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Nov 24, 2008 - Philodryas laticeps Werner, 1900 was previously known only from the holotype, which was believed to be lost during the World Wars. We found ...
Zootaxa 1940: 25–40 (2008) www.mapress.com / zootaxa/

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Copyright © 2008 · Magnolia Press

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ZOOTAXA

Rediscovery and redescription of the type of Philodryas laticeps Werner, 1900 and the taxonomic status of P. oligolepis Gomes, 1921 (Serpentes, Colubridae) HUSSAM ZAHER1,4 GUSTAVO SCROCCHI2 & ROBERTA MASIERO3 1

Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42494-970, São Paulo, SP, Brasil. E-mail: [email protected] Instituto de Herpetología, Fundación Miguel Lillo, Miguel Lillo 251, 4000 San Miguel de Tucumán, Argentina. E-mail: [email protected] 3 Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42494-970, São Paulo, SP, Brasil. E-mail: [email protected] 4 Corresponding author 2

Abstract Philodryas laticeps Werner, 1900 was previously known only from the holotype, which was believed to be lost during the World Wars. We found the holotype to be housed in the Museum für Naturkunde, Berlin and here redescribe it and report on three additional specimens from the collections of the Instituto Butantan, São Paulo and Museum für Naturkunde, Berlin. We conclude that Philodryas oligolepis Gomes, 1921 and P. affinis Müller, 1928 are junior synonyms of Philodryas laticeps Werner, 1900. All specimens share the presence of a black mucosa surrounding the larynx and trachea in the floor of the mouth and distal rows of middle sized spines disposed in a typical “V-shaped” configuration on the asulcate surface of the hemipenial body, two uniquely derived features absent in all other species of the genus Philodryas. Intraspecific variation of external morphology, hemipenes, and coloration are documented. Key words: Xenodontinae, Philodryas laticeps Werner, Philodryas oligolepis Gomes, Philodryas affinis Müller, taxonomy, hemipenis, distribution

Introduction The genus Philodryas is known to include 15 to 22 species (we recognize 18, see Table 1) of large diurnal South American snakes for which few taxonomic papers are available (Peters and Orejas-Miranda, 1970; Thomas, 1976, 1977, 1997; Thomas and Johnson, 1984; Thomas and Fernandes, 1996; see Table 1). Until recently, Philodryas laticeps was recognized as a subspecies of P. viridissima (Thomas, 1976), and represented one of the most poorly known taxa of the genus. The species was described by Werner (1900), who based his description on only one specimen. This specimen was thought to be housed in the Naturhistorisches Museum Wien, because Werner worked extensively with the Vienna collection and signed as a member of this museum the paper in which Philodryas laticeps is described. This fact might have misled previous workers who tried to locate the specimen. The holotype of P. laticeps was finally found to be housed in the Berlin Museum für Naturkunde under the collection number ZMB 15704, having been “donated” to the Museum by Ferdinand Weichberger. The information related to the specimen bears locality data from “Santa Catharina, Brasilien,” the same as the type locality published by Werner. Besides, the specimen matches perfectly the original description, showing the unusual scutellation of two preoculars and three postoculars mentioned by Werner. Werner (1900) distinguished Philodryas laticeps from the two uniformly bright green species P. olfersii and P. viridissima mainly by the presence of two preoculars and three postoculars, recognizing a closer rela-

Accepted by D. Gower: 25 Sept. 2008; published: 24 Nov. 2008

25

tionship of his species with P. viridissima. Philodryas laticeps could be further distinguished from the remaining species of Philodryas by the combination of two characters included in the original description: a low number of dorsal scale rows (17) and the presence of laterally angulated ventrals. These two characters are also valid to separate P. laticeps from P. olfersii and P. viridissima. However, two other poorly known green species of Philodryas—P. affinis Müller, 1928 and P. oligolepis Gomes, 1921—also merit consideration because they retain a similarly low number of dorsal scale rows (17 and 15, respectively) and laterally angulated ventrals (not angulated in P. affinis, according to the original description). The holotype of the former was collected in Buenavista, Department of Santa Cruz de la Sierra, Bolivia, whereas the latter is from Mariana, State of Minas Gerais, Brazil. As mentioned by Müller (1928) in his original description of P. affinis, the species is most similar to P. laticeps because they share the same number of dorsal scale rows, being distinguished only by the number of pre- and postoculars which are lower in P. affinis (one and two, respectively) than in P. laticeps (two and three). Thomas (1976) recognized the latter condition as an obvious anomaly and concluded that P. affinis was a junior synonym of P. viridissima laticeps. Because the holotypes of both P. laticeps and P. affinis were thought by Thomas (1976) to be lost, he designated as neotype of P. viridissima laticeps another Bolivian specimen with 17 dorsal scale rows, deposited at the Field Museum of Natural History, Chicago under the number FMNH 168003. Gomes (in Amaral, 1921) distinguished Philodryas oligolepis from the other species of Philodryas by the presence of divided internasal scales and dorsal scales in 15-15-11 rows. Thomas (1976) placed Philodryas oligolepis in the synonymy of Philodryas viridissima viridissima, considering the presence of two pairs of internasals an anomaly “occasionally encountered in most snakes” and the loss of dorsal scale rows as “not deemed a useful taxonomic character unless some geographic consistency is evident.” However, the difference in dorsal scale rows between these two species is substantial and entails the loss of four rows and not two as suggested by Thomas (1976: 232). Following Thomas’ argument, P. oligolepis would be actually more closely comparable to his P. viridissima laticeps, which has 17 dorsal scale rows. His decision to synonymize P. oligolepis with P. viridissima viridissima seems to be based on a criterion of geographical proximity instead of distinction in dorsal scale counts. Other mentions of Philodryas oligolepis in the literature are found in Chippaux (1986) and Starace (1998). Chippaux (1986) reported the presence of the species in French Guiana, based on a specimen (MNHN 1978.2600) that had been previously identified by Gasc and Rodrigues (1980) as P. olfersii. More recently, Starace (1998) followed Chippaux in reporting the presence of the species in French Guiana, based on the MNHN specimen, but illustrated the species in his book with a picture of a specimen of P. olfersii (Michel Blanc, pers. comm.). We had the opportunity to analyze pictures of MNHN 1978.2600, and found out that it belongs to the genus Drymoluber. We conclude that Philodryas oligolepis is not present in French Guiana. In the last few years, three additional specimens assignable to Philodryas laticeps were found in the collections of the Instituto Butantan (IBH 7600 and IBH 73141) and Berlin Museum für Naturkunde (ZMB 69958). Ferrarezzi and Franco (pers. comm.) pointed out that one of these specimens (IBH 7600) was figured by Amaral (1977: 174) as Philodryas viridissima. With the discovery of Werner´s holotype of Philodryas laticeps and of the three additional specimens mentioned above, it became evident that a re-evaluation of the taxonomic status of the species of Philodryas with 17–15 dorsal scale rows was in need. In the present study, we re-describe the holotype of Philodryas laticeps, and furnish additional information on the color pattern, scuttelation and hemipenial morphology of the species. We also conclude that P. oligolepis Gomes is a junior synonym of P. laticeps Werner, and follow Thomas (1976) in considering P. affinis Müller as a junior synonym of P. laticeps Werner.

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TABLE 1. List of species and subspecies of the genus Philodryas recognized in three traditionally cited sources and in the present study. Numbers correspond to following notes: (1) Thomas’ PhD Dissertation is an unpublished work according to articles 8 and 9 of the ICZN. Several subsequent authors simply followed the taxonomic arrangements suggested by Thomas, thus validating these new arrangements without offering any satisfactory explanation to support the changes. (2) Uetz et al. (2008) lists 22 species of Philodryas; however, two species – P. biserialis and P. inca – are included in the list without any justification; the former species clearly does not belong to Philodryas (see Zaher, 1999) while the latter was convincingly synonymized with P. simonsii by Thomas (1977). (3) New combination proposed by Barrio (1959). (4) Subspecies described by Barrio (1959) and subsequently synonymized with P. a. subcarinata by Barrio et al. (1977: 230). (5) Species described by Amaral (1932); correct spelling is P. arnaldoi, not arnoldoi. (6) A synonym of P. trilineata (see Cei, 1993: 643–644). (7) Considered a synonym of P. olfersii herbeus by Lancini and Kornacker (1989: 212), who probably followed Thomas (1976) by mentioning only “In früheren Arbeiten wird die Grüne Strauchnatter als Philodryas carbonelli bezeichner. Dieser Name ist somit als Synonym von Philodryas olfersii herbeus anzusehen.” (8) Species considered to belong to the genus Philodryas by Boulenger (1896), Dowling (1969), Peters and Orejas-Miranda (1970), Myers and Hoogmoed (1974) and Thomas (1977); more recently allocated to the genus Alsophis by Thomas (1997) and Zaher (1999). (9) Correct spelling is Alsophis elegans rufodorsatus, not rufidorsatus (see Vanzolini, 1986: 21). (10) Species allocated to the genus Tropidodryas by Thomas and Dixon (1977). (11) Subspecies informally described by Thomas (1976), and explicitly recognized by Giraudo (2001) who furnished a diagnosis for P. aestivus levisquamus, thus producing a nomen nudum according to Articles 13 and 16 of the ICZN. (12) Species informally resurrected by Thomas (1976) from the synonymy of Philodryas psammophidea. (13) Species validated by Thomas et al. (1977) and posteriorly synonymized with P. varia by Thomas and Johnson (1984). (14) Species previously allocated to Dromicus by Peters and Orejas-Miranda (1970) and in Alsophis by Maglio (1970), but subsequently transferred to Philodryas by Thomas (1977); the species was further considered a Philodryas by Zaher (1999) who furnished hemipenial characteristics supporting this allocation. (15) New combinations proposed informally by Thomas (1976). (16) New subspecies described informally by Thomas (1976). (17) Species removed from Leimadophis and allocated to Philodryas by Thomas (1977). (18) Species removed from Dromicus and allocated to Philodryas by Thomas (1977). (19) Species and subspecies listed under Philodryas by Uetz et al. (2008), apparently based on Thomas (1976). (20) Species allocated to Philodryas by Thomas (1999). (21) Species removed from the synonymy of Philodryas olfersii without any justification. Peters and Orejas-Miranda, 1970

Thomas, 1976 (1)

Uetz et al., 2008 (2)

Present study

Philodryas aestivus aestivus (3)

Philodryas aestivus aestivus

Philodryas aestiva aestiva (19)

Philodryas aestiva

Philodryas aestivus manegar- Philodryas aestivus subcarinatus (4) zoni (4)

Philodryas aestiva subcarinata (19)

Philodryas arnaldoi

Philodryas arnaldoi (5)

Philodryas aestivus levisquamus (11)

Philodryas arnaldoi

Philodryas baroni

Philodryas baroni

Philodryas arnoldoi (5)

Philodryas baroni

Philodryas chamissonis

Philodryas burmeisteri (6)

Philodryas baroni

Philodryas biserialis biseria- Philodryas cordata lis (2)

Philodryas carbonelli (7)

Philodryas bolivianus (12)

Philodryas biserialis eibli (2) Philodryas hoodensis

Philodryas elegans elegans (8)

Philodryas borellii (13)

Philodryas boliviana (19)

Philodryas laticeps

Philodryas elegans rufidorsa- Philodryas burmeisteri tus (9)

Philodryas chamissonis

Philodryas livida

Philodryas mattogrossensis

Philodryas cordata

Philodryas mattogrossensis

Philodryas chamissonis (14)

Philodryas nattereri

Philodryas mattogrossensis

Philodryas hoodensis (20)

Philodryas nattereri

Philodryas olfersii

Philodryas nattereri

Philodryas inca (2)

Philodryas olfersii

Philodryas oligolepis

Philodryas olfersi olfersi (15)

Philodryas laticeps (21)

Philodryas patagoniensis

Philodryas patagoniensis

Philodryas olfersi herbeus (15)

Philodryas livida

Philodryas psammophidea

Philodryas psammophideus

Philodryas olfersi latirostris (15)

Philodryas mattogrossensis

Philodryas simonsii

Philodryas pseudoserra (10)

Philodryas patagoniensis

Philodryas nattereri

Philodryas tachymenoides

Philodryas serra (10)

Philodryas psammophideus psammophideus (15)

Philodryas olfersii olfersii (19)

Philodryas trilineata

TAXONOMY AND MORPHOLOGY OF PHILODRYAS LATICEPS

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27

......continue TABLE 1. (continued) Peters and Orejas-Miranda, 1970

Thomas, 1976 (1)

Uetz et al., 2008 (2)

Present study

Philodryas viridissimus

Philodryas psammophideus lativittatus (15)

Philodryas olfersii herbeus (19)

Philodryas varia

Philodryas psammophideus andensis (16)

Philodryas olfersii latirostris (19)

Philodryas viridissima

Philodryas simonsi (17)

Philodryas oligolepis

Philodryas tachymenoides (18)

Philodryas patagoniensis

Philodryas viridissimus viridissimus (15)

Philodryas psammophidea

Philodryas viridissimus laticeps (15)

Philodryas simonsii Philodryas tachymenoides Philodryas trilineata(see 6) Philodryas varius (see 13) Philodryas viridissima

Materials and methods Specimens examined are deposited in the following institutions (acronyms used in the text are given in parenthesis): Centro de Pesquisas do Cacau, Ilhéus (CEPEC); Instituto Butantan, São Paulo (IBH); Museu de Zoologia da Universidade de São Paulo (MZUSP); Museum für Naturkunde, Berlin (ZMB); Muséum National d’Histoire Naturelle de Paris (MNHN). We were able to locate and examine six specimens of Philodryas laticeps: 1) an adult female from Santa Catarina, Brazil (ZMB 15704, Holotype of Philodryas laticeps); 2) an adult male from Mariana (20°22'S; 43°29'W), Minas Gerais, Brazil (MZUSP 1389, holotype of Philodryas oligolepis); 3) an adult female from Baixo Guandu (ex Itá) (19°30'S; 41°01'W), Espírito Santo, Brazil (IBH 7600); 4) an adult male from Regência (19°36'S; 39°49'W), Espírito Santo, Brazil (IBH 73141); 5) an adult male from “Brazil”, without additional information (ZMB 69958); 6) an adult male from the mouth of Rio Chaparé, at Rio Mamoré (circa 15°56'S 64°41'W), Department of Cochabamba, Bolivia (FMNH 168003). Hemipenes of four males of Philodryas laticeps (MZUSP 1389, IBH 73141, ZMB 69958, FMNH 168003) were prepared and compared with those available for Philodryas viridissima (MZUSP 7839, 5192; IBH 32948, 45819, 51998, 24778, 14758, 57195, 24127, 14758, 10724; MNHN 3837) and P. olfersii (IBH 35862 35868; MZUSP 14013, 9893, 11910, 11926, 11924, 11921, 11904, 10721, 11906; CEPEC 1016; MNHN 1993.1620). Methods of hemipenial preparation and terminology follow Zaher (1999), Myers and Cadle (2003), and Zaher and Prudente (2003). Ventral scales were counted according to Dowling´s method (1951), starting with the first scale bordered on each side by the first dorsal scales row. However, the number of preventrals is also given for the holotype. Preventral scales correspond to the gulars that are distinctly wider than long, anterior to the first ventral scale (Dowling, 1951). This more traditional method of counting has the advantage of being more readily comparable with older literature (Myers 2003). Total length (TTL) and tail length (TAL) were measured to the nearest 1 mm by stretching carefully the specimens along a ruler. Head, snout, and head scale lengths were measured to the nearest 0.01 mm with a digital caliper, and then converted to proportions for purposes of description. Images were taken with a digital camera and mounted using Adobe Photoshop. We counted every dorsal scale row in all six available specimens (Table 2) and illustrate those counts with a graph representation (Fig. 1). We chose to use this method instead of Dowling’s (1951) because we found it more appropriate to illustrate the anomalous dorsal scale counts found in some specimens (especially in

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ZAHER ET AL.

MZUSP 1389) instead of representing these atypical pholidosis with a formula. Complete dorsal scale counts for each specimen examined is given in Table 2, while the full spectrum of dorsal scale variation can be observed in Figure 1. Counting of each dorsal row was performed on a posteriorly oriented diagonal line, from left to right side of the specimen (Table 2). Complete scale row counts were performed using ventral scales as a marker, the first dorsal scale row including the dorsal scale that touches the first ventral scale on the left side.

......continue

TAXONOMY AND MORPHOLOGY OF PHILODRYAS LATICEPS

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FIGURE 1. Graphic representation of the dorsal scale counts of six specimens of Philodryas laticeps. a) MZUSP 1389 (holotype of Philodryas oligolepis), b) IBH 73141, c) IBH 7600, d) ZMB 69958 e) ZMB 15704 (holotype of Philodryas laticeps), f) FMNH 168003. The X axis represents the number of each ventral scale and the Y axis the number of scales composing the corresponding dorsal scale row (counted from the left side).

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TABLE 2. Number of dorsal scales for each scale row in six specimens of Philodryas laticeps (ZMB 15704 and MZUSP 1389 are the holotypes of P. laticeps and P. oligolepis, respectively). MZUSP 1389

IB 73141

ZMB 15704

ZMB 69958

FMNH 168003

Ventrals

Dorsals

Ventrals

Dorsals

Ventrals

IB 7600 Dorsals

Ventrals

Dorsals

Ventrals

Dorsals

Ventrals

Dorsals

1

17

1-2

19

1

19

1

19

1

19

1–111

17

2–8

16

3–6

18

2–4

18

2–4

18

2

17

112–113

16

9–13

15

7–94

17

5–102

17

5–101

17

3–116

16

114–117

15

14

16

95–99

16

103–106 16

102–103 16

117

15

118–119

14

15–17

15

100

15

107–108 15

104–105 17

118–121

14

120

13

18

16

101–102 16

109–115

14

106

16

122–183 13

121

14

19–60

15

103

116–212

13

107–108 14

184–185 12

122–205 13

61

16

104–108 14

109–160 13

186–187 11

62–66

15

109–188 13

161–164 12

188–199 12

67–68

16

189–191 12

165–166 13

200–210 11

69

17

192

13

167–169 12

211–215

70

16

193

12

170–176 13

71–75

15

194–195 11

177–178 11

76–80

16

196–197 12

179–185 12

81–83

15

198–202 11

186–187 11

84

16

203–204 12

188–194 12

85–115

15

205–212 13

195–198 13

116

16

117–119

15

120

14

121

13

122–125

12

126–147

11

148

12

149–153

11

154

12

155–175

11

176

12

177–181

11

182

12

183–211

11

212

12

213

11

214

12

215

14

216

12

217

11

218

12

15

12

199–204 12

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Results Philodryas laticeps Werner, 1900 (Fig. 2) 1900. Philodryas laticeps Werner. Beschreibung einiger noch unbekannter neotropischer und indischer Reptilien. Zool. Anz., 23: 196–198. 1928. Philodryas affinis Müller. Herpetologische Mitteilungen. Zool. Anz., 78: 61–84. Holotype: ZSM 8/1928 (Zoologische Staatssammlung München), an adult male from Buena Vista (80 km northwest from Santa Cruz de la Sierra), Bolivia, donated in 1926 by Carlos Marie and presently lost. 1921. Philodryas oligolepis Gomes. In: Amaral, A. Ultimos trabalhos ineditos de J. Florencio Gomes: Duas novas especies de Colubrideos opisthoglyphos brasileiros (Philodryas oligolepis e Apostolepis longicauda). Annaes Paulistas de Medicina e Cirurgia 12 (7–8): 110–113. Holotype: MZUSP 1389 (Museu de Zoologia da Universidade de São Paulo, Brasil; formerly Collecção do Museu Paulista), an adult male from Mariana, Minas Gerais, Brasil, collected in 1898 by J. B. de Godoy. 1977. Philodryas oligolepis, Amaral. Serpentes do Brasil. Iconografia colorida. Edições Melhoramentos and Editora da Universidade de São Paulo, Brasil. 246 pp.

Holotype: ZMB 15704 (Berlin Museum für Naturkunde), an adult female from Santa Catarina, Brazil, procured by Ferdinand Weichberger (Fig. 2). Diagnosis: Philodryas laticeps differs from all other species in the genus by the following combination of characters: head, body, and tail uniformly green, except for the gular region and anteriormost ventrals which are white; mucosa surrounding the larynx and trachea in the floor of the mouth covered with black pigmentation; lack of well-defined longitudinal rows of large and shallow body calyces on the distal half of the asulcate surface of the hemipenial body; lateral enlarged spines reduced to middle sized spines; distal rows of middle sized spines disposed in a typical “V-shaped” configuration on the asulcate surface of the hemipenial body; diameter of the central portion of the hemipenis larger than both proximal and distal extremities; dorsal scale rows reducing anteroposteriorly from 19–17 to 13–11; high number of ventral scales (204 to 218); ventrals angulated laterally. Re-description of the holotype: When our observations differ from the original description (Werner, 1900), the latter is indicated in square brackets. The specimen is an adult female of 1045 mm [1060] total length, 301 mm [295] tail length (28.8% of total length); head length 27 mm (2.6% of total length); head width 19 mm at broadest point. Head distinct from neck. Dorsal scales smooth, in 17-17-12 rows (see Table 2 for more details), with two apical pits. There are two preventrals (sensu Dowling, 1951), 204 ventrals, a divided anal scale, and 116 [117] paired subcaudals. Ventrals are angulated laterally. Rostral scale 1.1 times higher than wide, visible from above; paired subequal internasals; paired prefrontals 1.2 times wider than long, in contact with each other and with frontal, supraocular, upper preocular, loreal, nasal, and internasal; frontal sub-pentagonal nearly as wide as long; supraoculars longer than wide; parietals 1.7 times longer than wide; nasal divided above and below naris, in contact with the first and second supralabials; loreal nearly as wide as long; two preoculars, the upper one nearly twice as large as the lower preocular, while the left preoculars are completely separated, the right ones are only partially divided; three postoculars, the lower one being the smallest one, while the upper two are of equal size; one large anterior temporal and one slightly smaller posterior temporal; eight supralabial scales, with second and third bordering the loreal, and fourth and fifth bordering the orbit; mental scale triangular; 11 infralabial scales on the left side and 12 on the right, the first to fifth contacting the anterior genial; anterior and posterior genials paired, the former being wider and shorter than the latter genials. Coloration of the holotype: According to Werner (1900), the color pattern of the holotype is uniformly “grass” green dorsally (including supralabials) and whitish green on the ventral scales, turning “grass” green on subcaudals. Our observations on the holotype confirm that the ventral (gular) region of the head and anterior third of the belly are white, turning progressively into light green on the posterior two-third of the belly.

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The dorsal and lateral surfaces of the head and dorsum are uniformly dark green. Intraspecific variation: Five adult specimens of Philodryas laticeps in addition to the holotype were available for study (MZUSP 1389, IBH 73141, 7600, FMNH 168003, ZMB 69958; ZSM 8/1928 not included in the analysis) (Table 3). All specimens are similar in color pattern to the holotype. Largest specimen a male (IBH 73141) 1165 mm TTL, 314 mm TAL; largest female 1055 mm TTL, 293 mm TAL. Tail with 27–29% of TTL, slightly longer in females (28–29% of TTL; N=2) than in males (27%; N=3). Head distinct from neck, body slightly laterally compressed, ventrals laterally angulated. Ventral counts 204–218 (mean = 211.0; SD = 5.51; N = 6) not significantly different between males and females (males with 205–218, mean = 212.0; SD = 5.57; N = 4; females with 204–212, mean = 208; SD = 5.66; N = 2). Anal always divided. Subcaudals in paired rows (divided) throughout the tail. Subcaudal counts 112–123 (mean = 115.60; SD = 4.23; N = 5), not significantly different between males and females (males with 112–123, mean = 116.0; SD = 6.08; N = 3; females with 114–116, mean = 115.0; SD 1.41; N = 2). Dorsal scales smooth with two apical pits and undifferentiated vertebral row. Dorsal scale counts ranging from 19 scale rows on the neck to a minimum of 11 rows just anterior to the cloaca, with reductions from 19 to 15 dorsal scale rows occurring between ventrals 94–119, and from 13 to 11 dorsal scale rows between ventrals 109–126 (see Tables 2, 3 and Figure 1 for details).

FIGURE 2. Holotype of Philodryas laticeps (ZMB 15704). Head in (a) dorsal, (b) ventral, (c) right lateral, and (d) left lateral views. Body in (e) dorsal and (f) ventral views. TAXONOMY AND MORPHOLOGY OF PHILODRYAS LATICEPS

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FIGURE 3. Heads of Philodryas laticeps in (a) right lateral, (b) left lateral, (c) dorsal, and (d) ventral views. MZUSP 1389 (top; holotype of P. oligolepis Gomes); IBH 73141 (upper middle); IBH 7600 (lower middle); ZMB 69958 (bottom).

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TABLE 3. Comparison of measurements (in mm) and standard characters in six specimens of Philodryas laticeps (ZMB 15704 and MZUSP 1389 are the holotypes of P. laticeps and P. oligolepis, respectively). Data for ZSM 8/1928 from the original description. Numbers in bold and highlighted in gray represent scale anomalies. TTL = total length; TAL = tail length. Specimens ZMB 15704 IB 7600 MZUSP 1389 IB 73141 ZMB 69958 FMNH 168003 ZSM 8/1928

Sex

& & % % % % %

ventrals

subcaudals

TTL

TAL

internasals

supralabials

infralabials

204

116

1045

301

1/1

8/8

11/12

212

114

1055

293

1/1

9/9

12/12

218

-

850

-

2/2

8/8

11/11

212

112

1165

314

1/2

10/9

11/11

215

123

1080

295

1/1

9/9

12/12

205

113

1081

296

1/1

8/8

11/11

212

113

1198

325

1/1

8/8

-

continued.

Specimens ZMB 15704 IB 7600 MZUSP 1389 IB 73141 ZMB 69958 FMNH 168003 ZSM 8/1928

Sex

& & % % % % %

preoculars

postoculars

ant temporals post temporals Dorsal scale rows

2/2

3/3

1/1

1/1

19–18–17–16–14–13–12–11

1/1

2/2

1/1

1/1

19–18–17–16–15–14–13

1/1

2/2

1/1

1/1

17–16–15–14–13–12–11

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Rostral visible from above. Prefrontals paired and only slightly wider than long; prefrontal contacts its mate, frontal, supraocular, preocular, loreal, nasal and internasal. Internasals paired (MZUSP 1389 with both internasals divided and IBH 73141 with right internasal divided); when internasal is divided, prefrontal contacts both scales. Nasal divided, contacts supralabials 1–2. Loreal present, width and length nearly the same size and contacts nasal, preocular, and supralabials 2–3. Preocular single (ZMB 15704 with two preoculars), 2.0 times higher than wide, and contacting supraocular, prefrontal, loreal and supralabials 3–4. Two postoculars of same size (ZMB 15704 with three postoculars). One anterior and one posterior temporal (ZMB 69958 with 1+2 temporals on the right side). Supraoculars 2.1 times longer than wide. Frontal pentagonal, 1.1 to 1.3 times longer than maximum width. Parietals paired, 1.6 to 1.7 times longer than wide. Eight, nine, or 10 supralabials (ZMB 15704, MZUSP 1389, FMNH 168003 with eight supralabials, IBH 7600 and ZMB 69958 with nine supralabials, IBH 73141 with 10 supralabials on the left side and nine on the right side), supralabials 4–5 in contact with orbit (IBH 73141 with left supralabials 4-5-6 in contact with orbit); 11 or 12 infralabials (ZMB 15704 with 11 infralabials on the left side and 12 on the right side, IBH 7600 and ZMB 69958 with 12 infralabials, the remaining three specimens with 11 infralabials), infralabials 1–5 in contact with anterior genials. Posterior genials slightly longer than anterior genials. Scale anomalies: All six specimens of Philodryas laticeps examined show a peculiarly high number of scale anomalies that include extra-numerary supralabials, infralabials, preoculars, postoculars, internasals, and highly irregular dorsal scale counts (Figs. 1, 2, 3; Tables 2, 3). Additionally, some specimens show an intriguing combination of anomalous scales or share the same anomalies (Table 3). All specimens present variable dorsal scale counts that vary from 19 or 17 scales on the anteriormost rows to 13 or 11 scales on the more posterior ones. However, all counts are included within 19 and 11 dorsal scale rows (Table 3). The holotype of Philodryas oligolepis (MZUSP 1389, Fig. 2) shows the highest number of changes in dorsal counts

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FIGURE 4. Hemipenes of Philodryas laticeps. (a, b) MZUSP 1389, (c, d) IBH 73141, and (e, f) ZMB 69958, in asulcate (upper) and sulcate (lower) views. All hemipenes are fully everted and maximally expanded.

whereas FMNH 168003 has the least variable counts (Table 3). Three specimens (IBH 7600, ZMB 69958, and the right side of IBH 73141) share the presence of nine supralabial scales, instead of eight as in the remaining specimens (Fig. 3). The additional scale is always small, with a triangular shape, and interposed between the

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sixth and the penultimate supralabials. Whether its presence resulted from the division of the sixth or the seventh supralabials is hard to define. The shape of the supernumerary scale differs significantly in the three specimens: the dorsal border of the supernumerary scale touches the postocular and first temporal in IBH 7600 and ZMB 69958 but only the first temporal in IBH 73141. Additionally, IBH 73141 shows another supernumerary supralabial scale on the left side, interposed between the fourth and fifth supralabials as a third scale bordering the eye. MZUSP 1389 and IBH 73141 share the presence of paired (divided) internasals. In IBH 73141, the right internasal is fully divided while the left internasal is only divided on its anterolateral border. The remaining specimens have single internasal scales on both sides. Two specimens (IBH 7600, ZMB 69958) have 12 infralabials instead of 11, while ZMB 15704 retains 11 infralabials on the left side and 12 on the right side.

FIGURE 5. Distribution map of Philodryas laticeps. Star, type locality (ZMB 15704); cross, specimen ZSM 8/1928 (not analysed).

Coloration in life: According to Gomes (in Amaral, 1921), the live specimen (holotype of Philodryas oligolepis) is “green above, greenish beneath; with inferior part of the head white.” The color in life described by Gomes (op. cit) and Werner (1900) can be identified also in preserved specimens. The dorsal surfaces of the head and body are nearly uniform green in all specimens. The ventral region of the head and anteriormost ventral scales are white whereas the rest of the belly is lighter green. Hemipenial description: The following description is based on four fully everted and maximally expanded hemipenes (MZUSP 1389, IBH 73141, ZMB 69958, FMNH 168003) (Fig. 4). Philodryas laticeps has a short hemipenial body with short but clearly bilobed lobes, Thomas’ (1976) type 2 hemipenis. The organ is semicalyculate and only slightly semicapitate. The diameter of the central portion of the hemipenis is expanded (inflated) with respect to both proximal and distal extremities, conferring an oval shape to the organ. The sulcus spermaticus divides at the base of the calyculate area, and the branches terminate centrolineally on

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the portion of the sulcate surface of the lobes, but not at their tip. Both capitulae are formed from papillate calyces and are totally confluent in the intrasulcar region, forming an uninterrupted calyculate area mostly restricted to the sulcate side of the organ. The calyces invade only marginally the distal and lateral borders of the asulcate side of the lobes. Most of the asulcate surface as well as the medial surface of the lobes and the short lobular crotch are ornamented with medium-sized body calyces that extend onto the distal one-third of the asulcate surface of the hemipenial body. Unlike all other species of Philodryas, large and shallow body calyces are absent on the asulcate surface of the hemipenis. Body calyces gradually grow in size from the tip of the lobes towards the distal one-third of the hemipenial body, resulting in medium-sized body calyces that are significantly smaller than the same structures present in the other species of Philodryas. Both capitula and body calyces are restricted to the distal one-third of the hemipenis, being bordered proximally by uniform rows of medium sized spines that cover the rest of the hemipenial body. Spines are larger around the borders of the capitula and along the sulcus spermaticus, decreasing gradually in size towards the proximal end of the hemipenis where they are replaced by spinules. Spinules cover most of the proximal half or third of the hemipenial body. The most distal rows of spines that border the body calyces on the asulcate surface are disposed in a typical “V-shaped” configuration. Distribution: Two apparently disjunct populations restricted to the tropical rainforest of central Bolivia in the Departments of Santa Cruz and Cochabamba, and the Atlantic rainforest of Southern and Southeastern Brazil in the States of Santa Catarina, Espírito Santo, and Minas Gerais (Fig. 5).

FIGURE 6. Hemipenis of Philodryas viridissima (IBH 45819) in asulcate (left) and sulcate (right) views.

FIGURE 7. Mouth of Philodryas laticeps (MZUSP 1389) opened in left lateral view to show the mucosa surrounding the larynx and trachea in the floor of the mouth covered with black pigmentation.

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Discussion Until presently, the types of Philodryas laticeps and P. affinis were thought to be lost and Philodryas oligolepis was known only from the holotype and a dubious record for French Guiana (Gomes in Amaral, 1921; Chippaux, 1986), which has been now correctly identified as belonging to the genus Drymoluber. In addition to the scarcity of material, the few specimens known until recently showed an unusually high number of scale anomalies that prevented systematists to define accurately their taxonomic status. Philodryas oligolepis was originally described by Gomes (in Amaral, 1921) who distinguished it from the other green species of Philodryas on the basis of its fewer dorsal scales (17-17-11 instead of 19-19-13) and the presence of two pairs of (divided) internasal scales. The species was synonymized by Thomas (1976) with P. v. viridissima, who distinguished the latter from P. v. laticeps by its higher number of dorsal scale rows at midbody (19 instead of 17) and deeper distal body calyces on the hemipenes. According to Thomas (1976), P. v. laticeps would have “rather shallow, distal asulcate calyces on the hemipenes”. Actually, P. viridissima differs considerably in its hemipenis from P. laticeps (compare Figs. 4 and 6), the former showing a longer hemipenial body relative to the lobes, larger calyces that extend throughout most of the asulcate side of the body, lack of capitulum, and a calyculate area restricted to the sulcate side of the lobes. However, considering the information available at that time, Thomas’ conclusions were appropriate. During the analysis of ZMB 15704, ZMB 69958, MZUSP 1389, IBH 73141, and IBH 7600, we found that all specimens have dorsal scale rows reducing anteroposteriorly from 19–17 to 13–11, and males share the same hemipenial pattern with the presence of middle sized spines disposed in a typical “V-shaped” configuration on the asulcate surface of the hemipenial body. Additionally, we found that they share the presence of a black mucosa surrounding the larynx and trachea on the floor of the mouth (Fig. 7), an unusual derived condition absent in all other species of Philodryas (confirmed in FMNH 168003; M. Kearney, pers. comm.). Besides the dorsal scale counts, hemipenial features, and black mucosa, all six specimens also share an overall green dorsum and laterally angulated ventrals. The presence of these derived features in both holotypes of P. oligolepis and P. laticeps support our conclusion that the former species is a junior synonym of the latter. Although the type of P. affinis is presumed to be lost, the description given by the author provides sufficient information for us to believe that this is also a junior synonym of P. laticeps.

Acknowledgements We are grateful to Alain Dubois, Annemarie Ohler, and Ivan Ineich (MNHN), Rainer Günther (ZMB), Maureen Kearney (FMNH), Francisco L. Franco and Hebert Ferrarezzi (IBH) for allowing us to examine specimens under their care. We are also deeply indebted to Roger Bour for sending photographs of specimen MNHN 1978.2600; to Michel Blanc for providing important information on his photograph of the specimen depicted in Starace’s book as Philodryas oligolepis; to Abbie Wolf for kindly preparing and photographing the hemipenis of FMNH 168003; to Maureen Kearney, Alan Resetar, and Tom Anton for confirming the black color of the throat of FMNH 168003 and sending scale counts and photographs of the latter; and to Alberto B. de Carvalho for editing the figures. The senior author is especially grateful to Christoph Kucharzewski for his precious help with the analysis of the type material during the visit to the Herpetological collection of the Berlin Museum für Naturkunde. This research was supported by grants from Fundação de Amparo à Pesquisa do Estado de São Paulo (BIOTA-FAPESP grant number 02/13602–4) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq grant number 303785/2004–7).

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References Amaral, A. (1921) Últimos trabalhos inéditos de J. Florencio Gomes: Duas novas especies de Colubrideos opisthoglyphos brasileiros (Philodryas oligolepis e Apostolepis longicauda). Annaes Paulistas de Medicina e Cirurgia, 12 (7– 8), 110–113. Amaral, A. (1977) Serpentes do Brasil. Iconografia colorida. Edições Melhoramentos e Editora da Universidade de São Paulo, Brasil. Barrio, A., Laurent, R.F. & Thomas, R.A. (1977) The status of Philodryas subcarinatus Boulenger (Reptilia, Serpentes, Colubridae). Journal of Herpetology, 11: 230–231. Chippaux, J.P. (1986) Les serpents de la Guyane Française. Institute Français de Recherche Scientifique pour le développement en Coopération. Collection faune Tropicale n° XXVII. Paris. Dowling, H.G. (1951) A proposed standard system of counting ventrals in snakes. British Journal of Herpetology, 1: 97– 99. Gasc, J.P. & Rodrigues, M.T. (1980) Liste préliminaire des serpents de Guyane Française. Bulletin du Muséum National d’Histoire Naturelle de Paris, 4e. série, 2 A (2), 559–598. Myers, C.W. (2003) Rare snakes – Five new species from Eastern Panama: Reviews of Northern Atractus and Southern Geophis (Colubridae: Dipsadinae). American Museum Novitates, 3391, 1–47. Myers, C.W. & Cadle, J.E. (2003) On the snake hemipenis, with notes on Psomophis and techniques of eversion: a response to Dowling. Herpetological Review, 34 (4), 295–302. Peters, J.A. & Orejas-Miranda, B. (1970) Catalogue of the Neotropical squamata. Part I. Snakes. Smithsonian Institution Press, Washington. Starace, F. (1998) Guide des serpents et Amphisbènes de Guyane. Ibis Rouge Editions. Thomas, R.A. (1976) A revision of the South-American Colubrid snake genus Philodryas Wagler, 1830. University Microfilms International. Ann Arbor, Michigan, U.S.A. Thomas, R.A. (1977) A new generic arrangement for Incaspis and mainland South American alsophis and the status of two additional Peruvian species. Copeia, 1977 (4), 648–652. Thomas, R.A. (1997) Galápagos terrestrial snakes: biogeography and systematics. Herpetological Natural History, 5 (1), 19–40. Thomas, R.A., Laurent, R.F. & Barrio, A. (1977) Philodryas borellii Peracca (Serpentes: Colubridae), a distinct species. Herpetologica 33, 82–86. Thomas, R.A. & Johnson, J.D. (1984) Philodryas varius (Jan, 1863), a senior synonym of Philodryas borellii Peracca (Serpentes: Colubridae). Journal of Herpetology, 18 (1), 80. Thomas, R.A. & Fernandes, R. (1996) The systematic status of Platyinion lividum Amaral, 1923 (Serpentes: Colubridae: Xenodontinae). Herpetologica, 52 (2), 271–275. Uetz, P., Goll, J. & Hallermann, J. (2008) The Reptile Database, http://www.reptile-database.org, accessed August 02, 2008. Zaher, H. (1999) Hemipenial morphology of the South American snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of Colubroid hemipenes. Bulletin of the American Museum of Natural History, 240, 1–168. Zaher, H. & Prudente, A.L.C. (2003) Hemipenis of Siphlophis (Serpentes: Xenodontinae) and techniques of hemipenial preparation in snakes: A response to Dowling. Herpetological Review, 34, 302–307.

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