Distribution extension for Leposoma rugiceps ( Cope , 1869 ) ( Squamata : Gymnophthalmidae ) in Panama , with first record from Bocas

The gymnophthalmid genus Leposoma Spix comprises small lizards that inhabit the leaf litter of tropical forests. Among the 16 species currently recognized within this genus of primarily South American distribution, only two are known to occur in Lower Central America (Köhler 2008; Pellegrino et al. 2011; Uetz 2011): Leposoma southi Ruthven and Gaige, 1924, and L. rugiceps (Cope, 1869). The latter species was described as Loxopholis rugiceps from “the Magdalena River region, New Grenada” (i.e., present day Colombia; Figure 1, locality 1). Later, Peters (1880) described Leposoma dispar from “Caceres am Cauca, Neu-Granada” (i.e., Caceres at the Cauca river, Colombia; Figure 1, locality 2). Ruibal (1952) included Cope’s genus Loxopholis in Leposoma Spix, and placed dispar in the synonymy of rugiceps, creating the new combination Leposoma rugiceps. The species is known from Colombia and central Panama (Köhler 2008; Arredondo 2010; Pellegrino et al. 2011; Uetz 2011). During recent field work in Bocas del Toro province of extreme western Panama, we encountered specimens of L. rugiceps at a locality far from the species’ documented range. Between 24 and 29 November 2009, we carried out baseline inventory work within the San San Pond Sak Wetland of International Importance (Ramsar site no. 611). Around 22:30 h. on 24 November 2009, AH, RDL, and SL collected two specimens of Leposoma rugiceps (SMF 90192 and MHCH 2340; Figure 2A–D) close to the facilities of the Asociación de Amigos y Vecinos de la Costa y de la Naturaleza (AAMVECONA) at Centro Boca San San (09°31’33” N, 82°30’36” W, 1 m asl; Figure 1, locality 7; Figure 3A, E–G), on the narrow peninsula east of the mouth of the San San river, located approximately one km from the latter and about 9 km north of the town center of Changuinola, Corregimiento de Changuinola, Distrito de Changuinola, Provincia de Bocas del Toro, Panama. The specimens were encountered just east of Centro Boca San San among leaf litter and debris within coastal vegetation including Cocos nucifera, Anacardium Abstract: Reporting on the first locality in Bocas del Toro province of extreme western Panama, we extend the known geographic distribution of the lizard Leposoma rugiceps (Cope, 1869) about 275 km westwards from the nearest locality in Panamá province. We provide photos of Panamanian specimens, comment on their morphology, and map the distribution of this binational endemism. 1 Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, 60325 Frankfurt am Main, Germany 2 Goethe-University, Institute for Ecology, Evolution & Diversity, Biologicum, Building C, Max-von-Laue-Straße 13, 60438 Frankfurt am Main, Germany 3 Universidad de Panamá, Centro Regional Universitario de Bocas del Toro, Panama * Corresponding author. E-mail: lotzkat@yahoo.com Sebastian Lotzkat 1,2,*, Andreas Hertz 1,2, Rosalba De Leon 3 and Gunther Köhler 1 Distribution extension for Leposoma rugiceps (Cope, 1869) (Squamata: Gymnophthalmidae) in Panama, with first record from Bocas del Toro Province

The gymnophthalmid genus Leposoma Spix comprises small lizards that inhabit the leaf litter of tropical forests.Among the 16 species currently recognized within this genus of primarily South American distribution, only two are known to occur in Lower Central America (Köhler 2008;Pellegrino et al. 2011;Uetz 2011): Leposoma southi Ruthven and Gaige, 1924, and L. rugiceps (Cope, 1869).The latter species was described as Loxopholis rugiceps from "the Magdalena River region, New Grenada" (i.e., present day Colombia; Figure 1, locality 1).Later, Peters (1880) described Leposoma dispar from "Caceres am Cauca, Neu-Granada" (i.e., Caceres at the Cauca river, Colombia; Figure 1, locality 2).Ruibal (1952) included Cope's genus Loxopholis in Leposoma Spix, and placed dispar in the synonymy of rugiceps, creating the new combination Leposoma rugiceps.The species is known from Colombia and central Panama (Köhler 2008;Arredondo 2010;Pellegrino et al. 2011;Uetz 2011).During recent field work in Bocas del Toro province of extreme western Panama, we encountered specimens of L. rugiceps at a locality far from the species' documented range.
Between 24 and 29 November 2009, we carried out baseline inventory work within the San San Pond Sak Wetland of International Importance (Ramsar site no.611).Around 22:30 h. on 24 November 2009, AH, RDL, and SL collected two specimens of Leposoma rugiceps (SMF 90192 and MHCH 2340;Figure 2A-D) close to the facilities of the Asociación de Amigos y Vecinos de la Costa y de la Naturaleza (AAMVECONA) at Centro Boca San San (09°31'33" N, 82°30'36" W, 1 m asl; Figure 1, locality 7; Figure 3A, E-G), on the narrow peninsula east of the mouth of the San San river, located approximately one km from the latter and about 9 km north of the town center of Changuinola, Corregimiento de Changuinola, Distrito de Changuinola, Provincia de Bocas del Toro, Panama.
The specimens were encountered just east of Centro Boca San San among leaf litter and debris within coastal vegetation including Cocos nucifera, Anacardium Abstract: Reporting on the first locality in Bocas del Toro province of extreme western Panama, we extend the known geographic distribution of the lizard Leposoma rugiceps (Cope, 1869) about 275 km westwards from the nearest locality in Panamá province.We provide photos of Panamanian specimens, comment on their morphology, and map the distribution of this binational endemism.
During the same survey period, we collected two specimens of Leposoma southi (SMF 90193, shown in Figure 2E-G, and MHCH 2341), and observed others, on the northern banks of Río Negro (09°30'29" N, 82°31'44" W; Figure 3A-D).This locality is only 2.8 km southwest of Boca San San, but exhibits a completely different habitat in the form of a mostly flooded river swamp forest dominated by Sangruillo (Pterocarpus officinalis) and Matomba (Rhaphia taedigera), bordered by a narrow strip of mangrove towards the river.
All specimens were encountered during opportunistic searches and caught by hand.The day after capture, they were preserved after euthanasia by pericardial injection of T61 (Intervet).The collecting permit SC/A-28-09 and the corresponding export permits were issued by the Dirección de Áreas Protegidas y Vida Silvestre of the Autoridad Nacional del Ambiente (ANAM), Panama City, Panama.The specimens have been deposited in the collection of the Senckenberg Forschungsinstitut Frankfurt, Germany (SMF 90192,90193), and in the Museo Herpetológico de Chiriquí (MHCH 2340, 2341), Universidad Autónoma de Chiriquí, Davíd, Chiriquí, Panama.Species identification was carried out employing the keys, figures, and descriptions provided by Ruibal (1952), Uzzell and Barry (1971), and Köhler (2008).The capitalized colors and color codes (the latter in parentheses) provided for referenced specimens follow those of Smithe (1975Smithe ( -1981)).Coordinates and elevation were recorded in the field using Garmin GPS receivers with built-in altimeters.All coordinates are in WGS 1984 datum.For distributional records of Leposoma rugiceps, we consulted relevant literature and directly  (Ruibal 1952;Telford 1971;Ibañez et al. 1995;1996;1997 (Dunn 1933;Ibañez et al. 1996;1997"1995").
Boca San San is by far the westernmost locality ever reported for Leposoma rugiceps.Our record extends the species' distributional range in a west-northwesterly direction, approximately 275 km from Cirí, 280 km from each Valle de Antón and Sherman, and 290 km from Altos de Campana.It seems logical that L. rugiceps should also be present in the intervening area, namely the Comarca Ngöbe-Buglé and Veraguas province.Since Boca San San is situated about five km from the border of Costa Rica, we strongly suspect that yet another addition to the fauna list of that country awaits discovery.
Given the abundance of Leposoma rugiceps in the coastal vegetation at Boca San San, it seems surprising that this species has never been reported between this locality and central Panama -especially along the coast and on the islands of Bocas del Toro, where similar habitats abound, and considerable sampling has taken place.Instead, all records of Leposoma between western Colón province and the Río San San are of L. southi.We can only guess whether, throughout this distributional gap, L. rugiceps (a) is completely absent, (b) is present but has never been collected, or (c) has been collected, but misidentified as L. southi.Assuming that the population at Boca San San is autochthonous, possibility (a) seems highly unlikely, given the presence of suitable habitat throughout most, if not all, of the region.However, it cannot be completely ruled out that the population of L. rugiceps found at San San Pond Sak might have been introduced, possibly from Colón, since in one case a specimen has even reached the USA in this fashion (MCZ R-93421, "found on bananas shipped from Panama").The species' presence on the Pearl Islands (Köhler 2008) constitutes another case of successful overwater dispersal, be it natural or anthropogenic.Possibility (b) could be explained by the two congeners exhibiting different habitat preferences.Although they have been reported to occur in sympatry (Ibañez et al. 1995;1996) or even syntopy (Gutierrez and Arredondo 2007) at certain sites, our observations suggest that the two species inhabit different environments at San San Pond Sak.Not a single L. southi was found along the sandy For field assistance at Boca San San, we thank J.-F.Bienentreu, F. Ortlieb and A. Uselis.We were kindly invited to San San Pond Sak by Chiquita, GIZ, and REWE, and are most grateful for the opportunity to work there as well as for finding everything arranged for us.Lodging at the Centro Boca San San and river transportation were provided by members of the Asociación de Amigos y Vecinos de la Costa y de la Naturaleza (AAMVECONA), whom we owe special thanks for their exceptional hospitality.This paper is based upon work funded to AH by the FAZIT-Stiftung, and to SL by the Studienstiftung des deutschen Volkes as well as the Vereinigung von Freunden und Förderern der Goethe-Universität.coastal strip at Boca San San, and no L. rugiceps in the swamp forest at Río Negro, indicating a sympatric, but not syntopic occurrence of the two species.However, since our surveys were of short duration and our samples are small, additional sampling is required to reliably assess this pattern of presumed ecological separation.Possibility (c) cannot be ruled out completely, although the species are distinguishable morphologically.The most conspicuous differences are the frontonasal plate (single in L. rugiceps, divided in L. southi; Figures 2D, G) and the lateral neck scales between ear opening and shoulder (imbricate and keeled, resembling the dorsals in L. rugiceps; non-imbricate and conical in L. southi; Figures 2C, F).However, the two species are very similar at first glance (Figures 2A, B, and E), so an occasional L. rugiceps might have been included in a series of L. southi.This requires reexamination of existing collections of L. southi as well as targeted sampling for L. rugiceps, preferably focussing on coastal vegetation, in Bocas del Toro and Veraguas provinces and the Comarca Ngöbe-Buglé.

Figure 2 .
Figure 2. Individuals of Leposoma from San San Pond Sak.(A-D) L. rugiceps: (A) MHCH 2340; (B) entire specimen, (C) close-up of head and neck, and (D) dorsal view of snout showing single frontonasal plate of SMF 90192; (E-G) L. southi: (E) entire specimen, (F) close-up of head and neck, and (G) dorsal view of snout showing divided frontonasal plate of SMF 90193.

Figure 3 .
Figure 3. Habitats of Leposoma at San San Pond Sak.(A) satellite image showing the collection localities (X) for L. rugiceps and L. southi at San San Pond Sak; (B-D) north bank of Río Negro where L. southi occurs: (B) view of Sangruillo forest with narrow strip of mangrove in the left; (C) roots of Rhizophora close to the river (D) roots of Sangruillo inside the forest; (E-F) coastal vegetation around Boca San San where L. rugiceps occurs: (E) view from the Río San San onto Centro Boca San San; (F) view east along the peninsula from the Centro; (G) manioc and sugar cane in the shrub layer.