Passos et al 2005 Allsteadt et al 2006 Passos amp

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 (eg McDiarmid 1968

Gardner amp 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 amp 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 (eg

Hoge et al 1977 Passos et al 2005 Passos amp 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 amp 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 posteriorndashanterior

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


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

NS = not significant = p lt 005 = p lt 001 = p lt 0001 Cum prop = cumulative proportion of eigenvalues in percentage () for the first three principal components For variable symbols see Fig 5 or Material and methods


SVL -0513 0124 0169 -0963 0085 NS 0093 NS

TL -0403 0683 -0539 -0801 0498 -0315

END -0392 -0108 0324 -0909 -0092 NS 0221

ID -0347 -0211 0269 -0855 -0191 0195

HL -0342 0028 0279 -0922 0028 NS 0221

HWS -0331 -0676 -0643 -0727 -0544 -0414

ED -0265 -0034 0103 -0876 -0041 NS 0100 NS




FIGURE 6 Projections of the Mahalanobis Dsup2 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


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 (eg D dichrous from Aripuanatilde (-1015 -5945) 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


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 (-1914 -4768) 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 Estaccedilatildeo Ecoloacutegica do Tripuiacute (-

2040 -4358 record of D dichrous) and Mariana (-2038 -4342 record of D brazili) Minas Gerais state

Caratinga RPPN Feliciano Miguel Abdala (-1971 -4181 record of D dichrous) and Alvarenga (-1941 -4172

record of D brazili) Minas Gerais state Missatildeo Velha Santo Antocircnio water spring (-741 -3921 record of D

dichrous) and Milagres (-731 -3895 record of D brazili) Cearaacute 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 Paraiacuteso Rondocircnia Brazil) B) Light stripe present although marked with two large spots (IBSP 16499 D brazili Turiba do Sul Satildeo Paulo Brazil) C) Light stripe absent (MZUSP 14298 D brazili UHE Luiacutes Eduardo Magalhatildees Tocantins Brazil) Photos Henrique C Costa

Drymoluber Amaral 1930

Drymoluber Amaral 1930 Memoacuterias 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) 157ndash

202 ventrals d) 84ndash127 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) 14ndash26 maxillary teeth l) ontogenetic

variation in the dorsal coloration of body and head (small specimens have dark and whitered 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


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 amp 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 (-2345) from tropical to subtropical seems to be a decisive factor limiting the austral distribution of

Drymoluber (Beacuternils 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 amp 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)


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 koumlniglich Akademie der Wissenschaften zu Berlin 29 p 284 Syntypes ZMB 1661 ZMB 1662 ZMB 2603

Herpetodryas occipitalis Guumlnther 1868 Annals and Magazine of Natural History Fourth Series 1 p 420 Holotype BMNH 194611461 formerly 186791728

Spilotes piceus Cope 1868 Proceedings of the Academy of Natural Sciences of Philadelphia 20 p 105ndash106 Holotype ANSP 3920

Coluber dichrousmdashBoulenger 1894 Catalogue of the Snakes in the British Museum (Natural History) Volume II British Museum of Natural History London p 30ndash31

Elaphe dichrousmdashGomes 1918 Memoacuterias do Instituto Butantan 1 p 67Drymoluber dichrousmdashAmaral 1930 Memoacuterias do Instituto Butantan 4 p 337

Lectotype (here designated in accordance with Article 74 of the International Code of Zoological Nomenclature)

Museum fuumlr 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 ldquoprobably from Surinamerdquo or ZMB 1662 which has a broken tail

Paralectotypes Museum fuumlr 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 fuumlr Naturkunde Berlin and on the oldest label of the specimen However a newer label (with the name

Drymoluber dichrous) indicates ldquoBrasilien Beckerrdquo The reason and source for the adjusted locality is unknown

even to the current curator (MO Roumldel 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 fuumlr Naturkunde Berlin of the type series of D dichrous (MO Roumldel pers com) Thus the type

locality of D dichrous was reported as ldquoBrazil and Surinamrdquo (eg Peters amp 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 GW 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 Proenccedila a path of the ancient royal road to Vila Rica (currently the

municipality of Ouro Preto) (Freyreiss 1907 Papavero 1971) (Fig 13 localities 1ndash12) In September 1814

Freyreiss explored the region around the Abaeteacute Indaiaacute and Satildeo Francisco Rivers later returning to Vila Rica

(Freyreiss 1907 Papavero 1971) (Fig 13 localities 12ndash18ndash12) In December 1814 he began a second trip to study

Indian tribes travelling from Vila Rica to the vicinities of Presiacutedio Satildeo Joatildeo Batista (currently the municipality of

Visconde do Rio Branco) (Fig 13 localities 12ndash23) 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 Espiacuterito Santo and Bahia provinces (Wied 1989 Papavero 1971) In

February 1816 the naturalists were in the Mucuri River southern Bahia (Fig 13 localities 1 24ndash37) There

Freyreiss decided to return to Espiacuterito Santo while Wied continued his travel northward (Wied 1989 p 186) In

Espiacuterito Santo Freyreiss visited again the localities of Satildeo Mateus and Linhares (Fig 13 localities 35 and 36) and


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 Colocircnia Leopoldina currently part of the municipality

of Nova Viccedilosa (Papavero 1971) (Fig 13 locality 38) There are some inconsistencies related to the place of death

of Freyreiss in 1825 Loumlfgren (1902) and Papavero (1971) wrote that the German naturalist died in Colocircnia

Leopoldina while Rocha (1972 ldquo1973rdquo) 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 Wiedrsquos expedition It is known that from his trips through Espiacuterito 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) 157ndash173 ventrals in males 160ndash

180 in females c) 87ndash110 subcaudals in males 86ndash109 in females d) 19ndash26 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 182ndash200 ventrals in males and 185ndash202 in

females 109ndash127 subcaudals in males and 109ndash126 in females Drymoluber apurimacensis is not distinguishable

from D dichrous based on ventrals and subcaudals counts having 158ndash164 ventrals in males and 166ndash182 in

females 84ndash93 subcaudals in males and 87ndash91 in females Drymoluber apurimacensis has 14ndash16 maxillary teeth

Small specimens of D dichrous have dark crossbands 15ndash7 scales wide (mean 36) and light interspaces 05ndash

25 scales wide (mean 08) while in D apurimacensis the dark crossbands are 1ndash2 scales wide and the pale

interspaces are 2ndash3 scales wide Juvenile specimens of D brazili have dark crossbands of similar width to those of

D dichrous (2ndash6 scales mean 36) but the pale interspaces are wider (05ndash5 scales mean 16)

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 246 mm length (42 of the SVL) greatest width of head 129 mm (52 of its length) width of head at

the supraoculars 93 mm internasal distance 51 mm eye diameter 445 mm eye-nostril distance 45 mm The

morphometric measurements were taken by Christoph Kucharzewski Museum fuumlr 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 15 times longer than wide supraoculars longer than

wide parietals about 15 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 (12+11) four anterior temporals (two uppers and

two lowers) and two posterior temporals (one upper and one lower) on the left side (22+11) 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


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) Espiacuterito 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 (Mageacute) 3 = Fazenda

Mandioca (Mageacute) 4 = Fazenda do Padre Correia (Correcircas Petroacutepolis) 5 = Fazenda das Sebollas (Inconfidecircncia Paraiacuteba do Sul) 6 = Rio Paraibuna 7 = Matias Barbosa 8 = Juiz de Fora 9 = Chapeacuteu DrsquoUvas (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 Satildeo Joanico (Maravilhas) 15 = Pompeu 16 = Rio Satildeo Francisco 17 = Fazenda do Comandante

de Indaiaacute (Quartel Geral) 18 = Quartel do Assunccedilatildeo (Coacuterrego dos Tiros Tiros) 19 = Mariana 20 = Santana dos Ferros (Guaraciaba) 21 = Santa Rita (Viccedilosa) 22 = Serra de S Beralde (Serra de Satildeo Geraldo Satildeo Geraldo) 23 = Presiacutedio de Satildeo

Joatildeo Batista (Visconde do Rio Branco) 24 = S Gonzalves (Satildeo Gonccedilalo) 25 = Freguesia de Maricaacute (Maricaacute) 26 = Araruama 27 = Cabo Frio 28 = Villa de S Joatildeo de Macaheacute (Macaeacute) 29 = Villa de S Salvador dos Campos dos Goytacazes (Campos dos Goytacazes) 30 = Itapemerim 31 = Povoaccedilatildeo de Piuma (Piuacutema) 32 = Nossa Senhora da Victoria (Vitoacuteria) 33 = Vila Nova do Almeida (Nova Almeida Serra) 34 = Quartel do Riacho (Riacho Aracruz) 35 = Linhares 36 = Barra do Satildeo Mateus (Satildeo Mateus) 37 = Rio Mucuri (Mucuri) 38 = Colocircnia Leopoldina (Nova Viccedilosa) 39 = Caravelas 40 = Canavieiras 41 = Salvador Dashed line = Travel from Rio de Janeiro to Quartel do Assunccedilatildeo (Tiros) in 1814 (Localities 1ndash12 12ndash11 11ndash18) Dotted line = Travel from Villa Rica (Ouro Preto) to the vicinities of Presiacutedio Satildeo Joatildeo Batista (Visconde do Rio Branco) from December 1814 to January 1815 (Localities 12ndash23) Dashed-dotted line = Travel from Rio de Janeiro to Mucuri from July 1815 to 1816 (Localities 1 24ndash38) Localities 39ndash41 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)


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 Roumldel


Coloration of the lectotype Peters (1863) described the coloration of Herpetodryas dichroa as ldquoDorsum

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 eyesrdquo 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 15) 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 15) The venter of some specimens is the same color as dorsal scales or a little paler (n=3 15) One

specimen (005) 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=1 005) the subcaudals may have small black dots in

larger numbers in the posterior region (n=1 005) 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 65)

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 85)

Coloration of adults in life Based on descriptions in the literature (eg Cunha amp Nascimento 1978 Martins

amp Oliveira 1998 Bartlett amp Bartlett 2003 Argocirclo 2004a) and some photographs (Fig 15) we noted some variation

in the color pattern of adult specimens of D dichrous in lifemdashbut 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=43 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 285ndash391 mm) the banded coloration has disappeared

completely The tail bands are rarely visible (only in some specimens with SVL lt 385 mm) Dorsal crossbands

vary from de 15ndash7 vertebralparavertebral scales wide (mean 36 SD=087 n=306 crossbands)

The last crossband anterior to the cloacal shield tends to be the narrowest (15ndash5 scales mean 29 SD=079

n=48) and the fifteenth after the head the widest (2ndash7 scales mean 36 SD=075 n=71) The light interspaces

between dark crossbands vary between 05ndash25 scales (mean 08 SD=043 n=306 interspaces) The interspaces

anterior to the last crossband and the fifth crossband anterior to the cloacal shield are the narrowest with 05ndash1

scales (mean 05 SD=01 n=48 and mean 05 SD=009 n=49) The interspace posterior to the first crossband and

the interspace anterior to the fifth crossband are the widest with 05ndash25 scales (mean 117 SD=049 n=67 and

mean 096 SD=046 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 15) or only on the posterior half (n=2 25)

The subcaudals in juveniles are the same pattern as those in adults (cream color with darkened lateral edges) (n=73

935) although in some specimens these plates are completely black (n=5 65)

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=19 261) maculate (n=5 68) or absent (n=49 671) (Fig 13)


Like in the adults the gular region of the juveniles usually is cream colored (n=48 615) but black marks

may occur on the infralabials and chinshields (n=30 385) The supralabials have distinct dark marks on their

lateral edges (n=51 654) but sometimes these marks are pale as in adults (n=8 1025) or even barely visible

(n=19 2435)

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 Choccedila Bahia Brazil B) Reserva Extrativista Riozinho da Liberdade Tarauacaacute Acre Brazil C) Reserva Ducke Manaus Amazonas Brazil D) Nouragues Station French Guiana E) Parque Estadual Guajaraacute-Mirim Rondocircnia Brazil F) Marabaacute Paraacute Brazil Photos Marco Antocircnio de Freitas (A) Paulo Seacutergio Bernarde (B) Rafael de Fraga (C) Maeumll Dewynter (D) Laurie J Vitt (E) and Pedro L V Peloso (F)


luber Amaral 1930 Most common (over 50 of

Drymoluber dichrous

oce

eas

Eastern Andes Amazonia Guiana Shield

Brazilian Atlantic Forest Atlantic Forestndash

Caatinga and Atlantic ForestndashCerrado

transitional areas

1050 mm (MPEG 17235)

15ndash15ndash15 (n=289) 15ndash15ndash17 (n=1)

15ndash15ndash13 ( n=1)

2

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

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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)

longer than high (n=193) as high as long

(n=92) higher than long (n=5)

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

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

ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)

helliphellipcontinued on the next page

CO

ST

A E

T A

L

370

middot Zootaxa 3716 (3) copy

2013 M

agnolia Press

TABLE 5 Summary of geographic distribution and morphological variation of the three species of Drymofrequency) character states are represented in bold Drymoluber apurimacensis Drymoluber brazili

Geographic distribution South of the Apurimaacutec and

Pampas Rivers in the Peruvian

Serraniacutea Esteparia ecoregion

Cerrado Caatinga Atlantic Forests of the lower D

River and Atlantic ForestndashCerrado transitional ar

Maximum snout-vent length 670 mm (MHNSM 18647) 1178 mm (IBSP 34369)

Number of dorsal scale rows 13ndash13ndash13 17ndash17ndash15

Dorsal apical pits 0 2

Cloacal shield Entire Entire (n=82) Divided (n=1)

Number of pre-ventrals 1 (n=1) 2 (n=2) 1 (n=74) 2 (n=8) 3 (n=1)

Number of ventrals in males 158ndash164 (n=2) 182ndash200 (mean 189 SD=361 n=46)

Number of ventrals in females 166ndash182 (n=2) 185ndash202 (mean 193 SD=368 n=37)

Number of subcaudals in males 84ndash93 (mean 88 SD=458 n=3) 109ndash127 (mean 118 SD=437 n=29)

Number of subcaudals in females 87ndash91 (mean 89 SD=208 n=3) 109ndash126 (mean 119 SD=445 n=26)

Loreal longer than high longer than high

Preoculars 1 1

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

Temporal formula 1+11 1+11 (n=9 sides) 1+23 (n=1 side) 11+1 (n=1 s

11+11 (n=90 sides) 11+12 (n=8 sides) 11+2

sides) 11+22 (n=1 side) 11+31 (n=1 side) 12

(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41

side) 21+11 (n=5 sides) 21+12 (n=3 sides) 2

(n=1 side) 22+42 (n=1 side) 23+12 (n=1 side

Zootaxa 3716 (3) copy

2013 Magnolia P

ress middot 371

Drymoluber dichrous

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

des) iiindashiv (n=1 side) iiindashv (n=454 sides)

iiindashvi (n=2 sides) ivndashv (n=88 sides)

ivndashvi (n=34 sides)


10 (n=7 sides)

1st pair

indashiv (n=334 sides) indashv (n=244 sides)

indashvi (n=2 sides)

ivndashv (n=12 sides) ivndashvi (n=322 sides)

vndashvi (n=240 sides) vndashvii (n=4 sides)

vindashvii (n=2 sides)

vndashviii (n=4 sides) vndashix (n=7 sides)

vindashviii (n=75 sides) vindashix (n=485 sides)

vindashx (n=2 sides) viindashx (n=6 sides)

19ndash26 (mean =23 SD=121)

) with at least three regions where the scale count is

647 which according to Lehr et al (2004) has nine

TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)

Drymoluber apurimacensis Drymoluber brazili

Number of supralabials 8 7 (n=3 sides) 8 (n=143 sides) 9 (n=18 sides)

Supralabials contacting the eye

orbit

ivndashv iiindashv (n=1 side) ivndashv (n=152 sides) ivndashvi (n=2 si

vndashvi (n=9 sides)

Number of infralabials 7 (n=2 sides) 8 (n=4 sides)

9 (n=2 sides)


Infralabials contacting the

mental

1st pair 1st pair

Infralabials contacting the first

pair of chinshields

indashiv (n=4 sides) indashv (n=2 sides) indashiii (n=1 side) indashiv (n=21 sides)

indashv (n=140 sides) indashvi (n=2 sides)


second pair of chinshields

ivndashv (n=4 sides) v (n=2 sides) iii (n=1 side) iv (n=1 side) ivndashv (n=20 sides)

v (n=134 sides) vndashvi (n=8 sides)

Infralabials contacting the gulars vndashvii (n=2 sides)

vndashviii (n=4 sides)

iiindashviii (n=1 side) ivndashviii (n=1 side)


vndashx (n=6 sides) vindashix (n=4 sides)

vindashx (n=4 sides)

Maxillary teeth 14ndash16 (mean =15 SD=1) 19ndash25 (mean =23 SD=105)

Lehr et al (2004) cited 182 ventrals for MTKD 44669 (male) but this specimen is in poor condition (Fig 23Bimpaired 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 18infralabials

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 Choccedila Bahia Brazil D) Ubajara Cearaacute Brazil Photos Rafael de Fraga (A) William Quatman (B) Marco Antocircnio 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 (5ndash10 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 60ndash70 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 6ndash10 spines (mean 7 SD=122) 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 5ndash7 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 (265) had broken tail The tail of 54 of

those specimens (185 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


already recorded in several other Neotropical Colubrinae including Dendrophidion Drymobius Mastigodryas and

Scaphiodontophis (Mendelson III 1992 Slowinski amp Savage 1995 Prudente et al 2007 Leite et al 2009) Tail

length of specimens with complete tail is 259ndash484 of the SVL in males (mean 406 SD=318 n=122) and

354ndash653 in females (mean 41 SD=375 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


and environments with a relative degree of anthropization (Cunha amp Nascimento 1978 Martins amp Oliveira 1998

Argocirclo 2004a Franccedila et al 2006) Drymoluber dichrous is diurnal forages on the ground and rests on the

vegetation at night (Cunha amp Nascimento 1978 Martins amp 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 amp Nascimento 1978 Duellman 1978

2005 Dixon amp Soini 1986 Avila-Pires 1995 Martins amp Oliveira 1998 Vitt et al 2000 Borges-Nojosa amp Lima

2001 Pinto 2006 Bernarde amp Abe 2010 Veriacutessimo 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 speciesrsquo diet

Information about reproduction in D dichrous is scarce and restricted to Amazonian specimens Drymoluber

dichrous is oviparous and there are reports of 2ndash6 eggs per clutch (Fitch 1970 Martins amp 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 amp Oliveira (1998)

Drymoluber dichrous has several defensive behaviors When handled individuals will rotate their body

vibrate their tail and occasionally bite (Martins amp 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 amp

Brodie Jr (2004) suggested that young cross-banded specimens might mimic coralsnakes (Elapidae)

As mentioned above the long tail of D dichrous (mean 406 of the SVL in males and 41 in females) with

a tail breakage ratio of 185 (considering only those specimens where breakage occurred before collection) is

evidence of the defensive behavior known as pseudoautotomy (Slowinski amp 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 ldquotwo-coloredrdquo or

ldquotwo-skinsrdquo 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 Memoacuterias 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)mdashAmaral 1929 Memoacuterias do Instituto Butantan 4 p 11Drymoluber brazilimdashStuart 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 Satildeo 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)


Holotype Instituto Butantan Satildeo Paulo IBSP 696 adult male SVL 1090 mm TL 473 mm collected on

September 1914 in the Estaccedilatildeo Ferroviaacuteria de Engenheiro Lisboa (currently inactive) near the municipality of

Uberaba (-1980 -4760) Minas Gerais Brazil Specimen personally examined by the senior author

Paratypes Instituto Butantan Satildeo Paulo IBSP 383 adult male SVL 863 mm TL 394 mm collected in

February 1913 in the Estaccedilatildeo Ferroviaacuteria Santa Eudoacutexia (currently inactive) municipality of Satildeo Carlos (-2177

-4778) Satildeo Paulo Brazil IBSP 573 adult female SVL 854 mm TL 422 mm collected in February 1914 in the

Estaccedilatildeo Ferroviaacuteria de Sampaio Vidal (currently inactive) municipality of Ribeiratildeo Bonito (-2207 -4818) Satildeo

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 Estaccedilatildeo Ferroviaacuteria

Java (currently inactive) municipality of Boa Esperanccedila do Sul (-2199 -4839) Satildeo Paulo Brazil IBSP 1286

adult female SVL 898 mm TL 402 mm collected in May 1917 in the Estaccedilatildeo Ferroviaacuteria Pedregulho (currently

inactive) municipality of Pedregulho (-2026 -4748) Satildeo 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 Estaccedilatildeo Ferroviaacuteria Engenheiro

Lisboa (type locality) was located at km 555 of the Tronco-Catalatildeo railroad which at the time of the collection of

the holotype (September 1914) went from Campinas Satildeo Paulo (-2209 -4705) to Ipameri Goiaacutes (-1772 -4815)

(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-Catalatildeo 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

Eudoacutexia 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 Satildeo Paulo from the southeast to the north and northwest

(Giesbrecht 2009 Cavalcanti 2010) On the other hand the Estaccedilatildeo Ferroviaacuteria Pedregulho was located at km 455

on the Tronco-Catalatildeo railroad

Diagnosis Drymoluber brazili can be distinguished from D apurimacensis and D dichrous by the following



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 158ndash164 ventrals in males and 166ndash182

in females 84ndash93 subcaudals in males and 87ndash91 in females Drymoluber dichrous has 157ndash173 ventrals in males

and 160ndash180 in females 87ndash110 subcaudals in males and 86ndash109 in females Drymoluber apurimacensis has 14ndash

16 maxillary teeth

Small specimens of Drymoluber brazili have dark crossbands 2ndash6 scales wide (mean 36) and light interspaces

05ndash5 scales wide (mean 16) while in a single specimen of D apurimacensis the dark crossbands are 1ndash2 scales

wide and the light interspaces 2ndash3 scales wide Young specimens of D dichrous have dark crossbands with similar

width to those of D brazili (15ndash7 scales mean 36) but the pale interspaces are on average narrower (05ndash25

scales mean 08)

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 351 mm long (32 of the SVL) greatest width of head 1520 mm (43 of head length) width of

head at the supraoculars 113 mm internasal distance 61 mm eye diameter 60 mm eye-nostril distance 59 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




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) (11+11) 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 Guimaratildees 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=49 778) is

uniform bluish-gray bluish-brown or yellowish-brown throughout A few specimens (n=13 206) are grayish-

brown 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=49 778) is immaculate yellowish-cream and the lateral edges of ventrals

have the same color as the dorsal scales In one specimen (16) the dorsal coloration extends into the ventral

region in other specimens (n=13 206) 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 (381) 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 79) In a

few the subcaudals have more than half their area dark like the dorsals (n=1 16) or have black spots throughout

their area (n=1 16) In others black spots are numerous only in the posterior region of the tail (n=24 381) or

the posterior part of tail is completely black (n=8 127)

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=59 936) but in some specimens visible markings are absent (n=4 64)

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 reddish-

brown This same pattern is seen in the illustration (Fig 19A) of one specimen from Visconde de Parnaiacuteba Satildeo

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 2ndash6

vertebralparavertebral scales (mean 36 SD=089 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 2ndash6 scales wide

(mean 4 SD=095 n=20 100) The widths of the pale interspaces vary between 05ndash5 scales (mean 16


SD=097 n=62 interespaces) The interspace anterior to the fifth crossband anterior to the cloaca is the narrowest

with 05 or 15 scales (n=2 10) The interspace posterior to the first crossband is the widest with 1ndash5 scales

(mean 232 SD=090 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 (AndashD F) and Ana Baacuterbara Barros (E)


FIGURE 19 Color in life of some adult specimens of Drymoluber brazili (Gomes 1918) A) Visconde de Parnaiacuteba Satildeo 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 Goiaacutes 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



single and centrolinear Lobe about half of the hemipenis length with papillate calyces (5ndash10 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 8ndash13 spines (mean 8

SD=197) 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


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 6ndash7 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 (337) had broken tail The tail of 24

of those specimens (289 of the total examined) was healed suggesting the presence of pseudoautotomy in this

species Tail length of specimens with complete tail is 381ndash627 of the SVL in males (mean 442 SD=423

n=29) and 4111ndash56 (mean 457 SD=35 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 (Canindeyuacute 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) Franccedila et al

(2006) reported the species in Vilhena Rondocircnia Brazil (-1274 -6015) in Amazonian savannas However that

specimen (CHUNB 12791) is actually a D dichrous There is a record from Aripuanatilde Mato Grosso Brazil (-

1015 -5945) 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 Cearaacute Brazil) actually is a young

Drymarchon corais MZUSP 11263 (from Gauacutecha 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 (-1954 -4064)

and Baixo Guanduacute (-1952 -4102) state of Espiacuterito Santo and in Aimoreacutes (-1950 -4106) 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 21000 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 amp 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 Franccedila amp Arauacutejo 2006 Franccedila et

al 2008) and seems to be absent from altered habitats (Franccedila amp Arauacutejo 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 (Franccedila amp Arauacutejo 2006 Franccedila 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 amp 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


FIGURE 21 Hemipenis of Drymoluber brazili IBSP 33668 Goiacircnia Goiaacutes Brazil A) Sulcate face B) Asulcate face MCNR 1736 Usina Hidreleacutetrica de Irapeacute Minas Gerais Brazil C) Sulcate face D) Asulcate face Scale bar = 1 cm Photos Henrique C Costa


There is a report of an unidentified Gymnophthalmidae as prey of Drymoluber brazili (Franccedila et al 2008)

Additionally Pavan amp 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

Franccedila amp Arauacutejo (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 ldquoWhen 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 tailrdquo (translation by the authors) Brodie III amp Brodie Jr (2004) suggested

that young cross-banded specimens might mimic coralsnakes (Elapidae)

The long tail of Drymoluber brazili (mean 442 of the SVL in males and 457 in females) with a tail

beakage ratio of 289 (considering only those specimens where the breakage certainly occurred before the

collection) is ample evidence of pseudoautotomy (Slowinski amp 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 amp 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 (-1364 -7288)

Abancay Province Apurimaacutec 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 (-1365 -7338) Huancarama district Andahuaylas Province

Apurimaacutec Department Peru 3300 m above sea level (specimen examined by the senior author Fig 23A)

Museum fuumlr Tierkunde Dresden MTKD 44669 adult female SVL 653 mm TL 253 mm killed by a local farmer

in May 2002 in Cconoc (-1306 -7264) Abancay Province Apurimaacutec 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 Apurimaacutec 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) 158ndash164 ventrals in males 166ndash

182 in females c) 84ndash93 subcaudals in males 87ndash91 in females d) 14ndash16 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 182ndash200 ventrals in males and 185ndash202 in

females 109ndash127 subcaudals in males and 109ndash126 in females Drymoluber dichrous is not distinguishable from

D apurimacensis based on ventrals and subcaudals counts having 157ndash173 ventrals in males and 160ndash180 in

females 87ndash110 subcaudals in males and 86ndash109 in females Drymoluber brazili has 19ndash15 maxillary teeth and D

dichrous has 19ndash16


FIGURE 22 Holotype of Drymoluber apurimacensis Lehr Carrillo amp Hocking 2004 (MHNSM 20672) Photo Claudia P Torres Gastello

The only known juvenile of Drymoluber apurimacensis (holotype) has dark crossbands 1ndash2 scales wide and

pale interspaces 2ndash3 scales wide Juveniles of D brazili have dark crossbands 2ndash6 scales wide (mean 36) and pale

interspaces 05ndash5 scales wide (mean 16) Juveniles of D dichrous have dark crossbands 15ndash7 scales wide (mean

36) and pale interspaces 05ndash25 scales wide (mean 08)

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 (HC Costa

pers obs M Auer pers comm CP 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+11 in the

present study) while D brazili and D dichrous have temporal formulae of 2+2 (11+11 in the present study) As

previously discussed marked polymorphism of temporal plates characterizes Drymoluber and the temporal

formula 1+11 also occurs in D dichrous and D brazili although the 11+11 formula is more frequent


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 23 of the body Subcaudals are bluish-gray and cream colored

FIGURE 23 Four of the five paratypes of Drymoluber apurimacensis Lehr Carrillo amp Hocking 2004 A) FMNH 81542 B) MTKD 44669 C) MTKD 45192 D) MTKD 45193 Photos Henrique C Costa (A) and Barbara Bastian (BndashD)

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 1ndash2 scales

wide) which become less visible in the posterior third of the body The light interspaces are 2ndash3 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)


single and centrolinear Lobe with just less than half of the hemipenis length with papillate calyces (5ndash10

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 60ndash70 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


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 344ndash417 of the SVL in males (mean 385 SD=375 n=3) and

354ndash409 (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 Apurimaacutec and Pampas Rivers department of Apurimaacutec in the Serraniacutea

Esteparia ecoregion The area is characterized by matorral seco and matorral sub-huacutemedo vegetation (Lehr et al

2004) Drymoluber apurimacensis probably is endemic to the deep valley of the Apuriacutemac 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 Apuriacutemac region (Lehr amp Catenazzi 2008

BirdLife International 2009 Knapp amp 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 Apurimaacutec Department Peru A) Sulcate face B) Asulcate face Scale bar = 05 cm Photos Henrique C Costa


Etymology The specific name apurimacensis means ldquofrom Apurimaacutecrdquo Apurimaacutec the name of the region

where the species is known to occur is a Quechuan (an indiginous language from the Andes) word meaning ldquothe

mountain spirit that speaksrdquo (Lehr et al 2004)

Key to the species of Drymoluber Amaral 1930

1A Seventeen (17) midbody scale rows with reduction to fifteen (15) rows anterior to the cloacal shield 182ndash202 ventrals 109ndash

127 subcaudals D brazili

1B Thirteen (13) or fifteen (15) midbody scale rows without posterior reduction 157ndash182 ventrals 84ndash110 subcaudals 2

2A Thirteen (13) dorsal scale rows at midbody without apical pits 158ndash182 ventrals 84ndash93 subcaudals 14ndash16 maxillary teeth

D apurimacensis

2B Fifteen (15) dorsal scale rows at midbody with two apical pits 157ndash180 ventrals 86ndash110 subcaudals 19ndash26 maxillary teeth

D dichrous

Acknowledgements

We are indebt to several persons and institutions the Coordenaccedilatildeo de Aperfeiccediloamento de Pessoal de Niacutevel

Superior (CAPES) and the Smithsonian Institution for schorlarships to the senior author Guarino R Colli Marcela

A Brasil and Mariana CM Viana (CHUNB) Paulo CA Garcia and Patriacutecia S Santos (CHUFMG) Jacques HC

Delabie (CZGB) Francisco Luiacutes Franco and Valdir Joseacute Germano (IBSP) Maria Rita S Pires and Bruna Imai

(LZVUFOP) Luisa Sarmento and Fabiano Lanschi (MBML) Glaacuteucia Maria F Pontes (MCP) Luciana B

Nascimento (MCN) Hussam Zaher and Carolina Mello (MZUSP) Antocircnio JS Argocirclo (MZUESC) Ronaldo

Fernandes and Marcelo Gomes (MNRJ) Ana Luacutecia C Prudente Joatildeo Fabriacutecio M Sarmento Alessandra

Travassos and Paula CR Almeida (MPEG) Paulo S Bernarde (UFACF) Diva Maria Borges-Nojosa and Daniel

C Lima (UFC) Marcos Andreacute 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 Roumldel 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 Maeumll

Dewynter Marco Antocircnio 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 Joseacute 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 Deacutebora C Rodrigues Fabiano

Godoy Fiona Wilkinson Jeremy F Jacobs Kathleen M Kelly Kenneth Tighe Larry Matheson Marcelo J

Sturaro Maacutercio M Morais Jr Pedro LV 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 Beacuternils Rocircmulo 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

References

Adler K (2007) Contributions to the History of Herpetology Volume 2 Society for the Study of Amphibians and Reptiles Saint Louis 380 pp

Alexander AA amp Gans C (1966) The pattern of dermal-vertebral correlation in snakes and amphisbaenians Zoologische

Mededelingen 41 171ndash190Allsteadt J Savitzky AH Petersen CE amp Naik DN (2006) Geographic variation in the morphology of Crotalus horridus

(Serpentes Viperidae) Herpetological Monographs 20 1ndash63Amaral A (1923) New genera and species of snakes Procceedings of the New England Zoological Club 8 85ndash105


Amaral A (1926) Ophidia from South America in the Carnegie Museum a critique of Dr LE Griffins ldquoCatalog of the Ophidia from South America at present (June 1916) contained in the Carnegie Museumrdquo Annals of Carnegie Museum 16 319ndash323

Amaral A (1929) Estudos sobre ophidios neoacutetropicos XVII Valor sistemaacutetico de varias formas de ophidios neotropicosMemoacuterias do Instituto Butantan 4 3ndash68

Amaral A (1930) Estudos Sobre Ophidios Neotroacutepicos XXII ndash Sobre a espeacutecie Coluber dichrous (Peters) Boulenger 1894Memoacuterias do Instituto Butantan 4 333ndash337

Amaral A (1978) Serpentes do Brasil Iconografia Colorida Editora Melhoramentos Satildeo Paulo 247 ppArgocirclo AJS (2004a) As Serpentes dos cacauais do sudeste da Bahia Editora da UESC Ilheacuteus 260 ppArgocirclo AJS (2004b) Geographic Distribution Drymoluber brazili Herpetological Review 35 191Arnold SJ amp Peterson CR (2002) A Model for optimal Reaction Norms The Case of the Pregnant Garter Snake and Her

Temperature-Sensitive Embryos The American Naturalist 160 306ndash316 httpdxdoiorg101086341522

Avila-Pires TCS (1995) Lizards of Brazilian Amazocircnia (Reptilia Squamata) Zoologische Verhandenlingen 299 1ndash706Avila-Pires TCS Vitt LJ Sartorius SS amp Zani PA (2009) Squamata (Reptilia) from four sites in southern Amazonia

with a biogeographic analysis of Amazonian lizards Boletim do Museu Paraense Emiacutelio Goeldi Ciecircncias Naturais 4 99ndash118

Bartlett RD amp Bartlett P (2003) Reptiles and Amphibians of the Amazon University Press of Florida Gainesville 291 ppBernarde PS amp Abe AS (2010) Haacutebitos alimentares de serpentes em Espigatildeo do Oeste Rondocircnia Brasil Biota Neotropica

10 167ndash173 httpdxdoiorg101590s1676-06032010000100017

Beacuternils RS Giraudo AR Carreira S amp Cechin SZ (2007) Reacutepteis das porccedilotildees subtropical e temperada da regiatildeo neotropical Ciecircncia amp Ambiente 35 101ndash136

BirdLife International (2009) Species factsheet Synallaxis courseni Available from httpwwwbirdlifeorg (Accessed 28 Apr 2010)

Bokermann W (1957) Atualizaccedilatildeo do itineraacuterio da viagem do Priacutencipe de Wied ao Brasil (1815ndash1817) Arquivos de Zoologia

Satildeo Paulo 10 209ndash251Borges-Nojosa DM amp Lima DC (2001) Dieta de Drymoluber dichrous (Peters 1863) dos Brejos-de-altitude do Estado do

Cearaacute Brasil (Serpentes Colubridae) Boletim do Museu Nacional do Rio de Janeiro 468 01ndash05Boulenger GA (1894) Catalogue of the Snakes in the British Museum (Natural History) Volume II British Museum of

Natural History London 382 ppBrodie III ED amp Brodie Jr ED (2004) Venomous Snake Mimicry In Campbell JA amp Lamar WW (Eds) Venomous

Reptiles of the Western Hemisphere Volume II Cornell University Press Ithaca pp 617ndash633Cacciali P Fernandeacutez S amp Ramireacutez F (2005) Geographic Distribution Drymoluber dichrous Herpetological Review 36

339Carnaval AC amp Moritz C (2008) Historical climate modelling predicts patterns of current biodiversity in the Brazilian

Atlantic forest Journal of Biogeography 35 1187ndash1201Cavalcanti FR (2010) Centro-Oeste Brasil Trens Ferrovias e Ferreomodelismo Available from httpvfcobraziliajorbr

(Accessed 18 Aug 2010)Cavalcanti MJ amp Lopes PRD (1993) Anaacutelise morfomeacutetrica multivariada de cinco espeacutecies de Serranidae (Teleostei

Perciformes) Acta Biologica Leopoldensia 15 53ndash64Cisneros-Heredia DF (2003) Herpetofauna de la Estacioacuten de Biodiversidad Tiputini Amazoniacutea Ecuatoriana Ecologiacutea de una

comunidad taxonoacutemicamente diversa con comentarios sobre metodologiacuteas de inventario In De la Torre S amp Reck G (Eds) Ecologiacutea y Ambiente en el Ecuador Memorias del I Congreso de Ecologiacutea y Ambiente Ecuador paiacutes megadiverso CD-Rom Universidad San Francisco de Quito Quito pp 1ndash21

Cope ED (1868) An examination of the Reptilia and Batrachia obtained by the Orton Expedition to Ecuador and the Upper Amazon with notes on other species Proceedings of the Academy of Natural Sciences of Philadelphia 20 96ndash140

Costa LP (2003) The historical bridge between the Amazon and Atlantic Forest of Brazil a study of molecular phylogeography with small mammals Journal of Biogeography 30 71ndash86 httpdxdoiorg101046j1365-2699200300792x

Cunha OR amp Nascimento FP (1978) Ofiacutedios da Amazocircnia X ndash As cobras da regiatildeo leste do Paraacute Publicaccedilotildees Avulsas do

Museu Paraense Emiacutelio Goeldi 31 1ndash218de Queiroz K amp Good DA (1997) Phenetic clustering in biology A critique The Quarterly Review of Biology 72 3ndash30

httpdxdoiorg101086419656Dixon JR amp Soini P (1986) The reptiles of the upper Amazon basin Iquitos region Peru 2nd ed Wisconsin Milwaukee

Public Museum Milwaukee 154 ppDowling HG amp Savage JM (1960) A guide to the snake hemipenis a survey of basic structure and systematic chacarteristcs

Zoologica 45 17ndash28Duellman WE (1978) The biology of an equatorial herpetofauna in Amazonian Ecuador Miscellaneous Publications of the

Museum of Natural History University Kansas 65 1ndash352


Duellman WE (2005) Cusco Amazoacutenico The lives of amphibians and reptiles in na Amazonian Rainforest Comstock Publishing Associates Ithaca and London 433 pp

Embert D (2007) Distribution diversity and conservation status of Bolivian Reptiles PhD Dissertation Rheinischen Friedrichs-Wilhelms-Universitaumlt Bonn 429 pp

Fitch HS (1970) Reproductive cycles of lizards and snakes Miscellaneous Publications of the Museum of Natural History University Kansas 52 1ndash247

Fitch HS (1981) Sexual Size Differences in Reptiles Miscellaneous Publications of the Museum of Natural History University Kansas 70 1ndash72

Fox W (1948) Effect of temperature on development of scutellation in the garter snake Thamnophis elegans atratus Copeia 1948 252ndash262 httpdxdoiorg1023071438712

Franccedila FGR amp Arauacutejo AFB (2006) The Conservation Status of Snakes in Central Brazil South American Journal of

Herpetology 1 25ndash36 httpdxdoiorg1029941808-9798(2006)1[25tcsosi]20co2

Franccedila FGR Mesquita DO amp Colli GR (2006) A checklist of snakes from Amazonian savannas in Brazil housed in the Coleccedilatildeo Herpetoloacutegica da Universidade de Brasiacutelia with new distribution records Occasional Papers of the Sam Noble

Oklahoma Museum of Natural History 17 1ndash13Franccedila FGR Mesquita DO Nogueira CC amp Arauacutejo AFB (2008) Phylogeny and ecology determine morphological

structure in a snake assemblage in the central Brazilian Cerrado Copeia 2008 23ndash38 httpdxdoiorg101643ch-05-034

Franco FL (2012) A Coleccedilatildeo Herpetoloacutegica do Instituto Butantan da suaorigem ao incecircndio ocorrido em 15 de maio de 2010 Herpetologia Brasileira 1 22ndash31

Freire EMX (2000) Geographic Distribution Drymoluber dichrous Herpetological Review 31 55Freitas MA Franccedila DPF amp Veriacutessimo D (2012) Distribution extension of Drymoluber brazili (Gomes 1918) (Serpentes

Colubridae) for the state of Piauiacute Brazil Check List 8 168ndash169Freyreiss GW (1907) Viagem ao Interior do Brasil nos annos de 1814-1815 pelo naturalista G W Freyreiss (traduzido pelo

Dr Alberto Loumlfgren) Revista do Instituto Histoacuterico e Geograacutephico do Estado de Satildeo Paulo 11 158ndash228Gardner SA amp Mendelson III JR (2004) Taxonomy and Geographic Variation in the Leaf-Nosed Snake Phyllorhynchus

decurtatus (Squamata Colubridae) Journal of Herpetology 38 187ndash196 httpdxdoiorg10167081-03a

Gaucher P Dewynter M amp Born M (2008) Reptile Checklist of Nouragues area Centre National de la Recherche

Scientifique Available from httpwwwnouraguescnrsfrreptilehtml (Accessed 28 Apr 2010)Giesbrecht RM (2009) Estaccedilotildees Ferroviaacuterias do Brasil Available from httpwwwestacoesferroviariascombrindexhtml

(Date of access 18 August 2010)Gomes JF (1918) Contribuiccedilatildeo para o conhecimento dos ofiacutedios do Brasil ndash III (1) Memoacuterias do Instituto Butantan 1 57ndash83Grazziotin FG Monzel M Echeverrigaray S amp Bonatto SL (2006) Phylogeography of the Bothrops jararaca complex

(Serpentes Viperidae) past fragmentation and island colonization in the Brazilian Atlantic Forest Molecular Ecology 15 3969ndash3982 httpdxdoiorg101111j1365-294x200603057x

Griffin LE (1916) A catalogue of the Ophidia from South America at present (June 1916) in the Carnegie Museum with descriptions of some new species Memoirs of the Carnegie Museum 7 163ndash228

Guimaratildees MR Bovo RP Kasperoviczus KN amp Marques OAV (2010) Bothrops insularis (Golden Lancehead) Maximum length Herpetological Review 41 89

Guumlnther A (1868) Sixth account of new species of snakes in the collection of the British Museum Annals and Magazine of

Natural History 4 (1) 413ndash429 httpdxdoiorg10108000222936808695725

Harvey MB (1998) Reptiles and Amphibians of Parque Nacional Noel Kempff Mercado In Killeen TJ amp TS Schulenberg (Eds) RAP Working Papers 10 A Biological Assessment of Parque Nacional Noel Kempff Mercado Conservation International Washington DC pp 144ndash166

Hoge AR Belluomini HE amp Fernandes W (1977) Variaccedilatildeo do nuacutemero de placas ventrals de Bothrops jararaca em funccedilatildeo dos climas (Viperidae Crotalinae) Memoacuterias do Instituto Butantan 4041 11ndash17

Humphries JM Bookstein FL Chernoff B Smith GR Elder RL amp Poss SG (1981) Multivariate discrimination by shape in relation to size Systematic Zoology 30 291ndash308 httpdxdoiorg1023072413251

Icochea J Quispitupac E Portilla A amp Ponce E (2001) Assessment of Amphibians and Reptiles of the Lower Urubamba Region Peru In Alonso A Dallmier F amp Campbell P (Eds) Urubamba The Biodiversity of a Peruvian Rainforest Smithsonian Institution Washington DC pp 129ndash142

Jackson JF (1978) Differentiation in the genera Enyalius and Strobilurus (Iguanidae) implications for Pleistocene climatic changes in eastern Brazil Arquivos de Zoologia 30 1ndash79

James FC amp McCulloch CE (1990) Multivariate Analysis in Ecology and Systematics Panacea or Pandorarsquos Box Annual

Review of Ecology and Systematics 21 129ndash166 httpdxdoiorg101146annureves21110190001021


Johnson RA amp Wichern DW (1998) Applied multivariate statistical analysis Prentice-Hall New Jersey 816 ppKnapp S amp Nee M (2009) Solanum anomalostemon (Solanaceae) an Endangered New Species from Southern Peru with

Unusual Anther Morphology Novon A Journal for Botanical Nomenclature 19 178ndash181 httpdxdoiorg1034172007108

Ledru MP (1993) Late quaternary environmental and climatic changes in central Brazil Quaternary Research 39 90ndash98 httpdxdoiorg101006qres19931011

Lehr E Carrillo N amp Hocking P (2004) A new species of Drymoluber (Reptilia Squamata Colubridae) from southeastern Peru Copeia 2004 46ndash52 httpdxdoiorg101643ci-02-248r1

Lehr E amp Catenazzi A (2008) A new species of Bryophryne (Anura Strabomantidae) from souther Peru Zootaxa 1784 1ndash10

Leite PT Nunes SF Kaefer IL amp Cechin SZ (2009) Reproductive biology of the swamp racer Mastigodryas bifossatus

(Serpentes Colubridae) in subtropical Brazil Zoologia 26 12ndash18 httpdxdoiorg101590s1984-46702009000100003

Loumlfgren A (1902) In Freyreiss GW Viagem a varias tribus de selvagens na Capitania de Minas Gerais permanencia entre ellas descripccedilatildeo de seus usos e costumes Revista do Instituto Histoacuterico e Geographico de Satildeo Paulo 6 pp 236ndash237

Manier MK (2004) Geographic variation in the long-nosed snake Rhinocheilus lecontei (Colubridae) beyond the subspecies debate Biological Journal of the Linnean Society 83 65ndash85 httpdxdoiorg101111j1095-8312200400373x

Manly BFJ (2004) Multivariate Statistical Methods A Primer 2nd ed Chapman and HallCRC Boca Raton USA 208 ppMartins M amp Oliveira ME (1998) Natural History of snakes in forests of the Manaus Region Central Amazonia Brazil

Herpetological Natural History 6 78ndash150Martins M Marques OAV amp Sazima I (2008) How to be arboreal and diurnal and still stay alive microhabitat use time of

activity and defense in Neotropical forest snakes South American Journal of Herpetology 3 58ndash67 httpdxdoiorg1029941808-9798(2008)3[58HTBAAD]20CO2

McDiarmid R (1968) Variation Distribution and Systematic Status of the Black-Headed Snake Tantilla yaquia Smith Bulletin

of the Southern California Academy of Sciences 67 159ndash177Mendelson III JR (1992) Frequency of tail breakage in Coniophanes fissidens (Serpentes Colubridae) Herpetologica 48

448ndash455Montero R Scrocchi G Montantildeo ME amp Fernaacutendez IM (1995) Nuevas citas de saurios anfisbenidos y ofidios para

Bolivia Cuadernos de Herpetologia 9 7ndash13Moreira LA Fenolio DB Silva HLR amp Silva-Jr NJ (2009) A preliminary list of the herpetofauna from termite mounds

of the Cerrado in the upper Tocantins river valley Papeacuteis Avulsos de Zoologia 49 183ndash189Olson DM Dinerstein E Wikramanayake ED Burgess ND Powell GVN Underwood EC DrsquoAmico JA Itoua I

Strand HE Morrison JC Loucks CJ Allnutt TF Ricketts TH Kura Y Lamoreux JF Wettengel WW Hedao P amp Kassem KR (2001) Terrestrial ecoregions of the world A new map of life on Earth Bioscience 51 933ndash938 httpdxdoiorg1016410006-3568(2001)051[0933TEOTWA]20CO2

Osgood DW (1978) Effects of temperature on the development of meristic characters in Natrix fasciata Copeia 1978 33ndash37 httpdxdoiorg1023071443819

Papavero N (1971) Essays on Neotropical Dipterology Vol I Museu de Zoologia Universidade de Satildeo Paulo Satildeo Paulo 216 pp

Passos P amp Brandatildeo F (2002) Geographic Distribution Drymoluber dichrous Herpetological Review 33 324Passos P amp Fernandes R (2008) Revision of the Epicrates cenchria complex (Serpentes Boidae) Herpetological

Monographs 22 1ndash30 httpdxdoiorg10165506-0031

Passos P Fernandes R amp Porto M (2005) Geographical variation and taxonomy of the snail-eating snake Dipsas albifrons

(Sauvage 1884) with comments on the systematic status of Dipsas albifrons cavalheiroi Hoge 1950 (Serpentes Colubridae Dipsadinae) Zootaxa 1013 19ndash34

Pavan D amp Dixo M (2004) A Herpetofauna da aacuterea de influecircncia do reservatoacuterio da Usina Hidreleacutetrica Luiacutes Eduardo Magalhatildees Palmas TO Humanitas 46 13ndash30

Paynter Jr RA (1982) Ornithological Gazetteer of Venezuela Museum of Comparative Zoology Harvard University Cambridge Massachusetts iv + 245 pp

Paynter Jr RA (1992) Ornithological Gazetteer of Bolivia (Second Edition) Museum of Comparative Zoology Harvard University Cambridge Massachusetts vi + 185 pp

Paynter Jr RA (1993) Ornithological Gazetteer of Ecuador (Second Edition) Museum of Comparative Zoology Harvard University Cambridge Massachusetts xi + 247 pp

Paynter Jr RA (1997) Ornithological Gazetteer of Colombia (Second Edition) Museum of Comparative Zoology Harvard University Cambridge Massachusetts ix + 537 pp

Paynter Jr RA amp Traylor Jr MA (1991) Ornithological Gazetteer of Brazil (2 vols) Museum of Comparative Zoology Harvard University Cambridge Massachusetts viii + 789 pp


Peracca MG (1897) Viagio del Dr Enrico Festa nellEcuador e regioni vicine Bolletino dei Musei di Zoologia ed Anatomia comparata dela R Universitagrave di Torino 300 1ndash20

Pereira DN Stender-Oliveira F Rodrigues MG amp Beacuternils RS (2007) Distribution and habitat use of Sordellina punctata

(Serpentes Colubridae) with a new record from State of Satildeo Paulo Brazil Herpetological Bulletin 100 18ndash22Pesantes OS (1994) A method for preparing hemipenis of preserved snakes Journal of Herpetology 28 93ndash95

httpdxdoiorg1023071564686Peters W (1863) Uumlber einige neue oder weniger bekannte Schlangenarten des zoologischen Museums zu Berlin

Monatsberichte der koumlniglich Akademie der Wissenschaften zu Berlin 29 272ndash289Peters JA (1964) Dictionary of Herpetology Hafner New York 393 ppPeters JA amp Orejas-Miranda B (1970) Catalogue of the Neotropical Squamata Part I Snakes United States National

Museum Bulletin 297 1ndash347Pinto RR (2006) Drymoluber dichrous (NCN) Ophiophagy Herpetological Review 37 231Pizzatto L Almeida-Santos SM amp Marques OAV (2007) Biologia reprodutiva de serpentes brasileiras In Nascimento

LB amp Oliveira ME (Eds) Herpetologia no Brasil II Sociedade Brasileira de Herpetologia Belo Horizonte pp 201ndash221

Prudente ALC Maschio GF Yamashina CE amp Santos-Costa MC (2007) Morphology Reproductive Biology and Diet of Dendrophidion dendrophis (Schlegel 1837) (Serpentes Colubridae) in Brazilian Amazon South American Journal of

Herpetology 2 53ndash58 httpdxdoiorg1029941808-9798(2007)2[53mrbado]20co2

Reis SF Cruz JF amp Von Zuben CJ (1988) Anaacutelise multivariada da evoluccedilatildeo craniana em roedores caviiacutedeos convergecircncia de trajetoacuterias ontogeneacuteticas Revista Brasileira de Geneacutetica 11 633ndash641

Ribon R (1995) Nova subespeacutecie de Caprimulgus (Linnaeus) (Aves Caprimulgidae) do Espiacuterito Santo Brasil Revista

Brasileira de Zoologia 12 333ndash337 httpdxdoiorg101590s0101-81751995000200011

Rocha L (1972 ldquo1973rdquo) Wied Freyreiss e Sellow no Espiacuterito Santo Revista do Instituto Histoacuterico e Geograacutefico Brasileiro 297 56ndash67

Rodrigues MT (1990) Os lagartos da Floresta Atlacircntica brasileira distribuiccedilatildeo atual e preteacuterita e suas implicaccedilotildees para estudos futuros II Simpoacutesio sobre ecossistemas da costa sul brasileira Estrutura manejo e funccedilatildeo Academia de Ciecircncias do Estado de Satildeo Paulo Satildeo Paulo pp 404ndash410

Rodrigues MT (1996) Lizards Snakes and Amphisbaenians from the Quartenary Sand Dunes of the Middle Rio Satildeo Francisco Bahia Brazil Journal of Herpetology 30 513ndash523 httpdxdoiorg1023071565694

Rodrigues MT (2003) Herpetofauna da Caatinga In Leal IR Tabarelli M amp Silva JMC (Orgs) Ecologia e

Conservaccedilatildeo da Caatinga 4 Universidade Federal de Pernambuco Recife pp 181ndash236Santana GG Vieira WLS Pereira-Filho GA Delfim FR Lima YCC amp Vieira KS (2008) Herpetofauna em um

fragmento de Floresta Atlacircntica no Estado da Paraiacuteba Regiatildeo Nordeste do Brasil Biotemas 71 75ndash84 httpdxdoiorg1050072175-79252008v21n1p75

Shine R Olsson MM Moore IT LeMaster MP amp Mason RT (1999) Why do male snakes have longer tales than females Proceedings of the Royal Society of London B 266 2147ndash2151 httpdxdoiorg101098rspb19990901

Silva MB Barbosa DBS Lima YCC Antunes JL Figueiredo FJ amp Nogueira-Paranhos JD (2007) Ocorrecircncia de Drymoluber cf brazili (Reptilia Squamata Colubridae) para o estado do Piauiacute Anais do VIII Congresso de Ecologia do

Brasil Caxambu - MG 1ndash2Silva Jr NJ Silva HLR Costa MC Buononato MA Tonial MLS Ribeiro RS Moreira LA amp Pessoa AM

(2007) Avaliaccedilatildeo preliminar da fauna silvestre terrestre do vale do rio Caiapoacute Goiaacutes Implicaccedilotildees para a conservaccedilatildeo da biodiversidade regional Estudos 34 1057ndash1094

Silveira AL Pires MRS amp Cotta GA (2010 ldquo2011rdquo) Serpentes de uma aacuterea de transiccedilatildeo entre o Cerrado e a Mata Atlacircntica no sudeste do Brasil Arquivos do Museu Nacional Rio de Janeiro 68 79ndash110

Slowinski JB amp Savage JM (1995) Urotomy in Scaphiodontophis evidence for the multiple tail break hypothesis in snakes Herpertologica 51 338ndash341

StatSoft Inc (2004) Statistica (data analysis software system) version 7 Tulsa USAStuart LC (1932) Studies on Neotropical Colubrinae I The taxonomic status of the genus Drymobius Fitzinger Occasional

Papers of the Museum of Zoology University of Michigan 236 1ndash16 Stuart LC (1933) Studies on Neotropical Colubrinae III The Taxonomic Status of Certain Neotropical Racers Copeia 1933

9ndash10Torres-Carvajal O (2001) Lizards of Ecuador CheckList Distribution and Systematic References Smithsonian

Herpetological information Service 131 1ndash35Urban DL (2003) Spatial analysis in ecology Mantelrsquos test National Center for Ecological Analysis and Synthesis Univ of

California Santa Barbara Available from httpwwwnceasucsbeduscicompdocuments (Accessed 29 Apr 2010)Vanzolini PE (1981) A quasi-historical approach of the natural history of the differentiation of reptiles in tropical geographic

isolates Papeacuteis Avulsos de Zoologia 24 189ndash204


Vanzolini PE Ramos-Costa AMM amp Vitt LJ (1980) Reacutepteis das Caatingas Anais da Academia Brasileira de Ciecircncias Rio de Janeiro 161 pp

Veriacutessimo D Freitas MM amp Rosa GM (2012) Drymoluber dichrous (northern woodland racer) and Anolis fuscoauratus

(slender anole) Predation Herpetological Bulletin 120 36ndash38Vervust B Van Dogen S amp Van Damme R (2009) The effect of preservation on lizard morphometrics ndash an experimental

study Amphibia Reptilia 30 321ndash329 httpdxdoiorg101163156853809788795209

Vitt LJ Souza RA Sartorius SS Aacutevila-Pires TCS amp Espoacutesito MC (2000) Comparative ecology of sympatric Gonatodes (Squamata Gekkonidae) in the western Amazon of Brazil Copeia 2000 83ndash95 httpdxdoiorg1016430045-8511(2000)2000[0083ceosgs]20co2

Vrcibradic D (2007) Geographic Distribution Drymoluber dichrous Herpetological Review 38 486Warrell DA Theakston RGD amp Wuumlster W (2010) Destruction of the collection of reptiles and arthropods at Butantan

Institute a view from the United Kingdom The Journal of Venomous Animals and Toxins including Tropical Diseases 16 534ndash536 httpdxdoiorg101590s1678-91992010000400003

Wied MP (1989) Viagem ao Brasil (Portuguese version of Reise nach Brasilien in den Jahren 1815 bis 1817 translated by Edgar Suumlssekind de Mendonccedila and Flavio Poppe de Figueiredo with notes by Oliveacuterio Pinto) Editora Itatiaia Editora da Universidade de Satildeo Paulo Belo Horizonte Satildeo Paulo Brazil 536 pp

Zaher H amp Prudente ALC (2003) Hemipenes of Siphlophis (Serpentes Xenodontinae) and Techniques of Hemipenal Preparation in Snakes A Response to Downling Herpetological Review 34 302ndash307

Zar JH (1999) Biostatistical Analysis Prentice Hall Upper Sadle River New Jersey USA 663 pp

Appendix I

Collections from which specimens of Drymoluber were personally examined for this study BRAZIL Coleccedilatildeo Herpetoloacutegica da Universidade de Brasiacutelia (CHUNB) Brasiacutelia DF Coleccedilatildeo Herpetoloacutegica da Universidade Federal de Minas Gerais (CHUFMG) Belo Horizonte MG Coleccedilatildeo Zooloacutegica Gregoacuterio Bondar (CZGB) Ilheacuteus BA Instituto Butantan (IBSP) Satildeo Paulo SP Laboratoacuterio de Zoologia de Vertebrados Universidade Federal de Ouro Preto (LZV) Ouro Preto MG Museu de Biologia Mello Leitatildeo (MBML) Santa Teresa ES Museu de Ciecircncias e Tecnologia PUC-RS(MCP) Porto Alegre RS Museu de Ciecircncias Naturais PUC-MG (MCNR) Belo Horizonte MG Museu de Zoologia da Universidade de Satildeo Paulo (MZUSP) Satildeo Paulo SP Museu de Zoologia da Universidade Estadual de Santa Cruz(MZUESC) Ilheacuteus BA Museu de Zoologia Joatildeo Moojen Universidade Federal de Viccedilosa (MZUFV) Viccedilosa MG Museu Nacional Universidade Federal do Rio de Janeiro (MNRJ) Rio de Janeiro RJ Museu Paraense Emiacutelio Goeldi(MPEG) Beleacutem PA Universidade Federal do Acre Campus Floresta (UFACF) Cruzeiro do Sul AC Universidade Federal do Cearaacute (UFC) Fortaleza CE Universidade Federal do Mato Grosso (UFMT) Cuiabaacute 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 DCCollections from which photographs of specimens of Drymoluber were examined for this study GERMANY Museum fuumlr Naturkunde Berlin (ZMB) Berlin Museum fuumlr Tierkunde (MTKD) Dresden PARAGUAY Museo Nacional de Historia Natural Del Paraguay (MNHNP) Asunciacuteon 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 dagger indicates specimens which hemipenis was also examined Geographic coordinates expressed in decimal degrees as ldquolatitude longituderdquo (eg 260 -7260 mean 260deg N 7260deg W)

Drymoluber apurimacensisPERU APURIacuteMAC Abancay Abancay (MHNSM 20672 [Holotype]) [-1364 -7288] Cconoc (MTKD 44669

45192 45193 [Paratypes]) [-1306 -7264] Andahuaylas Huancarama Hacienda Palmira (FMNH 81542 dagger [Paratype]) [-1365 -7338]

Drymoluber braziliBRASIL Without precise locality (IBSP 574 [Paratype of Drymobius brazili]) BAHIA Without precise locality


(IBSP 2717) Gentio do Ouro Santo Inaacutecio (MZUSP 9596) [-1110 -4273] Jacobina Caatinga do Moura (MZUSP 7544) [-1098 -4077] Joatildeo Dourado Gruta dos Brejotildees (MZUESC 3815) [-1100 -4125] Morro do Chapeacuteu (MZUSP 7807) [-1155 -4116] CEARAacute Milagres (IBSP 76968) [-731 -3895] DISTRITO FEDERAL Brasiacutelia (CHUNB 37345 CHUNB 3747 CHUNB 3748 CHUNB 6054) [-1578 -4793] ESPIacuteRITO SANTO Baixo Guanduacute (IBSP 8312 IBSP 8836 dagger) [-1952 -4102] Colatina (IBSP 37413) [-1954 -4063] GOIAacuteS Alto Paraiacuteso de Goiaacutes (CHUNB 3839) [-1413 -4751] Goiacircnia (IBSP 33668 dagger) [-1668 -4925] Mineiros Parque Nacional das Emas (IBSP 62682) [-1808 -5292] Satildeo Domingos Parque Estadual Terra Ronca (IBSP 62650) [-1365 -4635] Uruaccedilu Cana Brava (IBSP 26716 IBSP 9144) [-1453 -4914] Serra da Mesa (ponto 2) (MZUSP 11021) [-1390 -4831] MATO GROSSO Chapada dos Guimaratildees (UFMT 6970 dagger UFMT 728) [-1545 -5574] MATO GROSSO DO SUL Campo Grande (IBSP 10465 dagger IBSP 31692) [-2044 -5465] Miranda (IBSP 1845) [-2024 -5638] Ponta Poratilde (IBSP 25379 dagger) [-2254 -5573] Terenos (IBSP 10018 IBSP 9829) [-2044 -5486] Trecircs Lagoas (IBSP 51700) [-2075 -5168] Jupiaacute (IBSP 27199) [-2075 -5168] MINAS GERAIS Aimoreacutes (MCNR 1538) [-1950 -4106] Alvarenga (CHUFMG 2843) [-1941 -4172] Frutal (IBSP 52290) [-2003 -4894] Gratildeo Mogol (MZUSP 7988) [-1671 -4259] Nova Ponte (MZUFV 780) [-1914 -4768] Sabaraacute (IBSP 40475) [-1989 -4381] Santana do Riacho (CHUFMG 417) [-1911 -4368] Serra do Cipoacute (MZUSP 7692) [-1923 -4355] Uberaba (IBSP 1707) Uberaba Engenheiro Lisboa (IBSP 696 [Holotype of Drymobius brazili]) [-1980 -4760] UHE Irapeacute (MCNR 1736 dagger) [-1674 -4258] PARAIacuteBA Teixeira Serra do Teixeira (MZUSP 7562) [-720 -3725] SAtildeO PAULO Andradina (IBSP 40779) [-2090 -5138] Araccedilatuba (IBSP 26688) [-2121 -5043] Araraquara (IBSP 23693 IBSP 33120 IBSP 8103) [-2179 -4818] Barretos (IBSP 7675) [-2056 -4857] Birigui (IBSP 21928 IBSP 22415) [-2129 -5034] Boa Esperanccedila do Sul Estaccedilatildeo Ferroviaacuteria Java (IBSP 741 [Paratype of Drymobius brazili]) [-2199 -4839] Castilho (IBSP 24422) [-2087 -5149] Coroados (IBSP 17019) [-2135 -5028] Dourado (IBSP 18309) [-2210 -4832] Dracena (IBSP 25187) [-2148 -5153] Guaraccedilaiacute (IBSP 32897) [-2103 -5121] Guataparaacute Estaccedilatildeo Ferroviaacuteria de Monteiros (IBSP 12692) [-2150 -4804] Itaacutepolis (IBSP 18053) [-2160 -4881] Moreira Melo (IBSP 16595) [not found] Novo Horizonte Estaccedilatildeo Ferroviaacuteria Porto Ferratildeo (IBSP 22502 IBSP 22515) [-2147 -4922] Oswaldo Cruz (IBSP 32624 IBSP 32657) [-2180 -5089] Pedregulho (IBSP 1286 [Paratype of Drymobius brazili]) [-2026 -4748] Penaacutepolis (IBSP 1844 [Holotype of Drymobius rubriceps]) [-2142 -5008] Ribeiratildeo Bonito (IBSP 46996) [-2118 -4781] Estaccedilatildeo Ferroviaacuteria Sampaio Vidal (IBSP 573 [Paratype of Drymobius brazili]) [-2207 -4818] Ribeiratildeo Preto (IBSP 18992) [-2118 -4781] Santa Luacutecia Fazenda Santa Izabel (IBSP 34369) [-2169 -4808] Satildeo Carlos Estaccedilatildeo Ferroviaacuteria Santa Eudoacutexia (IBSP 383 [Paratype of Drymobius brazili]) [-2177 -4778] Satildeo Joaquim da Barra Estaccedilatildeo Ferroviaacuteria Jussara (IBSP 16231) [-2058 -4786] Satildeo Joseacute do Rio Preto Estaccedilatildeo Ferroviaacuteria Rio Preto (IBSP 10188 IBSP 8925) [-2082 -4938] Satildeo Simatildeo Fazenda Vale da Sauacutede (IBSP 33660) [-2148 -4755] Turiba do Sul (IBSP 10538 IBSP 16499) [-2375 -4928] Uracircnia (IBSP 17224) [-2025 -5064] Valparaiacuteso (IBSP 29221) [-2123 -5087] TOCANTINS UHE Luiacutes Eduado Magalhatildees (MZUSP 14298) [-975 -4840] UHE Peixe Angical (MZUSP 15506 MZUSP 15507) [-1240 -4825] PARAGUAICANINDEYUacute Reserva Natural del Bosque Mbaracayuacute (MNHNP 11025) [-2414 -5532]

Drymoluber dichrousBOLIVIA BENI Riberalta (AMNH 22491) [-1098 -6610] Rurrenabaque (AMNH 119926) [-1444 -6753]

Vaca Diez Tumi Chucua (USNM 280382) [-1115 -6617] 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) [-933 -7042] Rio Juruaacute (MPEG 20384 dagger) [-827 -7274] Rio Branco (IBSP 69567) [-997 -6780] Santa Rosa do Purus Alto Purus (MZUSP 2497) [-933 -7042] Tarauacaacute Reserva Extrativista Riozinho da Liberdade (UFACF 1333 UFACF 663 UFACF 720 UFACF 741 UFACF 801) [-800 -7200] AMAPAacute Macapaacute Zebratildeo (MNRJ 9240) [004 -5106] Serra do Navio (IBSP 25411) [090 -5200] Tartarugalzinho Reserva DNERu Rio Tracajatuba (MPEG 421) [100 -5100] Rio Maracaacute (MZUSP 7681) [-043 -5143] AMAZONAS Without precise locality (IBSP 7706) Arajatuba Rio Negro (IBSP 2198) [-310 -6049] Barcelos Rio Negro MZUSP 5467 [-097 -6293] Benjamin Constant Proacuteximo a igarapeacute (MPEG 18240 dagger) [-447 -7003] Madeirera SCHEFFER Rio Ituxi (MPEG 20330 MPEG 20331) [-820 -6542] Presidente Figueiredo UHE Balbina (IBSP 51703 IBSP 52003 IBSP 52140 dagger MPEG 17437 MPEG 17443 MPEG 17446 MPEG 17481 MPEG 17494 MPEG 17566 MZUSP 9644 MZUSP 9645) [-154 -5984] Reserva INPA-WWF (MZUSP 7609) [-242 -5972] Serra do Tapirapecoacute (MZUSP 14284 MZUSP 14285 MZUSP 14286) [120 -6480] BAHIA Without precise locality(MZUESC 5801) Arataca (CZGB 396) [-1526 -3941] Fazenda Alto Rocha (MZUESC 1528) [-1515 -3930] Barra do Choccedila (CZGB 7981 CZGB 7396) [-1486 -4056] Fazenda Cangussu (MZUESC 2540) [-1496 -4065] Barra do Rocha Fazenda Pedra Preta (MZUESC 1993) [-1406 -3960] Barro Preto (CZGB 822) [-1477 -3940] Buerarema (CZGB 339 CZGB 4441) [-1495 -3930] Caatiba (CZGB 7059) [-1498 -4041] Cairu Fazenda Ilha do Barro (MZUESC 636) [-1355 -3906] Camacan (CZGB 846) [-1542 -3950] Fazenda Uraiccedilu (MZUESC 5162) [-1539 -3956] Camamu (CZGB 5180) [-1395 -3910] Dario Meira (CZGB 3164 CZGB 3342) [-1444 -3991] Ibirataia (CZGB 7397) [-1407 -3964] Ilheacuteus (CZGB 1222 CZGB 1269 CZGB 3176 CZGB 666 dagger) [-1478 -3911 Itabuna (CZGB 324) [-1479 -3928] Itacareacute (CZGB 2101) [-1428 -3900] Itagi (MZUESC 5697 dagger) [-1407 -3995 Itagiba (CZGB 5430) [-1428 -3984 Ituberaacute (CZGB 5609 CZGB 6825 CZGB 6826) [-1373 -3915] Mutuiacutepe (CZGB 7717


dagger) [-1323 -3951] Nova Ibiaacute (CZGB 6809 CZGB 7385) [-1383 -3959] Poccedilotildees Fazenda Inveja (MZUESC 1383) [-1463 -4027] Una (CZGB 331) [-1527 -3907] Fazenda Ararauacutena (MZUESC 3739) [-1527 -3907] CEARAacuteMaranguape Serra de Maranguape (UFC 2101 UFC 2201 UFC 2204 UFC 2205 UFC 2211 UFC 2220 UFC 2712 UFC 2730) [-390 -3872] Trilha do Pico da Rajada Serra de Maranguape (UFC 2081 UFC 2083 dagger UFC 2104 UFC 2128) [-390 -3872] Pacatuba Serra do Aratanha (UFC 2233) [-397 -3862] Pacoti Serra do Baturiteacute (UFC 2221) [-423 -3892] Ubajara Planalto da Ibiapaba Fazenda Buriti-INCRA (UFC 2177 dagger) [-384 -4090] ESPIacuteRITO SANTOAracruz (MBML 483) [-1982 -4027] Linhares (CZGB 2121 dagger MNRJ 10661 MNRJ 10663) [-1939 -4007] Vitoacuteria (MNRJ 4858) [-2032 -4034] MARANHAtildeO BR 316 25 Km distante do Rio Gurupi Nova Vida (MPEG 11099 MPEG 11155 MPEG 12106 MPEG 12228 dagger MPEG 12229 MPEG 12230 MPEG 12257 MPEG 14466 MPEG 15321 MPEG 16207) [-200 -4599] Buriticupu (MPEG 17713) [-435 -4640] Santa Luzia do Paruaacute BR 316 (MPEG 10832) [-263 -4577] MATO GROSSO Alta Floresta (IBSP 41469) [-988 -5609] Aripuanatilde (UFMT 5184 UFMT 5474) [-1015 -5945] Ilha do Salto Dardanelos (UFMT 2772) [-1015 -5945] Juruena (MZUSP 11303) [-1032 -5836] Rio Teles Pires Paranaacute do Cristoacutevatildeo (MZUSP 10983) [-755 -5797] Vale do Satildeo Domingos (UFMT 1997 dagger UFMT 919) [-1529 -5906] Vila Rica (MZUSP 11443 MZUSP 11444) [-1001 -5112] MINAS GERAIS Almenara Fazenda Limoeiro (MZUFV 1208 dagger) [-1601 -4050] Caratinga RPPN Feliciano Miguel Abdala (Estaccedilatildeo Bioloacutegica de Caratinga) (CHUFMG 1397 CHUFMG 1398) [-1971 -4181] Nova Ponte (MZUFV 779 dagger) [-1914 -4768] Ouro Preto Estaccedilatildeo Ecoloacutegica do Tripuiacute (LZV-S 373) [-2040 -4358] Parque Estadual do Rio Doce (MNRJ 9297 MZUFV 1422) [-1943 -4245] PARAacute Almerim Reserva Itapioara (MPEG 20108) [-153 -5256] Beleacutem (IBSP 3035 IBSP 3122 dagger IBSP 3141 IBSP 5089) [-146 -4850] Benevides Santa Baacuterbara (MPEG 2647) [-136 -4825] PratinhaEstrada de Genipauba antiga estrada do Accedilucareiro (MPEG 8400 MPEG 8607) [-136 -4825] Canindeacute Rio Gurupi (MZUSP 4237) [-255 -4652] Castanhal Boa Vista (MPEG 2652 MPEG 2670 MPEG 4041) [-136 -4799] Conceiccedilatildeo do Araguaia (IBSP 41522) [-826 -4926] Juruaacute Rio Xingu (MZUSP 9351 MZUSP 9352) [-340 -5188] Marabaacute (CHUNB 30419 MPEG 23235) [-537 -4912] Serra Norte Carajaacutes (MPEG 17233 MPEG 17235) [-585 -5020] Serra Norte Carajaacutes Aacuterea do Caldeiratildeo (MPEG 16921) [-585 -5020] Serra Norte Carajaacutes Campo rupestre do N1 (MPEG 16551 MPEG 16982) [-585 -5020] Serra Norte Carajaacutes Proacuteximo as margens do Igarapeacute Azul Estrada N1 Caldeiratildeo (MPEG 17057) [-585 -5020] Serra Norte Carajaacutes Aacuterea do Pojuca (MPEG 17085) [-585 -5020] Melgaccedilo FLONA Caxiuanatilde (MPEG 19750 MPEG 20044 MPEG 20146 MPEG 20298 MPEG 20800 MPEG 21833 MPEG 21834) [-180 -5150] Monte Cristo Rio Tapajoacutes (MZUSP 1264) [-407 -5565] Novo Progresso (CHUNB 35062) [-715 -5538] Ourilacircndia do Norte Acampamento Onccedila (MPEG 20793) [-715 -5100] Paragominas Fazenda Agroeste (MPEG 20000) [-300 -4735] Peixe-Boi (MPEG 1838) [-120 -4732] Projeto Igarapeacute Bahia Carajaacutes (MPEG 20762 MPEG 20765 MPEG 20766) [-605 -5010] Reserva Biologica Rio Trombetas Igarape Jacare above Lago Jacareacute (USNM 289091) [-119 -5667] Santareacutem (MCP 7617 MCP 7618 MCP 7906) [-244 -5471] Satildeo Domingos do Capim Km 16 da estrada do Acaraacute (MPEG 10811 MPEG 11698 MPEG 15189 MPEG 8688) [-168 -4777] Serra de Kukoinhokren (MZUSP 10763 MZUSP 11488) [-777 -5195] Tucuruiacute (IBSP 46626) [-383 -4969] PERNAMBUCO Cabo de Santo Agostinho Mata do Cutio Reserva Florestal do sistema Gursauacute (MNRJ 17069) [-827 -3506] RONDOcircNIA Alto Paraiacuteso (MZUSP 8494) [-962 -6345] Jaru Santa Cruz da Serra (MZUSP 8500) [-1067 -6257] Ministro Andreazza Nova Brasiacutelia (MZUSP 8747) [-1115 -6157] Monte Negro (IBSP 67619) [-1024 -6329] Porto Velho UHE Samuel (IBSP 52828 IBSP 53256) [-886 -6343] Margem esquerda do Rio Jamariacute Aacuterea de inundaccedilatildeo da UHE Samuel (MPEG 17779 MPEG 17808 MPEG 17820 MPEG 17829 MPEG 17841 MPEG 17849) [-886 -6343] Margem do Rio Jamariacute Aacuterea de inundaccedilatildeo da UHE Samuel (MPEG 17897 MPEG 17915 MPEG 17932 MPEG 17995 MPEG 17996 MPEG 18759) [-886 -6343] Vilhena (CHUNB 12791) [-1274 -6015] RORAIMA Amajari Ilha de Maracaacute (MZUSP 8806) [342 -6167] BR 210 4Km do Rio Ajarani (MPEG 19007) [192 -6117] Caracaraiacute Missatildeo Catrimani (MZUSP 10298 MZUSP 10422 MZUSP 10423 MZUSP 10424) [168 -6228] Mucajaiacute Apiauacute (MZUSP 9777) [263 -6125] Cachoeira do Cujubim Rio Catrimani (MZUSP 6388 MZUSP 6389 MZUSP 6971) [175 -6228] COLOMBIA AMAZONAS Leticia (AMNH 91812) [-417 -6995] GUAVIARE Cantildeo Agua Bonita (FMNH 75688) [260 -7260] META Bellavista acima de Villavicencio (MZUSP 6098) [425 -7367] Villavicencio (IBSP 6208 IBSP 7221) [416 -7364] Finca El Buque (MZUSP 8077) [415 -7362] ECUADOR Without precise locality Napo or Maranon (ANSP 3920 [Holotype of Spilotes piceus]) CHIMBORAZO Riobamba (AMNH 23248) [-167 -7863] MORONA-SANTIAGO Arapicos Riacuteo Llhushin (USNM 204133) [-185 -7795] Macas (USNM 65475 AMNH 28830) [-232 -7812] Sucua (USNM 283952) [-247 -7817] Chiguaza + Macuma (USNM 204132) [-210 -7787] Riobamba Chamala Normandia (in the vicinities of Macas according to Savage 1960 Misc Pub Mus Zool Univ Mich 112) (AMNH 35927) [-231 -7812] NAPO Cabeceras del Rio Arajuno tributary of Rio Napo (USNM 204126) [-140 -7788] Loreto (USNM 204125) [-067 -7733] PASTAZA Andoas Rio Pastaza (AMNH 49074) [-257 -7663] Rio Conambo (FMNH 206028) [-212 -7605] Riacuteo Liguino (USNM 204127) [-148 -7737] Cabeceras Del Riacuteo Bobonaza (USNM 204128) [-147 -7788] Riacuteo Bufeo tributary of Riacuteo Bobonaza (USNM 204129) [-233 -7667] Riacuteo Conambo near mouth of Riacuteo Romarizo (USNM 204130) [-187 -7678] Paracachi Rio Curaray (USNM 204131) [-160 -7635 GUYANA CUYUNI-MAZARUNI Kartabo (AMNH 18160) [-635 -5868] East bank of Waruma River Camp 3 (USNM 549327) [534 -6077] EAST BERBICEca 18 mi SW Kwakwani ca 2 mi downriver from confluence of Berbice River and Kurudini River Berbice River camp


(USNM 566261) [509 -5824] PERU AMAZONAS Kayamas Riacuteo Cenepa (USNM 560425) [-447 -7817] San Antonio Riacuteo Cenepa (USNM 316601) [450 -7817] Huampami across the Rio Cenepa from Yusa Patagkamu (USNM 316602) [-447 -7817] HUAacuteNUCO Tingo Maria (USNM 193808 USNM 193813) [-930 -7598] Universidad Agraria de la Selva Rio Huallaga (USNM 193807 USNM 193812) [-930 -7598] JUNIN Perene (AMNH 23354 AMNH 23374) [-1095 -7523] Tarma Chanchamayo (FMNH 45593) [-1140 -7570] LORETO Contamana Suhuaya Isla (AMNH 52295) [-734 -7506] Iquitos (AMNH 52318 AMNH 52644 USNM 197267) [-376 -7326] Riacuteo 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 dagger AMNH 55193 AMNH 55276) [-377 -7325] Nauta Rio Samiyia Santa Elena (FMNH 109807) [-453 -7345] Pampa Hermosa Riacuteo Cushabatay Riacuteo Ucayali (AMNH 55359) [-721 -7526] Riacuteo Cushabatay (AMNH 55963) [-723 -7530] Pebas (BMNH 194611461 formerly 186791728 [Holotype of Herpetodryas occipitalis]) [-318 -7178] Requena Monte Carmelo (AMNH 55609 AMNH 55628 AMNH 55635) [-499 -7396] Yagua Indian Village Riacuteo Loretoyacu (AMNH 114706) [-382 -7043] Yurimagua (FMNH 11259 dagger) [-590 -7611] Estiroacuten Rio Ampiyacu (MZUSP 4390 MZUSP 4391) [-312 -7192] Riacuteo Samiria and Parinari Cantildeon (AMNH 57258) [-470 -7430] Upper Ucayali River (AMNH 71115) [-450 -7345] Upper Amazon (FMNH 11177) [-634 -7517] Yarinacocha (FMNH 45592) [-552 -7437] MADRE DE DIOSca 30 Km Puerto Maldonado Tambopata Reserve Explorers Inn (USNM 247501 USNM 247500 USNM 247694) [-1283 -6928] La Pampa (FMNH 40206 dagger) [Toponym not found] Maldonado (AMNH 56148) [-1260 -6918] West bank of Riacuteo Tambopata Zona Reservada Tambopata-Candamo Colpa de Guacamayo (USNM 332470) [-1314 -6961] SURINAME Without precise locality data (ZMB 2603 [syntype of Herpetodryas dichroa]) BROKOPONDOBrownsberg Brownsberg Nature Park Mazaroni Plateau (AMNH 119431) [495 -5517] COMMEWIJNE Plantation Ma Retraite (AMNH 130505) [574 -5487] MAROWIJNE Moengo (USNM 64634) [562 -5440] Anapaike Village (MZUSP 4207) [333 -5407] SIPALIWINI Temomairem Sierra Tumuc Humac (AMNH 104610) [300 -5538] VENEZUELA AMAZONAS Cerro Ya-Pacana (USNM 83946) [375 -6680] Mount Duida region (AMNH 36610) [348 -6558] Middle camp Mount Duida region (AMNH 36614) [338 -6594] Esmeralda 56 km NW of Rio Cunucunuma Belen (USNM 217185) [365 -6577]

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 braziliBRASIL MINAS GERAIS Mariana [-2038 -4342 Silveira et al 2010 ldquo2011rdquo] GOIAacuteS IporaacuteArenoacutepolis

Pequena Central Hidreleacutetrica de Mosquetatildeo [-1635 -5143 Silva-Jr et al 2007] PIAUIacute Parque Nacional da Serra das Confusotildees [-945 -4386 Silva et al 2007] Satildeo Gonccedilalo do Gurgueacuteia [-1004 -4529 Freitas et a 2012]

Drymoluber dichrousBOLIacuteVIA BENI Parque Nacional y Territorio Indiacutegena Isiboro Seacutecure [-1638 -6604 Embert 2007] Reserva de

la Bioacutesfera Estacioacuten Bioloacutegica del Beni [-1466 -6633 Embert 2007] Reserva de la Bioacutesfera y Tierra Comunitaria de Origen Piloacuten Lajas [-1500 -6731 Embert 2007] COCHABAMBA Parque Nacional Carrasco [-1750 -6500 Embert 2007] Santuario de Vida Silvestre Cavernas del Repechoacuten [-1738 -6606 Embert 2007] Villa Tunari [-1697 -6543 Lehr et al 2004] LA PAZ Aacuterea Natural de Manejo Integrado Apolobamba [-1500 -6900 Embert 2007] Parque Nacional y Aacuterea Natural de Manejo Integrado Cotapata [-1617 -6800 Embert 2007] Parque Nacional y Aacuterea Natural de Manejo Integrado Madidi [-1385 -6857 Embert 2007] SANTA CRUZ Guarayos Campamento Riacuteo San Martin [-1505 -6193 Montero et al 1995] Parque Nacional Noel Kempff Mercado [-1480 -6064 Harvey 1998 Embert 2007] Parque Nacional y Aacuterea Natural de Manejo Integrado Amboroacute [-1784 -6408 Embert 2007] PANDO Reserva Nacional de Vida Silvestre Amazoacutenica Manuripi [-1169 -6761 Embert 2007] BRASIL ALAGOAS Maceioacute [-967 -3572 Freire 2000] Rio Largo [-950 -3583 Freire 2000] AMAZONAS Manaus Reserva Adolpho Ducke [-298 -5993 Martins amp Oliveira 1998] CEARAacute Missatildeo Velha Santo Antocircnio water spring [-741 -3921 Veriacutessimo et al 2012] PARAacute Agropecuaacuteria Treviso [-315 -5483 Aacutevila-Pires et al 2009] Braganccedila Parada Bom Jesus [-107 -4679 Cunha amp Nascimento 1978] Colocircnia Nova Km 264 BR 316 [-180 -4648 Cunha amp Nascimento 1978] Curuccedilaacute Vila Marauaacute [-075 -4787 Cunha amp Nascimento 1978] Maracanatilde Km 23 da estrada de Maracanatilde [-078 -4746 Cunha amp Nascimento 1978] Oureacutem Limatildeo Grande [-155 -4711 Cunha amp Nascimento 1978] Santo Antocircnio do Tauaacute PA-140 estrada da Vigia [-111 -4813 Cunha amp Nascimento 1978] Vigia Santa Rosa PA-140 estrada da Vigia [-097 -4808 Cunha amp Nascimento 1978] Viseu Bela Vista [-121 -4619 Cunha amp Nascimento 1978] Fazenda Real [-127 -4622 Cunha amp Nascimento 1978] PARAIacuteBA Joatildeo Pessoa Aacuterea de Preservaccedilatildeo Permanente Mata do Buraquinho [-715 -3487 Santana et al 2008] RIO DE JANEIRO Campos dos Goytacazes [-2184 -4167 photograph of a juvenile specimen taken by Carlos Henrique Oliveira Nogueira] Rondocircnia Espigatildeo DrsquoOeste [-1139 -6074 Bernarde amp Abe


2006] Parque Estadual Guajaraacute-Mirim [-1032 -6455 Avila-Pires et al 2009] COLOcircMBIA BOYACAacute 8 Km SSE Corocito [553 -7306 Lehr et al 2004] EQUADOR AZUAY San Joseacute Cuchipamba [-302 -7875 Peracca 1897] MORONA-SANTIAGO Gualaquiza [-340 -7858 Peracca 1897] NAPO Alintildeahui [-107 -7760 Lehr et al 2004] ORELLANA Estaciacuteon de Biodiversidad Tiputini [-062 -7617 Cisneros-Heria 2003] PASTAZA Shell Mera [-151 -7806 Lehr et al 2004] SUCUMBIOS Santa Cecilia [005 -7596 Duellman 1978 Lehr et al 2004] Puerto Libre Riacuteo Aguarico [005 -7748 Duellman 1978 Lehr et al 2004] FRENCH GUIANA CAYENNE Nouragues Rserve [407 -5273 Gaucher et al 2008] PERU CUSCO Cashiriari-2 Lower Urubamba Region [-1186 -7278 Icochea et al2001] HUAacuteNACO Ambo [-1007 -7724 Lehr et al 2004] Huancapallac [-990 -7639 Lehr et al 2004] Rio Llullapichis Panguana [-962 -7493 Lehr et al 2004] JUNIacuteN Chanchamayo [-1104 -7532 Lehr et al 2004] LORETO Iquitos Centro Unioacuten [-378 -7312 Dixon amp Soini 1986] Mishana [-401 -7358 Dixon amp Soini 1986] Moropon [-372 -7333 Dixon amp Soini 1986] Riacuteo Maniti [-354 -7270 Dixon amp Soini 1986] Sarayacu [-673 -7510 Boulenger 1894] Yurimagua Rio Huallaga [-591 -7609 Boulenger 1894] PASCO Pozuzo [-1015 -7556 Lehr et al 2004] Iscozacin (Bosque de Villa America) [Toponym not found Lehr et al 2004] SAN MARTIN Moyobamba [-605 -7694 Boulenger 1894] UCAYALI Yarinacocha [-833 -7462 Lehr et al 2004]


Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III

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false PDFXNoTrimBoxError true PDFXTrimBoxToMediaBoxOffset [ 000000 000000 000000 000000 ] PDFXSetBleedBoxToMediaBox true PDFXBleedBoxToTrimBoxOffset [ 000000 000000 000000 000000 ] PDFXOutputIntentProfile () PDFXOutputConditionIdentifier () PDFXOutputCondition () PDFXRegistryName () PDFXTrapped False Description ltlt CHS ltFEFF4f7f75288fd94e9b8bbe5b9a521b5efa7684002000500044004600206587686353ef901a8fc7684c976262535370673a548c002000700072006f006f00660065007200208fdb884c9ad88d2891cf62535370300260a853ef4ee54f7f75280020004100630072006f0062006100740020548c002000410064006f00620065002000520065006100640065007200200035002e003000204ee553ca66f49ad87248672c676562535f00521b5efa768400200050004400460020658768633002gt CHT ltFEFF4f7f752890194e9b8a2d7f6e5efa7acb7684002000410064006f006200650020005000440046002065874ef653ef5728684c9762537088686a5f548c002000700072006f006f00660065007200204e0a73725f979ad854c18cea7684521753706548679c300260a853ef4ee54f7f75280020004100630072006f0062006100740020548c002000410064006f00620065002000520065006100640065007200200035002e003000204ee553ca66f49ad87248672c4f86958b555f5df25efa7acb76840020005000440046002065874ef63002gt DAN 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 DEU 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 ESP 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FRA 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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 ldquosupposedly bought in Surinamrdquo Five years later Cope (1868)

and Guumlnther (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 ldquoElaphe dichroardquo 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 ldquoElaphe dichrousrdquo 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 Penaacutepolis state of Satildeo

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 amp 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 Apuriacutemac 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 (eg Borges-Nojosa amp Lima 2001

Passos amp Brandatildeo 2002 Santana et al 2008) Amazonian savannas (Franccedila et al 2006) Caatinga (Vanzolini 1981

Rodrigues 2003 Argocirclo 2004b) Cerrado (Pavan amp Dixo 2004 Franccedila amp Arauacutejo 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 (eg 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


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)

























































1015 -5945)













































































Passos et al 2005)




Sexual dimorphism













































0434 p lt 0001) (Figs 7 and 8)





VE 0812 0188 -0511 0967 0102 NS -0225

SC 0557 -0422 0702 0864 -0298 0403275

ILGl -0084 -0023 0039 -0721 -0090 NS 0125

ILGr -0079 -0022 0043 -0737 -0094 NS 0151

MD 0063 0015 -0010 0898 0100 NS -0055 NS

ATl 0061 0595 0285 0176 0777 0302

PTr 0055 0126 -0072 0304 0315 -0147

ATr 0049 0633 0384 0137 0796 0393

PTl 0023 0109 -0075 0137 0287 -0159

SLOl 0023 0014 0007 0399 0116 NS 0049 NS

SLOr 0022 0005 -0005 0378 0039 NS -0033 NS

ILC1r 0016 -0005 -0004 0396 -0058 NS -0037 NS

ILC1l 0015 0005 -0007 0350 0056 NS -0063 NS

LO -0007 -0001 -0008 -0223 -0016 -0090 NS

ILC2r 0005 -0019 0006 0065 NS -0115 NS 0032 NS

ILr 0004 0005 0006 0149 0084 NS 0078 NS

ILl 0003 0008 00009 0119 NS 0116 NS 0010 NS

SLl 00009 0005 0006 0048 NS 0125 NS 0118 NS

ILC2l -00006 0002 -0006 -0007 NS 0013 NS -0028 NS

SLr 00005 00004 -0003 0025 NS 0010 NS -0062 NS




VE 0746 -0468 0439 0950 -0242 0190

SC 0626 0720 -0288 0896 0419 -0140 NS

ATr 0132 -0378 -0724 0392 -0454 -0728

ATl 0100 -0270 -0420 0367 -0402 -0523

ILGl -0083 0060 0009 -0757 0223 0029 NS

ILGr -0072 0070 0013 -0688 0272 0044 NS

MD 0065 -0024 -0001 0904 -0137 NS -0007 NS






TABLE 2 (Continued)


PTr 0043 -0124 -0077 0267 -0315 -0163

PTl 0038 -0135 -0120 0256 -0366 -0272

SLOl 0032 -0036 0029 0426 -0193 0132 NS

SLOr 0031 -0015 0010 0453 -0092 NS 0051 NS

LO -0019 0006 0001 -0451 0060 NS 0013 NS

ILC1r 0012 -0005 00002 0317 -0053 NS 0001 NS

ILC1l 0010 -0001 0001 0252 -0013 NS 0015 NS

ILC2l -0009 0013 0003 -0118 NS 0064 NS 0013 NS

ILr 0009 0012 0009 0265 0142 NS 0087 NS

ILl 0005 0007 0013 0197 0107 NS 0154 NS

ILC2r -0004 0011 -0003 -0057 NS 0057 NS -0013 NS

SLr 0001 00007 0001 0077 NS 0013 NS 0021 NS

SLl 00004 -0004 0006 0017 NS -0064 NS 0081 NS








SVL -0508 0277 0199 -0938 0198 0102 NS

HL -0445 -0894 0024 -0788 -0614 0011 NS

TL -0367 0154 -0910 -0815 0133 NS -0561

END -0365 0182 0237 -0912 0177 0164

ID -0337 0178 0098 -0864 0176 0070 NS

HWS -0307 0149 0178 -0922 0174 0148 NS

ED -0258 0105 0183 -0862 0136 NS 0170
















1948)




















SVL -0513 0124 0169 -0963 0085 NS 0093 NS

TL -0403 0683 -0539 -0801 0498 -0315

END -0392 -0108 0324 -0909 -0092 NS 0221

ID -0347 -0211 0269 -0855 -0191 0195

HL -0342 0028 0279 -0922 0028 NS 0221

HWS -0331 -0676 -0643 -0727 -0544 -0414

ED -0265 -0034 0103 -0876 -0041 NS 0100 NS



























Taxonomy





dichrous)












Peters 1863












hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III

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 ESP 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FRA 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ITA 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 JPN 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 KOR 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 SUO 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 SVE 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 ENU (Use these settings to create Adobe PDF documents for quality printing on desktop printers and proofers Created PDF documents can be opened with Acrobat and Adobe Reader 50 and later) gtgt Namespace [ (Adobe) (Common) (10) ] OtherNamespaces [ ltlt AsReaderSpreads false CropImagesToFrames true ErrorControl WarnAndContinue FlattenerIgnoreSpreadOverrides false IncludeGuidesGrids false IncludeNonPrinting false IncludeSlug false Namespace [ (Adobe) (InDesign) (40) ] OmitPlacedBitmaps false OmitPlacedEPS false OmitPlacedPDF false SimulateOverprint Legacy gtgt ltlt AddBleedMarks false AddColorBars false AddCropMarks false AddPageInfo false AddRegMarks false ConvertColors NoConversion DestinationProfileName () DestinationProfileSelector NA Downsample16BitImages true FlattenerPreset ltlt PresetSelector MediumResolution gtgt FormElements false GenerateStructure true IncludeBookmarks false IncludeHyperlinks false IncludeInteractive false IncludeLayers false IncludeProfiles true MultimediaHandling UseObjectSettings Namespace [ (Adobe) (CreativeSuite) (20) ] PDFXOutputIntentProfileSelector NA PreserveEditing true UntaggedCMYKHandling LeaveUntagged UntaggedRGBHandling LeaveUntagged UseDocumentBleed false gtgt ]gtgt setdistillerparamsltlt HWResolution [1200 1200] PageSize [612000 792000]gtgt setpagedevice
























































1015 -5945)













































































Passos et al 2005)




Sexual dimorphism













































0434 p lt 0001) (Figs 7 and 8)





VE 0812 0188 -0511 0967 0102 NS -0225

SC 0557 -0422 0702 0864 -0298 0403275

ILGl -0084 -0023 0039 -0721 -0090 NS 0125

ILGr -0079 -0022 0043 -0737 -0094 NS 0151

MD 0063 0015 -0010 0898 0100 NS -0055 NS

ATl 0061 0595 0285 0176 0777 0302

PTr 0055 0126 -0072 0304 0315 -0147

ATr 0049 0633 0384 0137 0796 0393

PTl 0023 0109 -0075 0137 0287 -0159

SLOl 0023 0014 0007 0399 0116 NS 0049 NS

SLOr 0022 0005 -0005 0378 0039 NS -0033 NS

ILC1r 0016 -0005 -0004 0396 -0058 NS -0037 NS

ILC1l 0015 0005 -0007 0350 0056 NS -0063 NS

LO -0007 -0001 -0008 -0223 -0016 -0090 NS

ILC2r 0005 -0019 0006 0065 NS -0115 NS 0032 NS

ILr 0004 0005 0006 0149 0084 NS 0078 NS

ILl 0003 0008 00009 0119 NS 0116 NS 0010 NS

SLl 00009 0005 0006 0048 NS 0125 NS 0118 NS

ILC2l -00006 0002 -0006 -0007 NS 0013 NS -0028 NS

SLr 00005 00004 -0003 0025 NS 0010 NS -0062 NS




VE 0746 -0468 0439 0950 -0242 0190

SC 0626 0720 -0288 0896 0419 -0140 NS

ATr 0132 -0378 -0724 0392 -0454 -0728

ATl 0100 -0270 -0420 0367 -0402 -0523

ILGl -0083 0060 0009 -0757 0223 0029 NS

ILGr -0072 0070 0013 -0688 0272 0044 NS

MD 0065 -0024 -0001 0904 -0137 NS -0007 NS






TABLE 2 (Continued)


PTr 0043 -0124 -0077 0267 -0315 -0163

PTl 0038 -0135 -0120 0256 -0366 -0272

SLOl 0032 -0036 0029 0426 -0193 0132 NS

SLOr 0031 -0015 0010 0453 -0092 NS 0051 NS

LO -0019 0006 0001 -0451 0060 NS 0013 NS

ILC1r 0012 -0005 00002 0317 -0053 NS 0001 NS

ILC1l 0010 -0001 0001 0252 -0013 NS 0015 NS

ILC2l -0009 0013 0003 -0118 NS 0064 NS 0013 NS

ILr 0009 0012 0009 0265 0142 NS 0087 NS

ILl 0005 0007 0013 0197 0107 NS 0154 NS

ILC2r -0004 0011 -0003 -0057 NS 0057 NS -0013 NS

SLr 0001 00007 0001 0077 NS 0013 NS 0021 NS

SLl 00004 -0004 0006 0017 NS -0064 NS 0081 NS








SVL -0508 0277 0199 -0938 0198 0102 NS

HL -0445 -0894 0024 -0788 -0614 0011 NS

TL -0367 0154 -0910 -0815 0133 NS -0561

END -0365 0182 0237 -0912 0177 0164

ID -0337 0178 0098 -0864 0176 0070 NS

HWS -0307 0149 0178 -0922 0174 0148 NS

ED -0258 0105 0183 -0862 0136 NS 0170
















1948)




















SVL -0513 0124 0169 -0963 0085 NS 0093 NS

TL -0403 0683 -0539 -0801 0498 -0315

END -0392 -0108 0324 -0909 -0092 NS 0221

ID -0347 -0211 0269 -0855 -0191 0195

HL -0342 0028 0279 -0922 0028 NS 0221

HWS -0331 -0676 -0643 -0727 -0544 -0414

ED -0265 -0034 0103 -0876 -0041 NS 0100 NS



























Taxonomy





dichrous)












Peters 1863












hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III































Passos et al 2005)




Sexual dimorphism













































0434 p lt 0001) (Figs 7 and 8)





VE 0812 0188 -0511 0967 0102 NS -0225

SC 0557 -0422 0702 0864 -0298 0403275

ILGl -0084 -0023 0039 -0721 -0090 NS 0125

ILGr -0079 -0022 0043 -0737 -0094 NS 0151

MD 0063 0015 -0010 0898 0100 NS -0055 NS

ATl 0061 0595 0285 0176 0777 0302

PTr 0055 0126 -0072 0304 0315 -0147

ATr 0049 0633 0384 0137 0796 0393

PTl 0023 0109 -0075 0137 0287 -0159

SLOl 0023 0014 0007 0399 0116 NS 0049 NS

SLOr 0022 0005 -0005 0378 0039 NS -0033 NS

ILC1r 0016 -0005 -0004 0396 -0058 NS -0037 NS

ILC1l 0015 0005 -0007 0350 0056 NS -0063 NS

LO -0007 -0001 -0008 -0223 -0016 -0090 NS

ILC2r 0005 -0019 0006 0065 NS -0115 NS 0032 NS

ILr 0004 0005 0006 0149 0084 NS 0078 NS

ILl 0003 0008 00009 0119 NS 0116 NS 0010 NS

SLl 00009 0005 0006 0048 NS 0125 NS 0118 NS

ILC2l -00006 0002 -0006 -0007 NS 0013 NS -0028 NS

SLr 00005 00004 -0003 0025 NS 0010 NS -0062 NS




VE 0746 -0468 0439 0950 -0242 0190

SC 0626 0720 -0288 0896 0419 -0140 NS

ATr 0132 -0378 -0724 0392 -0454 -0728

ATl 0100 -0270 -0420 0367 -0402 -0523

ILGl -0083 0060 0009 -0757 0223 0029 NS

ILGr -0072 0070 0013 -0688 0272 0044 NS

MD 0065 -0024 -0001 0904 -0137 NS -0007 NS






TABLE 2 (Continued)


PTr 0043 -0124 -0077 0267 -0315 -0163

PTl 0038 -0135 -0120 0256 -0366 -0272

SLOl 0032 -0036 0029 0426 -0193 0132 NS

SLOr 0031 -0015 0010 0453 -0092 NS 0051 NS

LO -0019 0006 0001 -0451 0060 NS 0013 NS

ILC1r 0012 -0005 00002 0317 -0053 NS 0001 NS

ILC1l 0010 -0001 0001 0252 -0013 NS 0015 NS

ILC2l -0009 0013 0003 -0118 NS 0064 NS 0013 NS

ILr 0009 0012 0009 0265 0142 NS 0087 NS

ILl 0005 0007 0013 0197 0107 NS 0154 NS

ILC2r -0004 0011 -0003 -0057 NS 0057 NS -0013 NS

SLr 0001 00007 0001 0077 NS 0013 NS 0021 NS

SLl 00004 -0004 0006 0017 NS -0064 NS 0081 NS








SVL -0508 0277 0199 -0938 0198 0102 NS

HL -0445 -0894 0024 -0788 -0614 0011 NS

TL -0367 0154 -0910 -0815 0133 NS -0561

END -0365 0182 0237 -0912 0177 0164

ID -0337 0178 0098 -0864 0176 0070 NS

HWS -0307 0149 0178 -0922 0174 0148 NS

ED -0258 0105 0183 -0862 0136 NS 0170
















1948)




















SVL -0513 0124 0169 -0963 0085 NS 0093 NS

TL -0403 0683 -0539 -0801 0498 -0315

END -0392 -0108 0324 -0909 -0092 NS 0221

ID -0347 -0211 0269 -0855 -0191 0195

HL -0342 0028 0279 -0922 0028 NS 0221

HWS -0331 -0676 -0643 -0727 -0544 -0414

ED -0265 -0034 0103 -0876 -0041 NS 0100 NS



























Taxonomy





dichrous)












Peters 1863












hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III








0434 p lt 0001) (Figs 7 and 8)





VE 0812 0188 -0511 0967 0102 NS -0225

SC 0557 -0422 0702 0864 -0298 0403275

ILGl -0084 -0023 0039 -0721 -0090 NS 0125

ILGr -0079 -0022 0043 -0737 -0094 NS 0151

MD 0063 0015 -0010 0898 0100 NS -0055 NS

ATl 0061 0595 0285 0176 0777 0302

PTr 0055 0126 -0072 0304 0315 -0147

ATr 0049 0633 0384 0137 0796 0393

PTl 0023 0109 -0075 0137 0287 -0159

SLOl 0023 0014 0007 0399 0116 NS 0049 NS

SLOr 0022 0005 -0005 0378 0039 NS -0033 NS

ILC1r 0016 -0005 -0004 0396 -0058 NS -0037 NS

ILC1l 0015 0005 -0007 0350 0056 NS -0063 NS

LO -0007 -0001 -0008 -0223 -0016 -0090 NS

ILC2r 0005 -0019 0006 0065 NS -0115 NS 0032 NS

ILr 0004 0005 0006 0149 0084 NS 0078 NS

ILl 0003 0008 00009 0119 NS 0116 NS 0010 NS

SLl 00009 0005 0006 0048 NS 0125 NS 0118 NS

ILC2l -00006 0002 -0006 -0007 NS 0013 NS -0028 NS

SLr 00005 00004 -0003 0025 NS 0010 NS -0062 NS




VE 0746 -0468 0439 0950 -0242 0190

SC 0626 0720 -0288 0896 0419 -0140 NS

ATr 0132 -0378 -0724 0392 -0454 -0728

ATl 0100 -0270 -0420 0367 -0402 -0523

ILGl -0083 0060 0009 -0757 0223 0029 NS

ILGr -0072 0070 0013 -0688 0272 0044 NS

MD 0065 -0024 -0001 0904 -0137 NS -0007 NS






TABLE 2 (Continued)


PTr 0043 -0124 -0077 0267 -0315 -0163

PTl 0038 -0135 -0120 0256 -0366 -0272

SLOl 0032 -0036 0029 0426 -0193 0132 NS

SLOr 0031 -0015 0010 0453 -0092 NS 0051 NS

LO -0019 0006 0001 -0451 0060 NS 0013 NS

ILC1r 0012 -0005 00002 0317 -0053 NS 0001 NS

ILC1l 0010 -0001 0001 0252 -0013 NS 0015 NS

ILC2l -0009 0013 0003 -0118 NS 0064 NS 0013 NS

ILr 0009 0012 0009 0265 0142 NS 0087 NS

ILl 0005 0007 0013 0197 0107 NS 0154 NS

ILC2r -0004 0011 -0003 -0057 NS 0057 NS -0013 NS

SLr 0001 00007 0001 0077 NS 0013 NS 0021 NS

SLl 00004 -0004 0006 0017 NS -0064 NS 0081 NS








SVL -0508 0277 0199 -0938 0198 0102 NS

HL -0445 -0894 0024 -0788 -0614 0011 NS

TL -0367 0154 -0910 -0815 0133 NS -0561

END -0365 0182 0237 -0912 0177 0164

ID -0337 0178 0098 -0864 0176 0070 NS

HWS -0307 0149 0178 -0922 0174 0148 NS

ED -0258 0105 0183 -0862 0136 NS 0170
















1948)




















SVL -0513 0124 0169 -0963 0085 NS 0093 NS

TL -0403 0683 -0539 -0801 0498 -0315

END -0392 -0108 0324 -0909 -0092 NS 0221

ID -0347 -0211 0269 -0855 -0191 0195

HL -0342 0028 0279 -0922 0028 NS 0221

HWS -0331 -0676 -0643 -0727 -0544 -0414

ED -0265 -0034 0103 -0876 -0041 NS 0100 NS



























Taxonomy





dichrous)












Peters 1863












hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III





TABLE 2 (Continued)


PTr 0043 -0124 -0077 0267 -0315 -0163

PTl 0038 -0135 -0120 0256 -0366 -0272

SLOl 0032 -0036 0029 0426 -0193 0132 NS

SLOr 0031 -0015 0010 0453 -0092 NS 0051 NS

LO -0019 0006 0001 -0451 0060 NS 0013 NS

ILC1r 0012 -0005 00002 0317 -0053 NS 0001 NS

ILC1l 0010 -0001 0001 0252 -0013 NS 0015 NS

ILC2l -0009 0013 0003 -0118 NS 0064 NS 0013 NS

ILr 0009 0012 0009 0265 0142 NS 0087 NS

ILl 0005 0007 0013 0197 0107 NS 0154 NS

ILC2r -0004 0011 -0003 -0057 NS 0057 NS -0013 NS

SLr 0001 00007 0001 0077 NS 0013 NS 0021 NS

SLl 00004 -0004 0006 0017 NS -0064 NS 0081 NS








SVL -0508 0277 0199 -0938 0198 0102 NS

HL -0445 -0894 0024 -0788 -0614 0011 NS

TL -0367 0154 -0910 -0815 0133 NS -0561

END -0365 0182 0237 -0912 0177 0164

ID -0337 0178 0098 -0864 0176 0070 NS

HWS -0307 0149 0178 -0922 0174 0148 NS

ED -0258 0105 0183 -0862 0136 NS 0170
















1948)




















SVL -0513 0124 0169 -0963 0085 NS 0093 NS

TL -0403 0683 -0539 -0801 0498 -0315

END -0392 -0108 0324 -0909 -0092 NS 0221

ID -0347 -0211 0269 -0855 -0191 0195

HL -0342 0028 0279 -0922 0028 NS 0221

HWS -0331 -0676 -0643 -0727 -0544 -0414

ED -0265 -0034 0103 -0876 -0041 NS 0100 NS



























Taxonomy





dichrous)












Peters 1863












hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III






SVL -0508 0277 0199 -0938 0198 0102 NS

HL -0445 -0894 0024 -0788 -0614 0011 NS

TL -0367 0154 -0910 -0815 0133 NS -0561

END -0365 0182 0237 -0912 0177 0164

ID -0337 0178 0098 -0864 0176 0070 NS

HWS -0307 0149 0178 -0922 0174 0148 NS

ED -0258 0105 0183 -0862 0136 NS 0170
















1948)




















SVL -0513 0124 0169 -0963 0085 NS 0093 NS

TL -0403 0683 -0539 -0801 0498 -0315

END -0392 -0108 0324 -0909 -0092 NS 0221

ID -0347 -0211 0269 -0855 -0191 0195

HL -0342 0028 0279 -0922 0028 NS 0221

HWS -0331 -0676 -0643 -0727 -0544 -0414

ED -0265 -0034 0103 -0876 -0041 NS 0100 NS



























Taxonomy





dichrous)












Peters 1863












hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III














1948)




















SVL -0513 0124 0169 -0963 0085 NS 0093 NS

TL -0403 0683 -0539 -0801 0498 -0315

END -0392 -0108 0324 -0909 -0092 NS 0221

ID -0347 -0211 0269 -0855 -0191 0195

HL -0342 0028 0279 -0922 0028 NS 0221

HWS -0331 -0676 -0643 -0727 -0544 -0414

ED -0265 -0034 0103 -0876 -0041 NS 0100 NS



























Taxonomy





dichrous)












Peters 1863












hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III






SVL -0513 0124 0169 -0963 0085 NS 0093 NS

TL -0403 0683 -0539 -0801 0498 -0315

END -0392 -0108 0324 -0909 -0092 NS 0221

ID -0347 -0211 0269 -0855 -0191 0195

HL -0342 0028 0279 -0922 0028 NS 0221

HWS -0331 -0676 -0643 -0727 -0544 -0414

ED -0265 -0034 0103 -0876 -0041 NS 0100 NS



























Taxonomy





dichrous)












Peters 1863












hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III

























Taxonomy





dichrous)












Peters 1863












hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III




hook























































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III











































































































































and lateral edges












mean 096 SD=046 n=71)













(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III





(n=19 2435)







Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III



Drymoluber dichrous

oce

eas




transitional areas

1050 mm (MPEG 17235)



2


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

157ndash173 (mean 165 SD=335 n=172)

160ndash180 (mean 172 SD=372 n=114)

87ndash110 (mean 97 SD=420 n=122)

86ndash109 (mean 98 SD=479 n=89)





ide)

1 (n=10

+11

ides)

(n=1

2+22

)

1+1 (1 side) 1+11 (n=38 sides) 11+1 (n=2

sides) 11+11 (n=480 sides) 11+12 (n=7

sides) 11+21 (n=8 sides) 11+22 (n=1

side) 12+11 (n=20 sides) 21+11 (n=14

sides) 21+12 (n=1 side) or 22+11 (n=4

sides)


CO

ST

A E

T A

L

370


2013 M

agnolia Press

















Preoculars 1 1



11+11 (n=90 sides) 11+12 (n=8 sides) 11+2


(n=19 sides) 12+12 (n=5 sides) 12+21 (n=5 s

12+212 (n=1 side) 12+31 (n=1 side) 12+41




2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III



2013 Magnolia P

ress middot 371

Drymoluber dichrous






10 (n=7 sides)

1st pair












TA

XO

NO

MIC

RE

VIS

ION

OF

DR

YM

OL

UB

ER

TABLE 5 (Continued)




orbit




9 (n=2 sides)



mental

1st pair 1st pair


pair of chinshields





















































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III








































































































the senior author






















16 maxillary teeth





scales mean 08)


























































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III














































(Fig 19B)











(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III




(mean 232 SD=090 n=20 100)







































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III




































sparse spinules

























distribution range


























































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III



















































































































(Lehr et al 2004)











D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III










D apurimacensis


D dichrous

Acknowledgements



























References











































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III







































































































































Appendix I


Appendix II













Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III









Appendix III









Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III




Abstract

Resumo

Introduction








Acknowledgements

References

Appendix I

Appendix II

Appendix III


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

Documents

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