Cercosaura Vertebralis O’Shaughnessy, 1879 (Reptilia: Squamata

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n Geographic distribution of Cercosaura vertebralis O’Shaughnessy, 1879 (Reptilia: Squamata:

istributio Gymnophthalmidae) and the status of Cercosaura D ampuedai (Lancini, 1968) raphic g

eo Tiffany M. Doan 1, 2* and Juan C. Cusi 3 G n o 1 State College of Florida, Manatee-Sarasota, Department of Natural Science, 5840 26th Street West, Bradenton, FL 34207, USA. 2 Current address: University of Central Florida, Department of Biology, 4110 Libra Drive, Orlando, FL 32816, USA.

otes 3 Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Departamento de Herpetología, Av. Arenales 1256, Jesús María, Apdo N 14-0434, Lima 14, Perú. * Corresponding author. E-mail: [email protected]

Abstract: We report new locality records of Cercosaura vertebralis O’Shaughnessy, 1879 in Peru, after more than 40 years since the discovery of the single known Peruvian specimen. The Peruvian range of Cercosaura vertebralis, which was previously restricted to Piura, is extended into the Cajamarca region (Tabaconas Namballe and La Granja). We also

specimens. In addition, we comment on the population status of C. vertebralis in Peru, and provide data supporting the examinedsynonymy intraspecificof C. ampuedai variation with C. vertebralisof the species. throughout its range and provide a hemipenial description from the new

DOI: 10.15560/10.5.1195

The taxon Cercosaura vertebralis O’Shaughnessy, conducted with a combination of two collecting methods: 1879 is a poorly known lizard species of the family (1) Visual Encounter Survey (VES) and (2) Pitfall traps with Gymnophthalmidae with a variety of junior synonyms. drift fences. All possible microhabitats were examined in Uzzell (1973) synonymized Prionodactylus palmeri Boulenger, 1908, P. marianus Ruthven, 1921, and Euspondylus ampuedae Lancini, 1968 with P. vertebralis. The synonymies of P. palmeri and P. marinaus have not been disputed, but La Marca and García-Pérez (1990) resurrected E. ampuedae from within P. vertebralis, accepting the transfer to Prionodactylus and emending the ending to create P. ampuedai. Mijares-Urrutia (2000) presented new data relevant to the P. ampuedai question and considered it a valid taxon, but Doan (2003) and Doan and Lamar (2012) still considered the taxon to be a junior synonym. Doan (2003) transferred all Prionodactylus (and Pantodactylus) into Cercosaura based on evidence that Prionodactylus and Pantodactylus rendered Cercosaura paraphyletic. This generic change was accepted by all subsequent authors to date. Cercosaura vertebralis has a wide geographic range which includes Panama, Colombia, Ecuador, Venezuela, and Peru (Uzzell 1973). As such, it is the only species in the genus to extend into Central America (Doan and Lamar 2012). The southernmost extent of its range occurs in Peru, with, to date, a single specimen from Piura region. Recent expeditions revealed two populations of C. vertebralis in Cajamarca, Peru: one from Santuario Nacional Tabaconas Namballe (SNTN), northern Cajamarca, and the other one from La Granja village, central Cajamarca (Figure 1). These new data not only extend the range of the taxon, but also allow a re-examination of the synonymies of the included Figure 1. Map of northwestern South America showing the geographic distribution of Cercosaura vertebralis. Red squares represent previously junior taxa. known localities whereas yellow squares indicate the new Peruvian At the SNTN, the search for this lizard species was localities. EBC = Estación Biológica Chichilapa.

1195 Doan and Cusi | Geographic distribution of Cercosaura vertebralis diurnal VES, including movement of logs, rocks, shrubs, A and the mossy ground cover. Pitfall traps were arranged in Y-shaped arrays with one central bucket and three peripheral buckets spaced 10 m apart along a fence 30 cm high (Figure 2A). Three pitfall trap arrays, separated by about 500 m, were installed at SNTN and checked thrice daily, especially after rains. This arrangement was adopted because of the irregularity of the surface, installation effort, and higher capture probability. At La Granja village, we used diurnal VES only, along a riparian forest. The study was conducted under permit number 002-2012-SERNANP-SNTN issued by Ministerio del

10% formalin and preserved in 70% ethanol; they were subsequentlyAmbiente, SERNANP. deposited Specimens in the herpetological captured were collection fixed inof the Museo de Historia Natural, Universidad Nacional Mayor B de San Marcos (MUSM) in Lima, Peru. Body proportions were measured with a digital caliper (to the nearest 0.1 mm) under a stereomicroscope (Table 1). The preparation and staining of the hemipenes of one specimen of C. vertebralis (MUSM 32489) followed Pesantes (1994) and Zaher and Prudente (2003). In addition to six new specimens (2 male and 4 juvenile) from Tabaconas Namballe and La Granja, we also examined 38 additional preserved specimens from Panama, Colombia, Ecuador, and Peru from the collections of AMNH and LACM. Four of the new specimens (Figures 3, 4) were captured at the Estación Biológica Chichilapa (EBC) within the Santuario Nacional Tabaconas Namballe. The SNTN is a natural protected area located in the northernmost region of the Peruvian occidental Andes, Cajamarca, near the 2 Figure 2. (A) Pitfall traps with fences in Y-shaped arrays placed in Peru-Ecuador border. It covers an area of 295 km with an montane scrub. (B) Composition of montane scrub in Tabaconas river elevational gradient from 1800 to 3600 m and it is a unique valley, Cajamarca, Peru. Peruvian natural area that conserves the Andean páramos and montane forests (Amanzo et al. 2003; INRENA 2007). (Jalca) and montane forest. The vegetation is dominated by bushes (1–2 m) of the families Clusiaceae, Ericaceae, in the Tabaconas river valley, with a steep slope and rugged and Melastomataceae and also herbaceous plants of the mountains,The EBC (05°16′45.4″ surrounded S, 79°18′35.0″by montane W, scrub 2273 andm) is montane situated families Eriocaulaceae, Poaceae, and Cyperaceae. The most forest. That is, it is a transitional area between grassland representative genera and species of vegetation of the area are: Meriania, Vaccinium, Cavendishia, Chelonanthus alatus, Olyra, Paepalanthus, Epidendrum secundum (orchid), Table 1. Summary of standard measurements (in mm) for Cercosaura vertebralis from Cajamarca, Peru (Range, Mean). Abbreviations are: Blechnum (hard fern) and Lycopodium thyoides (Figure Snout-vent length (SVL), head length (HL), head width (HW), head height 2B). (HH), interlimb length (IL), humerus length (HUL), radius-ulna length An additional two specimens were collected at Checos creek, La Granja, Querocoto in Cajamarca, Peru. La Granja length(RUL), of metacarpus the longest lengthtoe (LTOE). (MCL), length of the longest finger (LFIN), is a village located in the interandean valley of the Río femur length (FL), tibia-fibula length (TFL), metatarsus length (MTL) and Cercosaura vertebralis, Cercosaura vertebralis, Chotano, Querocoto, Chota and covers an elevational males (n=2) juveniles (n=4) SVL (43.03–43.74) 43.39 (26.53–29.20) 28.21 HL (10.30–10.77) 10.54 (7.35–7.87) 7.57 areas,range secondaryfrom 2200 forest, to 2800 and m. patches Checos of creek montane (06°22′31.33″ forest. The HW (6.67–7.10) 6.89 (4.52–5.17) 4.86 terrainS, 79°07′01.12″W, has a moderate 2247 to m)steep is slope, surrounded with rocky by disturbed outcrops HH (3.90–4.43) 4.17 (2.56–3.49) 3.01 IL (19.14–20.63) 19.89 (12.58–14.83) 13.80 grassland) are dominated by grasses and Asteraceae, HUL (4.09–4.99) 4.54 (2.48–3.67) 3.08 andwhile scarce secondary soil. Disturbedforest has areasabundant (agricultural bamboo fieldsChusquea and RUL (4.97–5.16) 5.07 (2.72–3.56) 3.15 scandens, trees, and shrubs. Fragments of montane forest MCL (1.91–2.12) 2.02 (1.30–1.74) 1.50 are characterized by Lauraceae, Myrsinaceae, Myrtaceae, LFIN (4.53–4.85) 4.69 (2.99–3.41) 3.16 and Melastomataceae, an elevated proportion of FL (5.58–5.92) 5.75 (3.81–4.26) 4.02 epiphytes (Tillandsia complanata, Blechnum fragile, TFL (6.48–6.68) 6.58 (4.18–4.67) 4.37 Anthurium sp., and mosses), and scarce woody lianas. MTL (2.81–2.98) 2.90 (1.68–2.16) 1.97 Understory plants include the genera Niphidium, LTOE (6.68–7.76) 7.22 (4.33–5.26) 4.75 Blechnumy, and other grasses. It is an area with a high

1196 Doan and Cusi | Geographic distribution of Cercosaura vertebralis

A A

B B

Figure 3. Male individual of Cercosaura vertebralis (MUSM 31845) from Figure 4. (A) Male individual of Cercosaura vertebralis (MUSM 32489) Tabaconas Namballe. (A) Head in lateral view showing the lateral ocelli, from La Granja. Note the absence of the white labial bar. (B) Juvenile of orange spots, and white labial bar. (B) Dorsal color pattern of a live Cercosaura vertebralis (MUSM 32490) from La Granja showing absence of specimen. orange spots that are typical in adult individuals. level of anthropogenic habitat loss. in transverse series only, in 25–37 transverse rows. Scales Most lizard specimens were found under rocks, under around midbody 26–45. Frontonasal equal in length or moss, and active on the ground, with two caught in the shorter than frontal. Supraoculars 3–4, including fusion of pitfall arrays. From the nearest locality in Canchaque, Piura, Peru, the EBC is a range extension of approximately scales two. Six or eight longitudinal rows of ventral scales. 16 km ENE and La Granja is approximately 123 km SSE. Subdigitalthe first supraocular/supraciliary lamellae of Toe IV 13–24. in all individuals. Genial Table 2 contains a summary of relevant morphological region. Features of the new specimens agree broadly data of Cercosaura vertebralis throughout its range. The withThese other are theknown first specimens records of of the this species species, in with Cajamarca minor new specimens from Cajamarca, Peru differ from most of exceptions (mentioned below). the specimens from Ecuador, Colombia, Venezuela, and Panama by lacking prefrontal scales. As pointed out by Cercosaura vertebralis O’Shaughnessy, 1879 Uzzell (1973), the single specimen from El Oro, Ecuador Cercosaura (Pantodactylus) vertebralis O’Shaughnessy, and the single specimen previously known from Piura, Peru 1879: 298. also lack these scales, but all other specimens known from Prionodactylus vertebralis: Boulenger, 1885: 394. throughout the range of the species have prefrontal scales Prionodactylus palmeri Boulenger, 1908: 518. present. In all other characters, the Cajamarca specimens Prionodactylus marianus Ruthven, 1921: 1. are quite similar to those from Panama, Ecuador, and Euspondylus vertebralis: Dunn, 1944: 103. Colombia, with meristic character states being within Euspondylus ampuedae Lancini, 1968: 4. the typical ranges of the species (Table 2). Surprisingly, Prionodactylus ampuedai: La Marca & García-Pérez, 1990: the Cajamarca specimens have little in common with the 111. single previously known Peruvian specimen from Piura, Cercosaura vertebralis: Doan, 2003: 110. sharing only the absence of prefrontal scales. The one way Cercosaura ampuedai: Hernández Ruz, 2005: 1. that the EBC Cajamarca specimens are unique is that all of the specimens possess an oblique white bar that begins on Diagnosis: A medium-sized lizard of the genus Cercosaura with keeled hexagonal dorsal scales arranged the proximal pregular scale (Figure 3A). This white bar is the final infralabial and continues posteroventrally onto 1197 Doan and Cusi | Geographic distribution of Cercosaura vertebralis

not found on any of the La Granja specimens (Figure 4) or other specimens examined from throughout the range of the species. 0 0 0–4 7–10 26–30 17–22 33–37 20–23

(2005) In comparison to the hemipenis drawings of Hernández undivided Ruz (2005) and Hernández Ruz and Bernal González “C. ampuedai” long in drawing Hernández Ruz Ruz Hernández (2011), a detailed examination of MUSM 32489 from SNTN reveals that the clavate bilobed hemipenes possess 0 21

3–9 of the asulcate surface with a naked region separating long 9–11 17–22 26–31 32–37 11–16 18–22 (2000) divided four rows of inverted V-shaped flounces in basal position straight in straight photograph “C. ampuedai”

Mijares-Urrutia Mijares-Urrutia surrounding the lobules (Figure 5). This differs from the hemipenialthose flounces drawing from of the Hernández 14 longitudinal Ruz and rows Bernal of Gonzálezflounces

asulcate surface, whereas Hernández Ruz (2005) depicted 0 0 3–9

5–10 (2011), which depicted the flounces as continuous on the 27–30

(1990) a large naked region on the asulcate surface of “Cercosaura García-Pérez García-Pérez La Marca and La Marca

“C. ampuedai” ampuedai”. spicules as pinnate structures. On the sulcate surface, the sulcus spermaticus Each flounceis bifurcate has and transverse the medial diminutive region is naked. 5 5 0 50 29

2–3 Prefrontal scales are absent in specimens from the 100 21–22 15–19 17–19 35–43 divided Panama (3) Panama long/no suture from La Marca and García-Pérez (1990), Mijares-Urrutia (2000), and Hernández (2000), Mijares-Urrutia (1990), and García-Pérez La Marca from A 4 0 0–7 100 100 20–22 27–34 19–23 11–19 38–40 divided no suture/ no suture/ prefrontals 0–5 (95% 0) long/point/3 Colombia (24) Colombia Cercosaura ampuedai Cercosaura 6 0 43 0–3 4–5 100 point 20–24 25–30 17–22 14–20 31–39 divided absent/no suture/ long/ suture/ Ecuador (10) Ecuador 2 0 0 50 21 27 (2) 100 20–21 12–13 32–36 absent divided unknown La Granja, La Granja, Cajamarca, Peru Peru Cajamarca,

B 0 33 20 26 (4) 2–3 100 100 18–20 11–15 33–38 absent divided unknown Tabaconas, Tabaconas, Cajamarca, Peru Peru Cajamarca, throughout its range. Partial data on Partial its range. throughout vertebralis Cercosaura 0 0 0 0 21 31 22 18 29 100 absent unknown undivided Piura, Peru (1) Peru Piura,

Summary of relevant morphological characters of morphological characters Summary of relevant Figure 5. Hemipenis of Cercosaura vertebralis (MUSM 31845). (A) Frontal view of asulcate surface of the right organ, showing longitudinal Lateral ocelli Lateral % with pale vertebral line % with pale vertebral % with curved pale lip line % with curved Femoral pores Females (per leg) Females pores Femoral Prefrontal presence/suture Prefrontal IV Subdigital lamellae of Toe Males (per leg) pores Femoral Transverse rows of dorsal scales rows Transverse scales of ventral rows Transverse Longitudinal rows of dorsal scales Longitudinal rows midbody Scales around Palpebral eye disc eye Palpebral Character Oblique infralabial-pregular bar Oblique infralabial-pregular

Table 2. Table (2005). Ruz spermaticus. flounces. (B) Frontal view of sulcate surface, showing the bifurcate sulcus 1198 Doan and Cusi | Geographic distribution of Cercosaura vertebralis southernmost part of the range (N=8, 19% of the 42 and García-Pérez (1990), Mijares-Urrutia (2000), specimens with prefrontal data). Specimens from central and Hernández Ruz (2005) state that C. ampuedai Ecuador to Venezuela have a wide variety of contacts of has sexual dimorphism in the number of femoral their prefrontal scales from a tiny point suture (N=2, pores but that C. vertebralis does not display sexual 5%), to a longer suture (N=22, 52%), to having no contact dimorphism. Our examination of 44 specimens from between the two prefrontal scales (N=10, 24%) (see Fig. 6). Panama, Colombia, Ecuador, and Peru found no sexual One specimen has three prefrontal scales, which includes dimorphism (with 0–6 per leg in males and 0–7 in a tiny third scale medial to the larger lateral scales (Figure females). Although there does seem to be a difference 6). There does not appear to be any trend associated with in ranges between males and females in the specimens locality for prefrontal suture contact. allocated to C. ampuedai, and the overall number of Four publications plus the new data collected in femoral pores appears to be slightly higher in those this study come to bear on the question of validity specimens than in the C. vertebralis specimens, the of Cercosaura ampuedai (La Marca and García- ranges are overlapping, which makes the utility of Pérez 1990; Mijares-Urrutia 2000; Hernández Ruz this feature as a diagnostic character questionable. 2005; Hernández Ruz and Bernal González 2011). La Marca and García-Pérez (1990) discussed the From those publications, five potentially diagnostic shape of a pale lip line as important, with C. ampuedai characters may be gleaned: sexual dimorphism of having a straight line and C. vertebralis possessing a femoral pores, shape of the pale lip line, presence of curved line. In our current sample of 44 specimens, pale paravertebral line, number of scales around the we found a wide variety of curvature among the midbody, and division of palpebral eye scales. La Marca samples, without a clear trend that correlates with

Figure 6. Five patterns of prefrontal scales found in Cercosaura vertebralis. (A) No suture between prefrontal scales (AMNH R-60591); (B) prefrontal scales absent (MUSM 31845); (C) long contact (AMNH R-119370); (D) point contact (AMNH R-60587); (E) anomalous three prefrontal scales (LACM 45019).

1199 Doan and Cusi | Geographic distribution of Cercosaura vertebralis locality. Some C. vertebralis specimens had very Authors Contribution Statement: did morphological descriptions of some specimens, and contributed straight lip lines (see Figure 3), while others curved to the writing. TMD did most of the morphological JCC conducted descriptions all field work, and to varying degrees. Therefore, we do not consider this comparisons and wrote most of the manuscript. character to be diagnostic for distinguishing between the species. A pale paravertebral line was found to be Acknowledgments: J. Cusi thanks J. Cordova for allowing access to absent in C. ampuedai and present in C. vertebralis MUSM. We thank Ing. D. Cotrina for facilitating the collecting permits by La Marca and García-Pérez (1990), Hernández funded by a grant from the Rufford Small Grants Foundation. J. Cusi is Ruz (2005), and Hernández Ruz and Bernal González sincerelyand the parkthankful rangers to V. Vargas, for the F. assistance Vargas, C. Ramirez, in the field. and FieldworkA. Mendoza was for supporting and participating in the project at SNTN. Nanette Vega kindly (2011). Mijares-Urrutia (2000) found that C. ampuedai helped with the habitat description. We would also like to thank the was variable in this character, with some specimens museum personnel who allowed us to examine specimens under their having it and some not. In our current sample of C. care including Charles. J. Cole and David A. Kizirian at AMNH and Neftali vertebralis we found the paravertebral line present Camacho at LACM. in all specimens except for the one from Piura, Peru. Literature Cited La Marca and García-Pérez (1990) and Mijares- Amanzo J., R. Acosta, C. Aguilar, K. Eckhardt, S. Baldeón and T. Pequeño. Urrutia (2000) stated that scales around the midbody 2003. Evaluación Biológica Rápida del Santuario Nacional Tabaconas Namballe y Zonas Aledañas. Lima: Informe WWF–OPP: of C. ampuedai (26–31) are lower than C. vertebralis QM-91. 212 pp. (31–45). However, the specimen from Piura, Peru had Boulenger G.A. 1885. Catalogue of the Lizards in the British Museum 29 midbody annuli (Uzzell, 1973, counted 28) and the (Natural History) Volume II. London: British Natural History Museum. 497 pp. (doi: 10.5962/bhl.title.21097). Colombian C. vertebralis sample of Hernández Ruz and Boulenger G.A. 1908. Descriptions of new batrachians and reptiles Bernal González (2011) encompassed the entire scale discovered by Mr M. G. Palmer in south-western Colombia. range of both putative species (27–46), which discards Annals and Magazine of Natural History 8(2): 515–522 (http:// that character as diagnostic. Hernández Ruz (2005) biodiversitylibrary.org/page/26494615). and Hernández Ruz and Bernal González (2011) stated genera Cercosaura, Pantodactylus, and Prionodactylus (Reptilia: that the palpebral eye disc is undivided in C. ampuedai Doan,Squamata). T.M. 2003. AZoological new phylogenetic Journal classificationof the Linnean for Society the gymnophthalmid 137(1): 101– but divided in Mijares-Urrutia (2000) 115 (doi: 10.1046/j.1096-3642.2003.00043.x). C. vertebralis. Doan, T.M. and W.W. Lamar. 2012. A new montane species of Cercosaura found divided palpebrals in his C. ampuedai sample. In (Squamata: Gymnophthalmidae) from Colombia, with notes on the our sample of C. vertebralis all specimens had divided distribution of the genus. Zootaxa 3565: 44–54 (urn:lsid:zoobank. palpebrals except the one specimen from Piura, Peru. org:pub:40C56FA5-1827-4F8F-B236-FCE0D3AFAB53). Because all of the five putative diagnostic characters Segunda parte: reptiles, orden de los Saurios. Caldasia 3(11): 73–110. to distinguish Cercosaura ampuedai from C. vertebralis Dunn,Hernández E.R. Ruz, 1944. E.J. Los 2005. géneros Notas detaxonómicas anfibios y y reptilesbiológicas de de Colombia, Cercosaura II. ampuedai (Lancini, 1968) (Squamata: Gymnophthalmidae) en are variable within and among the putative species, la vertiente oriental de la Cordillera Oriental de Colombia. we designate C. ampuedai as a junior synonym of C. Publicações Avulsas do Instituto Pau Brasil de Historia Natural (São vertebralis, following Uzzell (1973), Doan (2003), Paulo) 8–9: 1–14. and Doan and Lamar (2012). Thus, the distribution of Hernández Ruz, E.J. and C.A. Bernal González. 2011. Variación morfológica en Cercosaura vertebralis (Sauria: Gymnophthalmidae) en Colombia. C. vertebralis includes populations in Peru, Ecuador, Ingenierías & Amazonia 4(1): 48–57. Colombia, Panama, and Venezuela (Figure 1). INRENA. 2007. Plan Maestro del Santuario Nacional Tabaconas Namballe Finally, we can point out that the SNTN populations 2007–2011. Lima: INRENA. 265 pp. La Marca, E. and J.E. García-Pérez. 1990. Prionodactylus ampuedai of C. vertebralis are relatively stable and abundant, (Sauria: Teiidae): a new lizard record for Colombia, with taxonomic because in 2011 (unpublished data, N=9) and 2012– comments on the species. Bulletin of the Maryland Herpetological 2013 (this article, N=4) it was possible encounter a Society 26(3): 111–115. Lancini A.R. 1968. El género Euspondylus (Sauria: Teiidae) en Venezuela. large number of individuals of this species at SNTN. Publicaciones Ocasionales del Museo de Ciencias Naturales, Zoología Tabaconas Namballe is a natural protected area where 12: 1–8. additional surveys could provide information about Mijares-Urrutia, A. 2000. Taxonomía de algunos microtéidos de Venezuela (Squamata), II: situación nomenclatural de Prionodactylus its natural history and ecology, including long-term ampuedai y Prionodactylus phelpsorum. Revista de Biología Tropical monitoring of this species. On the other hand, La Granja 48(2/3): 681–688 (http://www.scielo.sa.cr/scielo.php?script=sci_ arttext&pid=S0034-77442000000200041&lng=en&nrm=iso). is a village with many anthropogenic impacts, where O’Shaughnessy A.W.E. 1879. Descriptions of new species of lizards in the habitat loss caused by agriculture and cattle grazing are collection of the British Museum. Annals and Magazine of Natural History factors that appear to disturb the population levels of 5(4): 295–303 (http://biodiversitylibrary.org/page/25159870). these lizards. Pesantes, O. 1994. A method for preparing the hemipenis of preserved snakes. Journal of Herpetology 28(1): 93–95 (doi: 10.1670/10-139). Specimens examined: : Antioquia: 10 km W Ruthven A.G. 1921. Description of an apparently new lizard from Andes (town) (LACM 72790), San Pedro (AMNH R-32737– Colombia. Occasional Papers of the Museum of Zoology University of 32744), Santa Rita (LACMColombia 45017), Sonson (AMNH Michigan 103: 1–4 (http://hdl.handle.net/2027.42/56542). Uzzell, T.M. 1973. A revision of lizards of the genus Prionodactylus, with R-32725–32736); Cundinamarca: above Sasaima on a new genus for P. leucostictus and notes on the genus Euspondylus Bogota-Honda road (LACM 45018–45019); : El (Sauria, Teiidae). Postilla 159: 1–67 (http://biodiversitylibrary.org/ : El Chiral (AMNH R-18312); Mera, Río Pastaza page/12179719). Oro Pastaza: Zaher H. and A. Prudente. 2003. Hemipenes of Siphlophis (Serpentes, (AMNH R-60587, 60589, 60591–60597); Ecuador: Darién: Xenodontinae) and techniques of hemipenial preparation in snakes: SW sector Cerro Tacarcuna massif (AMNH R-119367), a response to Dowling. Herpetological Review 34(4): 302–307. Cerro Malí (AMNH R-119368, 119370); Panama: Cajamarca: Santuario Nacional Tabaconas Namballe (MUSM 31844– 31846, 31862), Caserio La Granja (MUSMPeru 32489–32490); May 2014 August 2014 Piura: 11 miles E Canchaque (on road), west slopes of Received: October 2014 Huancabamba Mountains (LACM 58811). Accepted: Ross MacCulloch Published online: Editorial responsibility: 1200