Zootaxa, Notes on Distribution, Variation and Characterization Of

Home , El Escondido, Erythrolamprus, Erythrolamprus bizona, Micrurus

Notes on distribution, variation and characterization of Erythrolamprus pseudocorallus Roze, 1959 (Serpentes: Colubridae) with the first records from Colombia

FELIPE FRANCO CURCIO1, SANTIAGO J. SÁNCHEZ-PACHECO3, JONH JAIRO MUESES-CISNEROS4 & MIGUEL TREFAUT RODRIGUES1, 2 1Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11.461, CEP 05422-970, São Paulo, SP, Brazil. E-mail: [email protected], [email protected] 3Programa de Pós-Graduação em Biologia Animal, Universidade Federal do Rio Grande do Sul–UFRGS, Avenida Bento Gonçalves, 9500, 91501-970, Porto Alegre, RS, Brasil & Conservación Internacional Colombia, Bogotá, D. C. Colombia. E-mail: [email protected] 4Laboratorio de anfibios, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, D. C. E-mail: [email protected]

Abstract

The Maracaibo false coral snake Erythrolamprus pseudocorallus, previously known only from Venezuela, is recorded from five departments in Colombia. These new data include the westernmost and the southernmost records presently known for the species. Two specimens previously identified as E. aesculapii, from the localities of El Valle, Distrito Federal, Venezuela, and Yarumal, Antioquia, Colombia, are now attributed to E. pseudocorallus, the first one representing the northeasternmost record of the species. Morphological characterization of E. pseudocorallus is expanded based on the new specimens.

Key words: Maracaibo false coral snake, new records, range extension, South America

Resumen

La serpiente “falsa coral de Maracaibo” Erythrolamprus pseudocorallus, previamente conocida solamente de Venezuela, es registrada en cinco departamentos de Colombia, incluyendo los registros más hacia el oeste y hacia el sur conocidos actualmente para esta especie. Dos especímenes previamente identificados como E. aesculapii, provenientes de El Valle, Distrito Federal, Venezuela y Yarumal, Antioquia, Colombia, son asignados a E. pseudocorallus, el primero representando el registro más hacia el noreste de la especie. Con base en estos nuevos especímenes, se amplía la caracterización morfológica de E. pseudocorallus.

Introduction

As presently understood, the genus Erythrolamprus Boie (Serpentes; Xenodontinae) includes six species of false coral snakes that occur in Central and South America, and also in the Caribbean islands of Trinidad and Tobago (Roze, 1959a, 1966; Peters & Orejas-Miranda, 1970; Cunha & Nascimento, 1980; Hardy & Boos, 1995; Boos, 2001). In Colombia, the genus is represented by the taxa E. aesculapii aesculapii (Linnaeus), E. bizona Jan and E. mimus micrurus Dunn & Bailey (Peters & Orejas-Miranda, 1970; Pérez-Santos & Moreno, 1988; Sánchez et al., 1995). Roze (1959a) described E. pseudocorallus (Maracaibo false coral snake) from the state of Zulia, Venezuela, and reported its occurrence in highlands next to Maracaibo and Perijá (Roze, 1966). Later, range

Accepted by D. Gower: 23 Jan. 2009; published: 18 Mar. 2009 33 extensions were published for the Venezuelan states of Mérida and Amazonas (La Marca & García, 1986; Fuentes & Barrio, 1999; La Marca & Soriano, 2004). Even though the species had never been formally recorded beyond the borders of Venezuela, Roze (1959a) mentioned the possibility of E. pseudocorallus occurring also in northwestern Colombia, and Pérez-Santos & Moreno (1988) suggested its possible occurrence in the Colombian department of Guajira, in Perijá region. Field surveys in the “Área Natural Única Los Estoraques”, municipality of La Playa de Belén, Norte Santander, Colombia, and examination of specimens housed in collections of Brazilian, Colombian and North American institutions, resulted in the detection of 22 specimens of Erythrolamprus from at least 15 different localities in Colombia, all matching most of the coloration and scale count features presented in the original description of E. pseudocorallus. These vouchers represent the first records of the species in Colombia and, along with a new record from Venezuela, allow us to more precisely characterize the species and report new information on its distribution, coloration, and lepidosis.

Materials and methods

All specimens examined in this study (see Appendix 1) were identified through direct comparison with the type series housed in the collections of the Museo de Biología, Universidad Central de Venezuela (MBUCV), and the Museo de Historia Natural La Salle [MHNLS (also SCN, Leviton et al., 1985)], both in Caracas, Venezuela. Specimens recorded from Colombia are deposited in the following collections: Academy of Natural Sciences of Philadelphia (ANSP; Philadelphia, PA, USA), American Museum of Natural History (AMNH; New York, NY, USA), Instituto Butantan (IBSP; São Paulo, SP, Brazil), Instituto de Ciencias Naturales–Universidad Nacional de Colombia (ICN), Museo de la Universidad La Salle (MLS) and Pontificia Universidad Javeriana (MUJ) (last three institutions in Bogotá, D.C., Colombia). One specimen of E. a. aesculapii from the Museu Paraense Emílio Goeldi, (Belém, PA, Brazil) is illustrated to allow comparison of color patterns (MPEG 19649, Boa Vista, Roraima, Brazil). Ventral scale counts follow the wider-than-long system (Peters, 1964). Measurements were taken to the nearest mm with a graduated ruler. Terminology regarding color patterns follows Savage & Slowinski (1992). Monads on the body and tail were counted separately; the dark nuchal collar was not included in monad counts. In all specimens, there is always one monad placed exactly on the region of the vent; thus its anterior and posterior halves were included in body and tail monad counts, respectively. Monads and red ring lengths are somewhat difficult to measure precisely in preserved coral snakes because red pigment tends to fade in ethanol and thereby blends with the narrow outer pale rings of monads. For that reason we provide four different band-length measurements, always counted as the number of dorsal scales in the 8th vertebral scale row (Table 1). The length of a “monad” refers herein to the central black ring together with its respective narrow outer pale rings; when we refer strictly to black ring length, the indistinguishable outer pale rings are not considered. Accordingly, when we refer to red ring length, we only consider the red space between two monads and do not take into account the pale outer rings of the latter. When red coloration is indistinguishable from the outer pale rings, we use the measurement of “interblack rings”, i.e. the distance between two black rings, including the undetected scales of the pale rings of the monads. Monad and black ring lengths were measured for the first, fourth, and last monads; the last monad was considered as the one prior to the vent. Red and interblack rings were measured in the first, fourth, and last rings; the first red ring was considered as the one immediately behind the first monad, whereas the last red ring was considered as the one between the second-to-last and the last monad, according to the criterion explained above. Coordinate data of localities (included in Appendix 1) were taken from gazetteers (Paynter, Jr., 1982, 1997), provided by the institutions, or directly taken with GPS devices at the localities of concern. The software Google Earth was used to confirm and to refine coordinate data of most of the localities.

Characterization and variation. A monadal patterned (sensu Savage & Slowinski, 1992) Erythrolamprus with symmetrical black rings, possibly showing pale spots on its ventrolateral surface (Fig. 1D); red rings generally varying from as wide as to more than three times longer than monads. Ventral scales 182–198 in males and 180–197 in females; subcaudal scales 46–62 in males and 45–57 in females. Dorsal (15/15/15), supralabial (7/7, 3rd and 4th pairs in contact with the orbit), infralabial (9/9), temporal (1+2) and ocular (1/2) counts are highly conservative for the genus and most specimens have these features, although abnormal fusions might occur in head scales. Maxillary teeth 9–11 slightly increasing in size posteriorly, followed by 1–2 strongly grooved fangs; diastema present. In most specimens, red pigment has faded and the rings show a yellowish, light red, or light pink color, making it difficult or impossible to distinguish it from contiguous outer pale rings of monads. Specimens collected more recently still maintain red coloration, allowing precise identification of pale bands (Table 1).

FIGURE 1. Dorsal and ventral views of Erythrolamprus pseudocorallus (A-B: holotype, MBUCV 03789; C-D: a Colombian specimen, ICN 8932) and a monadal patterned E. a. aesculapii (E-F: MPEG 19649) from Brazilian Amazon. Head coloration allows distinction between both species.

Snout pattern consists mostly of a pale rostral scale, marked by black on the region of its upper vertex; internasals mostly black, with narrow pale contours along their anterior margins; anterior margins of prefrontals marked with pale contours for about the first 1/3 of the total length of the scale; black head cap covering up to posterior 2/3 of prefrontals, the entire frontal, supraoculars, and anteromedial 1/2–2/3 of parietals; light head band anteriorly covering the lateroposterior 1/3–1/2 of parietals, all the temporals, 3rd to 7th supralabials and often the postoculars, and extending 0–2 dorsals posteriorly; pale head band not uniformly colored, with black contours on the posterior and lateral borders of all scales it covers; black nuchal collar

ERYTHROLAMPRUS PSEUDOCORALLUS Zootaxa 2045 © 2009 Magnolia Press · 35 covering 4.5–7 scales, rarely attaining the level of parietals tips; postnuchal light collar weakly or not distinguishable from first red ring, probably not exceeding 1.5 scales in length; 1st–2nd and 5th–7th supralabials mostly pale (whitish cream), marked by varying amounts of black on their upper-posterior portions, 3rd–4th supralabials marked by black mostly on upper areas, due to invasion of pigment from the black head cap.

TABLE 1. Measurements and scale counts of Colombian and type/paratype specimens of E. pseudocorallus. BRL = black ring length, IBRL = interblack ring length (outer pale rings of monads computed), ML = monad length (outer pale rings computed), NBM = number of body monads, NTM = number of tail monads, RRL = red ring length, SCAU = number of subcaudal scales, SVL = snout-vent length, TL = tail length, VEN = number of ventral scales; * = Holotype (Roze, 1959a), ** = Paratypes (Roze, 1959a), † = no “1363” of Roze (1959a), †† = present tag number of “MHNLS (ex. coll. Dr. A. Pons No 12)” of Roze (1959a).

Number Sex SVL TL VEN SCAU NBM NTM BRL (1st, 4th ML (1st, 4th RRL (1st, 4th IBRL (1st, 4th and last) and last) and last) and last) AMNH 12899 ♂ 407 83 182 61 15.5 3.5 3.5/3/3.5 -/-/- -/-/- 7.5/6/7.5 AMNH 17522 ♂ 570 115 186 57 15.5 4.5 3/3/4 -/-/7 -/-/- 8.5/5.5/- AMNH 35538 ♀ 696 122 190 51 13.5 3 4/4/4.5 -/-/- -/-/- 10.5/8.5/11.5 AMNH 35757 ♂ 588 124 191 61 15.5 3.5 4.5/4.5/5 -/-/- -/-/- 8.5/9/6.5 AMNH 59405 ♀ 632 110 193 57 12.5 3.5 3.5/3/4 -/-/6.5 -/-/- 11/8.5/8.5 ANSP 22786 ♂ 675 123 184 61 15.5 4.5 3/2.5/4 5/4/6 8/7/3 9.5/8/8.5 ANSP 22787 ♀ 231 37 188 49 12.5 2.5 3.5/3.5/4 4/?/5 ?/?/? 10/8/10 IBSP 09179 ♂ 409 102 188 58 11.5 3.5 4.5/4.5/4.5 -/-/- -/-/- 11.0/12.5/10.0 ICN 01889 ♀ 611 99 184 46 11.5 2.5 2.5/3.0/3.0 4.0/4.5/7.0 12.0/10.0/9.0 13.5/12.5/15.0 ICN 02573 ♂ 575 129 189 59 14.5 3.5 3.5/3.0/4.0 -/-/- -/-/- 8.0/8.0/10.0 ICN 03601 ♀ 274 42 180 49 11.5 2.5 3.0/3.5/4.5 4.0/5.0/6.0 11.0/9.0/9.0 12.0/10.5/10.0 ICN 06462 ♀ 661 109 188 51 14.5 3.5 3.0/2.5/3.5 5.0/5.0/5.0 7.0/7.0/7.0 9.5/10.0/9.5 ICN 08932 ♂ 534 110 188 62 12 3 4.5/5.5/6.0 6.0/6.0/8.0 9.0/10.0/8.0 10.5/11.5/10.0 ICN 11047 ♂ 571 - 188 - 14.5 - 2/2/3 3/4/- 10.0/7.0/11.0 11.0/9.0/8.5 ICN 11109 ♀ 635 112 180 49 9.5 2.5 4.5/4.0/5.0 6.0/7.0/8.0 10.0/9.0/11.0 11.5/12.5/14.5 MUJ 0904 ♂ 478 89 187 55 14 4 3.0/2.5/3.0 -/4.0/4.0 -/8.0/- 10.0/10.0/7.0 MUJ 0905 ♂ 585 106 198 61 15.5 4.5 3.0/3.0/3.5 -/-/- -/-/- 9.0/10.0/11.0 MLS 0666 ♂ 727 112 196 46 13 4.0 3.0/3.0/3.5 5.0/5.0/4.5 10.0/5.0/7.0 12.0/7.0/10.0 MLS 0667 ♂ 629 107 191 49 14 3.0 2.5/2.5/2.0 4.0/4.0/3.5 10.0/11.0/8.0 12.0/12.0/9.5 MLS 0673 ♀ 673 110 183 47 12.5 2.5 4.0/3.0/5.0 5.0/4.5/8.0 9.0/7.0/7.0 11.0/9.5/9.5 MLS 0676 ♀ 519 99 181 55 13.5 3.5 2.5/2.5/4.5 5.0/5.0/6.0 7.0/8.0/9.0 10.0/10.0/10.5 MLS 2244 ♂ 570 113 188 60 11.5 3.5 4.0/4.0/5.5 6.5/9.0/- 8.0/8.0/- 13.0/13.0/11.0 MLS 2341 ♀ 648 112 183 49 11.5 2.5 3.5/3.5/6.0 5.5/5.0/8.5 8.0/7.0/- 10.5/10.0/5.5 MBUCV III ♂ 616 115 192 55 14.5 3.5 4.0/3.0/4.5 5.0/-/- -/-/- 10.0/8.5/8.5 03789* MBUCV III ♀ 675 109 189 47 13.5 3.5 5.0/4.5/5.0 -/-/- -/-/- 10.0/8.0/8.5 03790** MBUCV III0 ♀ 699 124 192 50 9.5 2.5 3.0/3.0/3.0 5.0/5.0/7.0 14.0/12.0/12.0 16.5/15.0/15.5 3791** MHNLS 1**† ♂ 706 143 197 62 13.5 3.5 3.0/3.0/3.5 4.0/5.0/7.0 10.0/9.0/8.0 11.5/12.0/12.0 1332**†† ♀ 808 125 197 52 12 3 3.0/2.5/3.5 -/5.0/5.0 11.0/12.0/12.0 12.0/14.0/14.0

Body coloration composed of red rings alternating with complete and symmetrical monads (sensu Slowinski & Savage, 1992); 9.5–15.5 monads on body; 2.5–4.5 monads (or dyads) on tail; posterior body monads (mostly the ones placed on the posterior half of the trunk) sometimes tend to form triads due to higher

36 · Zootaxa 2045 © 2009 Magnolia Press CURCIO ET AL. concentration of black pigment on the distal portions of red rings, forming irregular black contours contiguous with the outer pale rings (e.g. ICN 2573); tail monads sometimes tending to form dyads; red rings equal to or more than three times as long as monads (in some specimens, some body monads can be slightly longer than some of the red rings, e.g. MLS 0666 and MLS 2341). The only information on color patterns in life is a photograph by E. La Marca of a juvenile specimen (Campbell & Lamar, 2005: plate 1116), showing an evidently white tonality of the pale rings of monads as well as of the light head band. The same color was also present in specimen ICN 8932 immediately after collection. Because Roze (1959a) describes such features as “yellow”, one must consider that tonality variations might occur, a fact that led us to keep the term “pale” when describing color patterns of preserved specimens.

FIGURE 2. Updated distribution of Erythrolamprus pseudocorallus. Star = approximated type locality, in the Sierra de Perijá, west to Maracaibo, Venezuela [based on Roze (1959a)]; triangles = localities of paratypes; square = specimen of La Marca & García (1986) from El Amacao, Mérida, Venezuela; ? = specimen of Fuentes & Barrio (1999) form Gavilán region, Amazonas, Venezuela (in need of revision); white circle = specimen of Roze (1959b) from El Valle, Federal District, Venezuela; black circles = records from Colombia (1 = Fusagasugá, Cundinamarca; 2 = Sasaima, Cundinamarca; 3 = Fresno, Tolima; 4 = Medellín, Antioquia; 5 = Landázuri, Santander; 6 = Puerto Berrío, Antioquia; 7 = Yarumal, Antioquia; 8 = Piedecuesta, Santander; 9 = Bucaramanga, Santander; 10 = Segovia, Antioquia; 11 = Durania, Norte Santander; 12 = Gramalote, Norte Santander; 13 = El Zulia, Norte Santander; 14 = Ocaña, Norte Santander; 15 = La Playa de Belén, Norte Santander).

Notes on paratypes. In the original description, Roze (1959a) designated four paratypes, two of which presently are in the collections of MHNLS and two are housed in MBUCV together with the holotype. After

ERYTHROLAMPRUS PSEUDOCORALLUS Zootaxa 2045 © 2009 Magnolia Press · 37 consulting catalogues of the Museo de Historia Natural La Salle we concluded that the specimen cited by Roze (1959a: 530) as “MHNLS No (ex. col. Dr. A. Pons No 12)” now has the number “1332” printed on its tag which lacks any institutional identifier or standard symbolic code. The specimen reported by Roze (1959a: 530) as “No 1363” presently has the presumably “new” institutional tag number MHNLS 1; the number cited in the original description apparently refers to the old catalogue series. The corrected data for the tags of these specimens are included in the Appendix 1, as well as in Table 1. Regarding the locality of paratype “1332” [MHNLS (ex col. Dr. A. Pons No 12) of Roze (1959a: 530)], the original description provides the following information: “Escondido, Perijá, Estado Zulia, Venezuela, coleccionado a 1.075 metros de altura”. Paynter, Jr. (1982) provides the locality “El Escondido, Zulia, 1075 m”, as “not located”, but placed in the middle of Sierra de Perijá; the area West of the municipality of “Machiques” (10o 04’ N, 72o 34’ W) is provided as an approximation based on three short expeditions of the “Sociedad de Ciencias Naturales La Salle” that took place in the region between the years of 1947 and 1951. The updated catalogue of MHNLS provides the locality of specimen “1332” as “Hacienda El Escondido, Río Negro, Sierra de Perijá”, at an elevation of 100 m (09o 54’ N, 72o 30’ W); the records assign Moises Nava as the collector, and the date of collection as August 1951. We were informed that these data actually correspond to the base camp during the expedition, which would explain the remarkable altitudinal difference between the data provided by Roze (1959a) and those of the updated catalogue of MHNLS. Colleagues who have worked in the region provided the correct name for the locality of specimen “1332” as “El Escondido, Cañon del Río Negro, Sierra de Perijá, Estado Zulia” at an elevation of 1075 m, along with the precise coordinates 10o 03’ N, 72o 47’ W (see Appendix 1). The only additional information we have is that a community of native Indians (“Yukpa de Ayaspaina”) inhabits the area (Jaramillo, 1987; Herrera et al. 1997).

FIGURE 3. Dorsal (A) and ventral (B) view of specimen from El Valle, Districto Federal, Venezuela (AMNH 59405), tentatively determined as Erythrolamprus aesculapii by Roze (1959b) but identified herein as E. pseudocorallus.

Discussion

La Marca & García (1986) reported the southernmost record of E. pseudocorallus in El Amacao, Andrés Bello district, state of Mérida, Venezuela (8o 40’ N, 71o 24’ W; see Fig. 2). Considering the new Colombian specimens herein reported, the southernmost record is now in the municipality of Fusagasugá, Cundinamarca department (Fig. 2); this locality is about 400 km southwest of the record of La Marca & García (1986). The locality of Medellín, Antioquia, Colombia, represents the westernmost occurrence of E. pseudocorallus (4 on map, Fig. 2). The presumed easternmost occurrence of the species was reported by Fuentes & Barrio (1999), who claimed to have recorded E. pseudocorallus in the region of Gavilán, state of Amazonas, Venezuela (05o 40’ N, 67o 30’ W). This record, based on a specimen presently deposited in the Museo de Ciencias Naturales de Caracas (MCNC 7676), lies far to the east of other confirmed points of occurrence of E. pseudocorallus, which seems to be a species primarily associated with the Andes, particularly to Cordillera Central de

38 · Zootaxa 2045 © 2009 Magnolia Press CURCIO ET AL. Colombia, Cordillera Oriental de Colombia, Cordillera de Mérida and the Sierra de Perijá (Fig. 2). Additionally, the specimen reported by Fuentes & Barrio (1999) represents the only Cis-Andean record of the species; all Colombian and other Venezuelan specimens are attributed to Trans-Andean localities (Fig. 2). Finally, E. a. aesculapii, which may show monadal patterns in some populations throughout the Amazon Basin (Fig. 1 E–F) and happens to be strikingly variable in color pattern at a single locality (see Martins & Oliveira, 1998), is expected to be common in the area of Gavilán. Such patterns of local geographic variation are usually associated with the influence of mimicry complexes due to co-occurrence with poisonous coral snakes of the genus Micrurus (Greene & McDiarmid, 1981; Marques & Puorto, 1991). In the region of Manaus, Amazonas, Brazil, monadal patterns in E. a. aesculapii were associated with sympatric occurrence of the monadal patterned poisonous coral snake Micrurus averyi (Martins & Oliveira, 1998). Similar associations between coral snakes and their mimics are to be expected in the Gavilán region. After examining a photograph of the specimen from Gavilán kindly provided by C. Barrio-Amorós, we noticed the presence of a solid and immaculate white head band, not invaded by black from the head cap, as well as a distinctive white snout pattern covering most of the internasal and prefrontal scales. These features contrast with the pattern herein described for E. pseudocorallus, but seem to be in agreement with the pattern of E. a. aesculapii of the Amazon basin (compare head patterns among Figs. 1 A, C and E). We have not had access to measurements and scale counts of this specimen, but head coloration reinforces the idea that the record by Fuentes & Barrio (1999) might in fact be a misidentification of a monadal patterned specimen of E. a. aesculapii. Therefore, we consider the record of Fuentes & Barrio (1999) as doubtful and recommend that this specimen receives further attention; we have plotted it on the distribution map as a “?” (Fig. 2). The type locality of E. pseudocorallus is somewhat imprecise [“Cerca de Maracaibo, Estado Zulia, Venezuela, en la región montañosa” (Roze, 1959a)]. Thus, we included on the map an approximate locality in the Sierra de Perijá, to the west of Maracaibo, only as a reference to the region where the actual type locality must be (Fig. 2). In the original description, Roze (1959a) claimed that E. pseudocorallus was characterized by a high number of subcaudals (55–62 in males and 50–53 in females) and strongly grooved fangs, in contrast to E. mimus micrurus (the only other monadal patterned potentially co-occurring Erythrolamprus) that would have lower subcaudal counts (46–52 in males and 42–46 in females) and aglyphous or weakly grooved fangs (Roze, 1959a, 1966; Pérez-Santos & Moreno, 1988). The new Colombian specimens show that subcaudal counts in E. pseudocorallus are more variable than previously presumed (see Table 1 and comments above), and therefore is a less useful character in distinguishing that species from E. mimus micrurus. However, we believe that it is quite easy to distinguish both species based on characters of coloration and dentition. Regarding color patterns, we note that E. pseudocorallus always has complete monads, regularly distributed throughout the body. In contrast, E. mimus micrurus has at least some of the body monads divided at the level of the vertebral scale row and its halves are alternately arranged throughout the body (Dunn & Bailey, 1939). Tooth grooving is also a somewhat helpful aid to identification, with a distinctively strong groove present in E. pseudocorallus, compared to the aglyphous or weakly grooved fangs present in specimens of E. mimus micrurus (Cope 1868, Dunn & Bailey, 1939; Roze, 1959a). Most specimens were recorded from localities at elevations above 800 m, except for MLS 0666, 0673 and ICN 11109 (Appendix 1). Ventral scale counts do not show any noticeable distinction between the three lowland specimens and those collected at higher elevations, and no correlation between these parameters is apparent (Pearson’s Correlation Coefficent = -0.040, p = 0.885, n = 23). Subcaudal counts of specimens MLS 0666 and 0667 seem low for male E. pseudocorallus, especially considering only the data in the original description. Nonetheless, these specimens were collected at elevations of 100 and 1100 m, respectively, suggesting that variation of subcaudal counts is not associated with elevation. Similarly to this pattern, two females from low elevations (MLS 0673 and ICN 11109) have subcaudal counts comparable to that of ICN 03601 (also a female) from one of the highest elevations reported in the sample. Although one might detect statistical correlation between the parameters “number of subcaudals” and “elevation” (Pearson Correlation Coefficent = 0.474, p = 0.026, n = 22), we consider that

ERYTHROLAMPRUS PSEUDOCORALLUS Zootaxa 2045 © 2009 Magnolia Press · 39 effect likely a result of the small size of our sample, which deters any consideration on possible morphological differences between lowland and highland populations of E. pseudocorallus. Because color pattern appears to be conservative and no other objective character was found to distinguish more than one taxonomic entity within what is presently called E. pseudocorallus, we propose that the species is strongly associated with the Cordillera Central and the Cordillera Oriental de Colombia, occurring at a wide elevational range, being probably more frequent in areas above 800 m. Finally, we want to point out that E. pseudocorallus had been reported by previous authors, though referred by the name of other species, prior to the original description by Roze (1959a). Amaral (1935) identified a Colombian specimen from Yarumal, Antioquia, deposited in the collection of the Instituto Butantan (IBSP 9179) as E. aesculapii; this author provided data on scale counts and called attention to the monadal color pattern similar to that of some Micrurus species. Twenty four years later, in the same year when the original description was published, Roze (1959b) himself mentioned a monadal patterned Erythrolamprus from El Valle, Federal District, Venezuela (AMNH 59405; Fig. 3), that he could not assign to any species previously reported from that country. Erythrolamprus bizona, of rather common occurrence along the northern coast of Venezuela (Roze, 1957: 190), would be absolutely out of the question as the possible identity of this specimen, because that species has black rings arranged in dyads on the whole body extension, as well as on the neck (Dunn & Bailey, 1939). Thus, Roze (1959b) tentatively attributed the name E. aesculapii to that specimen, because the body monads resembled the form illustrated by Jan & Sordelli (1866) as E. aesculapii var. monozona (a subspecies of E. aesculapii, sensu Peters & Orejas-Miranda, 1970). After having examined these specimens, scale counts and general color pattern combined with the data of the present sample led us to conclude that the records of Amaral (1935) and Roze (1959b) in fact are both E. pseudocorallus, respectively representing the oldest confirmed Colombian record of the species in the literature (Amaral, 1935) and the northeasternmost record of E. pseudocorallus, located beyond the eastern limit of the Cordillera de Mérida (Roze, 1959b), in mountains along the northern coast of Venezuela. Regarding other possible records, we call attention to another Colombian specimen mentioned by Cope (1899) with uncertain locality data, possibly from the vicinity of Bogotá, that we were not able to locate. Though determined by Cope as E. aesculapii, a short comment mentions that “No. 3 has the rings on the posterior part of the body arranged in threes, which is very unusual in this species” (Cope, 1899: 15). As mentioned previously, irregular triads on the posterior part of the body may occur in some specimens of E. pseudocorallus, and we are not aware of that feature being previously reported in any specimen of the E. aesculapii complex. Accordingly, we suspect that the report by Cope (1899) could represent in fact another (and the first) record of E. pseudocorallus from Colombia. Erythrolamprus pseudocorallus has also been confused with E. mimus micrurus, and it was not unusual to find specimens of E. pseudocorallus mistakenly identified as E. mimus in some of the collections we consulted. For example, Aleman (1953) listed specimens that were later included in the type series of E. pseudocorallus as the first record of E. mimus micrurus from Venezuela before Roze’s (1959a) description had been published; in so doing, this author ignored the differences in the disposition of the monads throughout the body mentioned by Cope (1868) and Dunn & Bailey (1939). Thus, the records of the present study along with these sparse indications of literature lead us to conclude that E. pseudocorallus is a widespread species throughout the Colombian (not recorded to the west of the Cordillera Central de Colombia) and Venezuelan Andes; other records from these areas are expected as workers continue to collect and examine material in regional and other collections containing representatives of the Andean herpetofauna of these countries.

Acknowledgements

We are grateful to Francisco L. Franco, Valdir José Germano (IBSP), John D. Lynch (ICN), Andrés Acosta (MUJ), Hno. Roque Casallas and Arturo Rodriguez (MLS), Gilson Rivas Fuenmayor, Josefa Celsa Señaris

40 · Zootaxa 2045 © 2009 Magnolia Press CURCIO ET AL. (MHNLS), Mercedes Salazar (MBUCV), Darrell Frost, David Kizirian, Rob Pascocello (AMNH), Ned Gilmore, Ted Deaschler (ANSP) and Ana Lúcia Prudente (MPEG) for access to specimens under their care. Dr. Libardo Suárez, director of the “Área Natural Única Los Estoraques”, Elibardo Bayona Tarazona and Joselito Vargas provided all the facilities during the field work. Enrique La Marca, Gilson Rivas Fuenmayor, Tito Barros and Cesar Barrio-Amorós provided important information on localities and morphological details of Venezuelan specimens. Thanks are also due to Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for support. FFC was supported by a scholarship provided by the Coordenadoria de Aperfeiçoamento de Pessoal de Ensino Superior (CAPES). SJS thanks Programa de Becas para la Iniciativa de Especies Amenazadas IEA Jorge Hernández Camacho (Convenio 245–2). Luís F. Silveira provided important suggestions and allowed access to his personal library and catalogues. Vítor Q. Piacentini helped with the map and provided constructive comments. John D. Lynch and Carla Piantoni kindly revised and criticized the manuscript. The careful reviews and the suggestions provided by Zootaxa referees, Janis Roze and Roy McDiarmid, as well as by the editor David Gower resulted in considerable improvement for the final version. José Cassimiro da Silva Jr. helped to review the proofs.

References

Aleman G., C. (1953) Contribución al Estudio de los Reptiles y Batracios de la Sierra de Perijá. Memoria de la Sociedad de Ciencias Naturales La Salle, Caracas, 13, 205–225. Amaral, A. (1935) Estudos sobre Ophidios Neotrópicos. XXXII. Apontamentos sobre a fauna da Colombia. Memórias do Instituto Butantan, 9, 209–216. Boos, H.E.A. (2001) The snakes of Trinidad & Tobago. Texas A & M University Press, College Station, 270 pp. Campbell, J.A. & Lamar, W.W. (2004) The venomous Reptiles of the Western Hemisphere. Vol.2. Comstock Publishing Associates, Cornell University Press, Ithaca & London, xiv + 477–870. Cope, E.D. (1868) Sixth contribution to the herpetology of Tropical Americas. Proceedings of the Academy of Natural Sciences of Philadelphia, 1868, 305–313. Cope, E.D. (1899) On a collection of Batrachia and Reptilia from New Granada. The Philadelphia Museums Scientific Bulletin, 1, 3–19. Cunha, O.R., & Nascimento, F.P. (1980) Ofídios da Amazônia. XI. Ofídios de Roraima e notas sobre Erythrolamprus bauperthuisii Duméril, Bibron & Duméril 1854, sinônimo de Erythrolamprus aesculapii aesculapii (Linnaeus, 1758). Boletim do Museu Paraense Emílio Goeldi, nova série, Zoologia, Belém, 102, 1–21. Dunn, E.R. & Bailey, J.R. (1939) Snakes from the uplands of the Canal Zone and Darien. Bulletin of the Museum of Comparative Zoology, 86, 1–22. Fuentes, O. & Barrio, C.L. (1999) Erythrolamprus pseudocorallus (Falsa Coral de Maracaibo, Maracaibo’s False Coral Snake) Geographic Distribution. Herpetological Review, 30, 53. Greene, H.W. & McDiarmid, R.W. (1981) Coral snake mimicry: Does it occur? Science, Washington D. C., 213, 1207–1212. Hardy, J.D., Jr. & Boos, H.E.A. (1995) Snakes of the genus Erythrolamprus (Serpentes: Colubridae) from Trinidad & Tobago, West Indies. Bulletin of the Maryland Herpetological Society, 31, 158–190. Herrera V, D.P., Santos, A.Y, Arlantt, B. & Arias, S. (1997) Estúdio sobre las condiciones y calidad de vida de la etnia Yukpa del Resguardo Iroka en Codazzi (Cesar, Colombia). Diagnósticos y alternativas. Fundación Cultural Sierra Nevada, Ministerio de Salud. Instituto Colombiano de Bienestar Familiar, Bogotá, Colombia. Jan, G & Sordelli, F. (1866) Iconographie générale des ophidiens. Tome second, livraison 19. Jaramillo, O.G (1987) Yuko-Yukpa. In: Introducción a la Colombia Ameríndia. Correa, F. & Pachón, X. (Eds.) Instituto Colombiano de Antropología, Bogotá, pp. 75–82. La Marca, E. & García, J.E. (1986) Erythrolamprus pseudocorallus (False Coral Snake) Geographic Distribution. Herpetological Review, 17, 27. La Marca, E. & Soriano, P.J. (2004) Reptiles de los Andes de Venezuela. Fundación Polar, Conservación Internacional, CODEPRE-ULA, Fundacite Mérida, BioGeos, Mérida, Venezuela, 173 pp. Leviton, A.E., Gibbs, R.H., Jr., Heal, E., & Dawson, C.E. (1985) Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia, 1985, 802–832. Marques, O.A.V. & Puorto, G. (1991) Padrões cromáticos, distribuição e possível mimetismo em Erythrolamprus

ERYTHROLAMPRUS PSEUDOCORALLUS Zootaxa 2045 © 2009 Magnolia Press · 41 aesculapii (Serpentes, Colubridae) no sudeste do Brasil. Memórias do Instituto Butantan, 53, 127–134. Martins, M. & Oliveira, M.E. (1998) Natural history of snakes in forests of the Manaus region, central Amazonia, Brasil. Herpetological Natural History, 6, 78–150. Paynter, R.A., Jr. (1982) Ornithological Gazetteer of Venezuela. Bird Department, Museum of Comparative Zoology, Harvard College, iv + 245 pp., 2 maps. Paynter, R.A., Jr. (1997) Ornithological Gazetteer of Colombia. 2nd Edition. Bird Department, Museum of Comparative Zoology, Harvard College, ix + 537 pp., 2 maps. Pérez-Santos, C. & Moreno, A.G. (1988) Serpientes de Colombia. Museo Regional de Scienze Naturali, Torino, Monografia VI, 515 pp. Peters, J.A. (1964) Dictionary of Herpetology. Hafner Publishing Company, New York, New York, USA, 392 pp. Peters, J.A., & Orejas-Miranda, B.R. (1970) Catalogue of the Neotropical Squamata. Part 1. Snakes. United States National Museum Bulletin, 297, viii + 347 pp. Roze, J.A. (1957). Ofidios coleccionados por la expedición Franco-Venezolana al Alto Orinoco, 1951 a 1952. Boletín del Museo de Ciencias Naturales, Caracas, 1, 179–195. Roze, J.A. (1959a) El género Erythrolamprus Wagler (Serpentes: Colubridae) en Venezuela. Acta Biológica Venezuélica, 2, 523–534. Roze, J.A. (1959b). Taxonomic notes on a collection of Venezuelan reptiles in the American Museum of Natural History. American Museum Novitates, 1934, 1–14. Roze, J.A. (1966) La Taxonomía y Zoogeografía de los Ofidios de Venezuela. Universidad Central de Venezuela, Ediciones de la Biblioteca, Caracas, Venezuela, 362 pp. Sánchez, H.M., Castaño, O.V. & Cárdenas, A.G. (1995) Diversidad de los reptiles en Colombia. In: Rangel, J.O. (Ed.), Colombia Diversidad Biótica I, Convenio Inderena–Universidad Nacional de Colombia, Bogotá, pp. 277–324. Savage, J.M. & Slowinski, J.B. (1992) The colouration of the venomous coral snakes (family Elapidae) and their mimics (families Aniliidae and Colubridae). Biological Journal of the Linnean Society, 45, 235–254.

Appendix 1. List of specimens examined

Erythrolamprus pseudocorallus: COLOMBIA: Antioquia: Medellín, 1470 m (06o 15’ N, 75o 35’ W)”: AMNH 35538; Puerto Berrío, 144 m (06º 25’ N, 74º 30’ W): ICN 11109; Segovia, 433 m (09º 14’ N, 75º22’ W): MLS 0673; Yarumal, 2300 m (06o 58’ N, 75o 24’ W): IBSP 9179; Cundinamarca: Fusagasugá, 1692 m (04º 21’ N, 74º 22’ W): ANSP 22786, ICN 02573, MLS 2341; Sasaima, 1140 m (04º58’ N, 74º 26’ W): MLS 0676; Norte Santander: Durania, Km 5 of the Durania–La Don Juana road, 876 m (07o 44’ N, 72o 40’ W): ICN 06462; El Zulia, Astillero, 100 m (07o 56’ N, 72o 35’ W): MLS 0666; Gramalote, 1100 m (07o 54’ N, 72o 07’ W): MLS 0667; La Playa de Belén, Vereda Piritama, vicinities of the “Área Natural Única Los Estoraques”, 1600 m (08o 15’ N, 73o 15’ W): MUJ 0904–05; Ocaña, 1265 m (08o 15’ N, 73o 20’ W): ANSP 22787; Santander: Bucaramanga, 935 m (07o 07’ N, 73o 06’ W): ICN 01889, ICN 11047; Landázuri, 997 m (06o 14’ N, 73o 46’ W): ICN 08932; Piedecuesta, 982 m (06o 59’ N, 73o 03’ W): MLS 2244; Tolima: Fresno, 1572 m (05o 02’ N, 75o 02’ W): ICN 03601; New Granada: San Pedro (NOT LOCATED): AMNH 12899, AMNH 35757; Unknown locality: AMNH 17522; VENEZUELA: Distrito Federal: El Valle, 1100 m (10o 27’ N, 66o 55’ W): AMNH 59405; Zulia: cerca de Maracaibo, en la región montañosa (NOT LOCATED): MBUCV III 03789–91*; El Escondido, Cañon del Río Negro, Sierra de Perijá, 1075 m (10º 03’ N, 72º 47’ W): 1332**; Kunana, Río Negro, Sierra de Perijá, 1100 m (10o 03’ N, 72o 48’ W): MHNLS 1***.

Erythrolamprus aesculapii aesculapii: BRAZIL: Roraima: Boa Vista, 85 m (02o49’ S, 60o40’ W).

*. MBUCV III 03789 (holotype), 03790–91 (paratypes). **. specimen no “1363” of Roze (1959a) (paratype). ***. old “MHNLS (ex. coll. Dr. A. Pons No 12)” of Roze (1959a) (paratype).