Distribution extension for Anolis pseudokemptoni Köhler , Ponce , Sunyer & Batista , 2007 ( Reptilia : Squamata : Dactyloidae ) , a microendemic species in the Serranía de Tabasará of the Comarca Ngöbe-Buglé of western Panama

Köhler et al. (2007) described Anolis pseudokemptoni on the basis of three specimens from “La Nevera, 8°29′45′′ N, 81°46′35′′ W, 1,600 m elevation, Serranía de Tabasará, Comarca Ngöbe Bugle, Distrito de Nole Düima, Corregimiento de Jadeberi, Panama.” Their slightly erroneous coordinates, also applied to the type localities of A. datzorum Köhler, Ponce, Sunyer & Batista, 2007 and A. pseudopachypus Köhler, Ponce, Sunyer & Batista, 2007, were later corrected to 8°30′ N, 81°46′20′′ W (Lotzkat et al. 2010a). The type locality (Figure 1, locality 1; Figure 2E and F) is situated on the continental divide of the Serranía de Tabasará, which constitutes the eastern continuous (unbroken above 1,000 m) portion of the Cordillera Central of western Panama. Ponce and Köhler (2008), at the time of their morphological revision of the Panamanian highland anoles related to A. kemptoni Dunn, 1940, still had only the type series of A. pseudokemptoni available. During recent field work in the highlands of western Panama, we collected 20 additional specimens of A. pseudokemptoni from several localities east and west of the type locality, which considerably improve our knowledge of the distribution and morphological variation of this little-known species. All specimens were encountered during opportunistic searches performed primarily at night, captured by hand, and preserved the day after capture. The collecting permits SE/A-30-08, SC/A-8-09, SC/A-28-09, and SC/A-21-10, as well as the corresponding exportation permits, were issued by the Dirección de Áreas Protegidas y Vida Silvestre of the Autoridad Nacional del Ambiente (ANAM), Panama City, Panama. Our specimens have been deposited in the collection of the Senckenberg Forschungsinstitut Frankfurt (SMF) and in the Museo Herpetológico de Chiriquí (MHCH) of the Universidad Autónoma de Chiriquí, Davíd, Chiriquí, Abstract: We report new localities for Anolis pseudokemptoni Köhler, Ponce, Sunyer & Batista, 2007, along the Serranía de Tabasará in the Comarca Ngöbe-Buglé of western Panama. These records extend the known geographic distribution of this lizard about 6 km eastward and 9 km northwestward, as well as the known vertical distribution approximately 470 m lower and 380 m higher. We also provide photos of specimens from different localities and comment on their morphology. No population of this Panamanian microendemic species, which is classifiable as Critically Endangered, lives inside a protected area. 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 Instituto de Ciencias Ambientales y Desarollo Sostenible, Universidad Autónoma de Chiriquí, Estafeta Universitaria, David, Chiriquí, Panamá. * Corresponding author. E-mail: lotzkat@yahoo.com Sebastian Lotzkat 1,2*, Abel Batista 1,2,3, Marcos Ponce 3 and Andreas Hertz 1,2 Distribution extension for Anolis pseudokemptoni Köhler, Ponce, Sunyer & Batista, 2007 (Reptilia: Squamata: Dactyloidae), a microendemic species in the Serranía de Tabasará of the Comarca Ngöbe-Buglé of western Panama


Distribution extension for Anolis pseudokemptoni
, a microendemic species in the Serranía de Tabasará of the Comarca Ngöbe-Buglé of western Panama Panama.Species identification was carried out employing the keys, figures, and descriptions provided by Köhler et al. (2007), Köhler (2008), and Ponce and Köhler (2008), and by comparison with the type series .All specimens strongly resemble the type specimens of Anolis pseudokemptoni and match the descriptions in Köhler et al. (2007) and Ponce and Köhler (2008) in terms of external morphology and dewlap coloration.Adult specimens were sexed according to dewlap size and coloration, by eversion of hemipenes in males, and by presence or absence of enlarged postcloacal scales, the last also applicable to juveniles.Characters of external morphology were recorded according to the methodology described by Köhler et al. (2007) and Ponce and Köhler (2008).Snout-vent length (SVL) and tail length (TL) were measured using a ruler.Head length (HL), Head width (HW), snout length, and shank length measurements were made using calipers.Measurements and scale counts are summarized in Table 1.Colors and color codes (the latter in parentheses) follow Smithe (1975Smithe ( -1981)).Coordinates and elevation were recorded using Garmin GPS receivers with built-in altimeters.All coordinates are in WGS 1984 datum; elevations are rounded to the nearest 10 m.Within the Comarca Ngöbe-Buglé, place names typically are in Ngäbere, the indigenous language of the Ngöbe people for which different spellings are available and used.For these names, we follow the spelling used on the Mapa General de la República de Panamá and in the Atlas Nacional de la República de Panamá (IGNTG 2000;2008).
Figure 1 shows the type locality of Anolis pseudokemptoni and the localities reported herein.Exact locality information for all collected specimens is provided in the Appendix.From west to east, we provide the following new records for A. pseudokemptoni: On the southeastern slopes of Cerro Saguí (also known as Cerro Ratón), between 2 and 2.5 km north of the settlement Ratón, we collected one individual along the banks of Quebrada Juglí (Figure 1, locality 2), and three more ascending the ridge E of this river (localities 3 and 4).Descending the trail leading from the continental divide N to Río Cricamola and Kankintú, we collected several specimens at Terminal de Hacha on the continental divide (locality 5), at the upper limit of the village of Hacha, also known as Río Hacha or Quebrada de Hacha (locality 6), and along Río Flor that is also known as Río Flores (locality 7), observing many more between these points (unnumbered localities).On the western slopes of Cerro Santiago near the type locality La Nevera, we collected individuals on the continental divide (locality 8) and adjacent Caribbean slopes (locality 9).Two specimens were collected east of Cerro Santiago in the Pacific drainage, about 3 km north of Buabidí, on the trail leading from Río Rey to the continental divide (localities 10 and 11).
The localities reported herein extend the known geographic and elevational range of Anolis pseudokemptoni.The records from Cerro Saguí, approximately 9 km northwest of the type locality, now constitute the westernmost records for the species, and raise the upper elevational limit to 2,010 m.The individuals from Hacha at 1,130 m set the lower elevational limit for A. pseudokemptoni.The trail ascending the southeastern slopes of Cerro Santiago above Río Rey, approximately 6 km east of the type locality, now constitutes the easternmost locality reported for the species, and the only locality situated below the continental divide on the Pacific slopes of the Serranía de Tabasará.
Selected specimens are shown in Figure 3.The examination of our additional specimens allows for a more comprehensive assessment of the variation in external morphology.Most notably, the minimum number of loreal scales is lowered to 56 arranged in 6 loreal rows, revealing an overlap in these characters with Anolis fortunensis Arosemena & Ibañez, 1994, A. kemptoni, and A. gruuo Köhler, Ponce, Sunyer & Batista, 2007(Ponce and Köhler 2008;Lotzkat et al. 2012a).Consequently, the high number of loreal scales and rows considered unique among these four closely related species by Ponce and Köhler (2008) loses its value as a non-overlapping diagnostic character.However, animals with 70 or more loreal scales are still highly likely to be representatives of A. pseudokemptoni.The species remains readily distinguishable from A. gruuo by the red male dewlap (orange in A. gruuo), and from A. fortunensis and A. kemptoni by the large bilobed hemipenis bulging the base of the tail in adult males (smaller and unilobed in the other two species).In Table 1, we provide an updated version of the table of Ponce and Köhler (2008) for selected morphological characteristics of A. pseudokemptoni, based on all known specimens except MHCH 1335 (n=22).
As typical for anoles, the coloration among our specimens is rather variable.Especially noteworthy is that a conspicuously contrasting dark and pale banding of the tail, stated as a diagnostic character for Anolis gruuo (Köhler et al. 2007, Köhler 2008, Ponce and Köhler 2008) and reported to not occur in all specimens of that species by Lotzkat et al. (2012a), is also present in some specimens of A. pseudokemptoni (Figure 3A and B).Since we also observed this banding among other betaanoles in western Panama, it should not be relied upon for species identification.Furthermore, we observed variation in female dewlap coloration ranging from almost white over the typical yellow shown by most females to orange (Figure 3K-N).The coloration in life of the male SMF 89515 (Figure 3A and G) was recorded as follows: as cloacal region Tawny Olive (223C); posterior portion of tail Pearl Gray (81) with a suggestion of Pale Horn Color (92), with Dark Brownish Olive (129) transverse bars grading into Grayish Horn Color (91) ventrally; iris True Cinnamon (139); dewlap Spinel Red (108B) with a suggestion of Poppy Red (108A) posteriorly, Spectrum Red (11) anteriorly, with a Burnt Orange (116) anterior border and dirty white scales, bordered by Dark Neutral Gray (83) except for the marginals.
All individuals were encountered at night, sleeping on vegetation between 1 and 4 m above ground.The collection sites encompass different habitats (compare photographs in Figure 3A-H) ranging from roadsides, riverbanks, pastures, plantations, and secondary forest to apparently pristine premontane and lower montane moist or wet forest, including cloud forest.While we found the species as low as 1,130 m on the Caribbean (i.e., northern) slopes, and down to 1,390 m atop the continental divide, our only two records from the Pacific (i.e., southern) slopes are much higher at 1,560 and 1,660 m, respectively, while still very close to the continental divide.At lower elevations on the Pacific slope, we found A. gruuo instead of A. pseudokemptoni (Lotzkat et al. 2012a).The parapatric distribution of our collection sites of these two species suggests that they do not occur in syntopy with each other.Rather, A. gruuo seems to be a species of the Pacific drainage, living at premontane elevations up to around 1,530 m, whereas A. pseudokemptoni appears to occupy the continental divide and adjacent Caribbean slopes, only descending onto the Pacific slopes down to somewhere near the upper elevational limit of A. gruuo.
The documented range of Anolis pseudokemptoni stretches over only 13 airline km, or 16 km along the course of the continental divide, from about 81°49'30'' W to 81°43' W at premontane and lower montane elevations of 1,130-2,010 m of the Serranía de Tabasará, Cordillera Central, in western Panama.Future fieldwork most probably will expand these range limits, but we are convinced that this expansion will be minor and mostly extend the current limits vertically as well as further to the east in the Comarca Ngöbe-Buglé, following our reasoning for A. gruuo in this regard (Lotzkat et al. 2012a) and considering the fact that we did not find A. pseudokemptoni during our extensive surveys around Santa Fé, Veraguas.
Accordingly, while Anolis gruuo ranges east into Veraguas province (Lotzkat et al. 2012a), A. pseudopachypus is distributed westerly to the Fortuna area in Chiriquí province (Lotzkat et al. 2010b), and A. datzorum has been reported from Costa Rica (Köhler and Vargas 2010), A. pseudokemptoni is the only of the four species described from the central Tabasará range by Köhler et al. (2007) that remains known only from near its type locality in the Comarca Ngöbe-Buglé.Our findings suggest that A. pseudokemptoni is a true microendemic species restricted to the central Serranía de Tabasará around the region known as Cerro Colorado.Its extent of occurrence of only 32.35 km 2 and the continuing deforestation (resulting in a decline of the obviously preferred forest habitats) we observed in the region qualify A. pseudokemptoni for the IUCN category "Critically Endangered" (CR), according to criterion B1b (IUCN 2001).A similar, yet even smaller range is documented for the recently discovered rainfrog

Figure 1 .
Figure 1.Distribution map showing the known localities for Anolis pseudokemptoni in the central Serranía de Tabasará, Comarca Ngöbe Buglé (outlined in the overview map), western Panama: The white circle (1) represents the type locality, black circles the localities reported herein.

Figure 2 .Figure 3 .
Figure 2. Habitats of Anolis pseudokemptoni along the Serranía de Tabasará in western Panama: (A) Cerro Saguí, looking north into the valley of Quebrada Juglí from approximately 1,800 m; (B) upper limit of Hacha, looking south towards junction of Río Flor and Río Hacha, 1,130 m; (C) valley of Río Flor 2.6 km north-northwest of type locality, 1220 m; (D) valley of Río Flor 1 km north-northeast of type locality, 1,500 m; (E) looking east-northeast from the continental divide at La Nevera, 1,700 m, across the type locality towards Cerro Santiago; (F) type locality at La Nevera, cloud forest streambed at approximately 1,600 m; (G) continental divide near La Nevera, elfin forest at 1,810 m; (H) crossing Río Rey on trail to continental divide, 1,550 m.

Table 1 .
Selected measurements, proportions, and scale characters of 11 males and 11 females of Anolis pseudokemptoni.Range is followed by mean value and standard deviation in parentheses.For abbreviations see text.