AMERICAN MUSEUM NOVITATES Number 3763, 19 pp. November 21, 2012 Discovery of the frog genus Anomaloglossus in Panama, with descriptions of two new species from the Chagres Highlands (Dendrobatoidea: Aromobatidae) CHARLES W. MYERS,1 ROBERTO IBANEZ D.,2 TARAN GRANT,3 AND CESAR A. JARAMILLO4 ABSTRACT The occurrence in Panama is documented for the South American frog genus Anomaloglossus (Dendrobatoidea: Aromobatidae). Two species are described from a low, forested uplift in east- central Panama, just northeast of Panama City. These low mountains, unnamed on maps, are designated the “Chagres Highlands” because a large part of the uplift lies in the Rio Chagres drain¬ age (which provides water critical to lock operation in the Panama Canal). The Chagres Highlands may be a lower montane forest refuge for some amphibians and reptiles, including the two Anom¬ aloglossus and Atelopus limosus, and the rare snakes Atractus depressiocellus, A. imperfectus, Geophis bellus, and Rhadinaea sargenti. Several other rare species are not endemic but include the Chagres Highland area as part of their individually fragmented or mosaic distributions (Adinobates fulgu- ritus, Anolis kunayalae, Coniophanes joanae, Geophis bracycephalus, Dipsas nicholsi). The two new frogs are at least broadly sympatric in the Chagres Highlands, but both spe¬ cies are rare. Anomaloglossus astralogaster, new species, is known only from the adult female holotype (22 mm SVL). Its ventral surfaces are covered overall with whitish dots (< 0.1 mm) somewhat similar to large chromatophores but possibly glandular; there is no appearance of 1 Division of Vertebrate Zoology (Herpetology), American Museum of Natural History. 2 Smithsonian Tropical Research Institute, Balboa, Republic of Panama, and Departamento de Zoologia, Universidad de Panama, Republic of Panama. 3 Division of Vertebrate Zoology (Herpetology), American Museum of Natural History, and Departamento de Zoologia, Instituto de Biociencias, Universidade de Sao Paulo, Brazil. 4 Facultad de Medicina, Departamento Histologia y Neuroanatomia Humana, Universidad de Panama, Republic of Panama, and Circulo Herpetologico de Panama, Republic of Panama. Copyright © American Museum of Natural History 2012 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3763 glandular structure at x50 magnification, but the edges of some of the pale dots can be “felt” with a fine (0.1 mm diameter) teasing needle and histological examination is needed. The other taxon is Anomaloglossus isthminus, new species, which is described from six speci¬ mens including four adult males (19-21 mm SVL), one adult female (23 mm SVL), and a juvenile female. Dorsal surfaces are basically brown mottled with darker brown. Small pale yellowish spots located proximally above the insertions of arm and thigh are not well defined and tend to disap¬ pear after preservation (unlike normal dendrobatid flash markings). Ventral surfaces are pale blue with some dark mottling but no pale dots. The vocalization of A. isthminus resembles calls of some South American species in being a train of “peeplike” notes, but there are fundamental interspe¬ cific differences in frequency modulation, note repetition rate, and call length. INTRODUCTION The Isthmus of Panama harbors a rich fauna of dendrobatoid frogs that have both northern (Central American) and southern (South American) affinities. Endemism is high in western Panama among both elements of this fauna. In comparison, there have been relatively fewer speciation events among dendrobatoids in eastern Panama (although we are aware of unnamed cryptic species). In this paper we describe two species with South American affinities from the east-central part of the isthmus—in the low highlands east of the Panama Canal (map 1). Myers et al. (2007: 14) commented that “It seems very odd that the low highlands northeast of Panama City have never received a widely accepted map name (usually no name is given, although ‘Sierra Maestra has been shown for part).” The names used in the herpetological literature (Pequeni-Esperanza Ridge, Piedras-Pacora Ridge, and Serranfa Piedras-Pacora) are descriptive only of the southern parts of this uplift, much of which is drained by headwaters of the Rfo Chagres and also included in the Chagres National Park. This protected watershed is essential for operating the locks that raise and lower ships for passage through the Panama Canal. We propose the name Chagres Highlands for this physiographic entity, although some of its small rivers drain directly north to the Caribbean Sea or south to the Bay of Panama. The relatively broad expanse of the Chagres Highlands is geologically complex (Montes et al., 2012). It is abutted immediately on the east by the Serranfa de San Bias—a narrow, broken range that carries the continental divide southward to the Serranfa del Darien on the Colombian border. The Chagres Highlands may be a lower montane forest refuge for some rare amphibians and reptiles (see Discussion). The first species described in this paper was discovered in 1974 by Catherine A. Toft. She collected a specimen from near the El Llano-Cartf Road, which crosses the low divide between the Chagres Highlands and the Serranfa de San Bias (see easternmost symbol in map 1). The next specimens were not collected until 1997, when Ibanez and Jaramillo discovered specimens at widely spaced localities in the Chagres Highlands (map 1). Toft’s specimen was identified by Myers (1991) as Colostethus chocoensis (Boulenger), a species then known only from its type locality in Pacific western Colombia. Myers also tenta¬ tively extended the range of C. chocoensis south into Ecuador. Later, however, discovery of the “median lingual process” in New World dendrobatoids revealed that this curious structure is 2012 MYERS ET AL.: ANOMALOGLOSSUS IN PANAMA 3 ecrw 7rvv Caribbean Sea Jf' < £7&$i’ 1 Jr, A f^F- •> m wm? -v ■ ‘s '' 9°H - .t* ^ PAM AM A CITY i-' Scry of Panama *££-i AT ■ t Yr KM 0 5 10 “o 50 40 MAP 1. Central Panama, showing locality records for two new species of Anomaloglossus in the Chagres Highlands. Black square, A. astralogaster. Black circles, A. isthminus (middle circle is the type locality). The Chagres Highlands comprise the upland northeastward of Panama City—between the Panama Canal (visible left of center) and the Serrania de San Bias (to the right of the easternmost A. isthminus locality). lacking in “Colostethus” (now Hyloxalus) chocoensis, but that it is present in the Ecuadorian frogs (Grant et al., 1997: 24, fn. 13) and also in Toft’s Panamanian specimen (this paper). The systematics of “Colostethus” sensu lato had seemed nearly intractable, but phylogenetic analyses of DNA sequences and morphological and behavioral characters recovered monophyletic groupings, and the lingual process provided a morphological synapomorphy for the new genus Anomaloglossus (Grant et al., 2006: 158). The Ecuadorian frogs allocated by Myers to chocoensis were named Anomaloglossus confusus (Myers and Grant, 2009) and the Panamanian frogs are treated in this paper. The first species of Anomaloglossus to be named is the one collected by Toft, Ibanez, and Jaramillo, as mentioned above. To our surprise, however, a second new species of Anomaloglos¬ sus also occurs in the Chagres Highlands. It was collected during an unpublished biological survey (Roldan, 1985). This second species is known only from a single specimen, and it has a unique ventral pattern of pale chromatophore-like (glandular?) dots that we do not recall seeing in any other frog. 4 AMERICAN MUSEUM NOVITATES NO. 3763 The type localities for the two new species are on Cerro Brewster, but on different sides of the political division between Provincia de Panama and Comarca Guna Yala.5 Methods: Measurements of specimens, hind-foot webbing formulae, and analysis of vocalizations follow the methods most recently discussed in Myers and Donnelly (2008: 12-13). Ocular-micrometer measurements labeled “sagittal” (from tip to snout to center of naris or to angle of jaw) are made in the plane parallel to the median or sagittal plane of the body Coordinates at the type locality of Anomaloglossus isthminus were determined by a hand¬ held GPS satellite receiver in 2009, after military restrictions had been lifted. (Prior to 2000, the point-position accuracy of commercially available GPS devices was limited to about 100 m). Coordinates for Cerro Bruja (a paratype locality) were taken in 1997 and are believed to be reasonably accurate because the reading coincided with map topography. Coordinates for the type locality of Anomaloglossus astralogaster are estimates based on information supplied by the collector, Jorge Roldan H. The datum is WGS84 for all coordinates in this paper. The locality map is based on a digital elevation model generated from the SRTM (Shuttle Radar Topography Mission) prepared by NASA and NGA (National Geospatial Intelligence Agency). Anomaloglossus isthminus, new species Figures 1A-D, 2-8; map 1 Colostethus chocoensis (Boulenger): Myers, 1991 (part: the single Panamanian specimen, KU 172790; figs. 3C, D, 4, 8D). Grant et al, 1997: 24, fn. 13 (part: only KU 172790; see Remarks). Holotype: MVUP 2345 (originally CH 4568), an adult male from forest stream at head¬ waters of Rio Piedras, Cerro Brewster, 810 m elevation, Provincia de Panama (9°19'17" N, 79°17'10" W), collected by Cesar A. Jaramillo, February 24, 1997. Paratopotypes (3): CH 4566, a juvenile female from forest stream, collected by Cesar A. Jaramillo, February 24, 1997. CH 4552 and 4557 (=AMNH A-187134), adult males from forest stream, collected by Cesar A. Jaramillo, February 25, 1997. Paratypes (2): CH 4512, an adult female from forest stream at headwaters of Rio Pie¬ dras, SSW slope of Cerro Bruja, 720 m elevation, Provincia de Colon (9°27'35" N, 79°34'29" W), collected by Roberto Ibanez, January 25, 1997. KU 172790, an adult male from forest stream west and down slope (150 m elev.) from km 11.7 (300 m) on road from El Llano to Carti, Provincia de Panama (approximately 9°18' N, 78°59' W), collected by Catherine A. Toft, July 1974). 5 This Comarca was shown on 20th century maps as the Comarca de San Bias, but it was changed to the “Comarca Kuna Yala” in the late 1990s. Apparently the letter “K” was officially removed from the Guna alphabet and in 2010 the name was again changed, to “Comarca Guna Yala” (or “Gunayala”). (See Ley fun¬ damental de la Comarca Gunayala available through http://www.congresogeneralkuna.com [accessed July 19, 2012]). See also Price (2005: 176). The word Kuna (or Cuna) is in wide usage, so this is a major name change for this comarca, although borders are only slightly affected. But possible elevation to complete pro¬ vincial status for other comarcas indigenas have profoundly changed the map of Panama, particularly as regards the provinces of Darien, Bocas del Toro, and Chiriquf. 2012 MYERS ET AL.: ANOMALOGLOSSUS IN PANAMA 5 FIG. 1. Two Panamanian species of Anomaloglossus in dorsal and ventral views. A-D: Anomaloglossus isthmi¬ nus, new species (left to right: CH 4512$, 4552 d, 4557 d [AMNH A-187134], MVUP 2345 d). E: Anomalo¬ glossus astralogaster, new species (MVUP 2346$). Scale = 10 mm. Etymology: The specific name is derived from Latin isthmus (< Greek isthmos) + -inus (-a, -um), an adjective-forming suffix meaning “belonging to,” in reference to the Isthmus of Panama. Diagnosis: A low median lingual process (fig. 2) or median pit (similar to fig. 11) distin¬ guishes Anomaloglossus isthminus from all other named Central American frogs except A. astralo¬ gaster, which is differentiated by a punctate ventral pattern of discrete pale spots (see fig. 10). A. isthminus seems related to an unnamed species from the Colombian side of Cerro Tacarcuna (Grant et al., 2011). Based solely on a few color photographs, the Tacarcuna species differs from A. isthminus in having weak indications of a pale dorsolateral line and pale spotted, lighter sides; the lateral spots are aligned as a broken, oblique lateral stripe extending about halfway along the sides; the blue ventral surfaces with variable dark markings are very similar to A. isthminus. AMERICAN MUSEUM NOVITATES NO. 3763 Measurements (in mm) of Holotype: The specimen is an adult male (fig. 3) with open vocal slits. Length from snout to vent 20.6; tibia length between heel and outer surface of flexed knee 10.3; greatest width of body 9.5; greatest head width between angles of jaws and width between outer edges of upper eyelids 8.1 and 6.8, respectively; approximate width of interor¬ bital area 3.5; head length (sagittal) from tip of snout to angle of jaw 5.6; tip of snout to center of naris (sagittal) 1.3; center of naris to anterior edge of eye 1.5; distance between centers of nares 2.9; eye length from anterior to posterior edge 3.3; tympanum nearly concealed; hand length from proximal edge of large medial pal¬ mar tubercle to tip of longest (third) finger 5.5; width of disc of third finger (and width of pen¬ ultimate phalanx below disc) 1.0 (0.6); width of discs (and penultimate phalanges below discs) of third and fourth toes 1.1 (0.7) and 1.0 (0.7), respectively. FIG. 2. The median lingual process (arrow) in Anom- DESCRIPTION aloglossus isthminus, new species (MVUP 2345 holo¬ type), approximately x!3. The type series comprises six specimens from three localities in east-central Panama. Four adult males, one adult female, and one juvenile female (in poor condition) are included. Measurements and proportions given in the following description are for the five adults. Morphology: Adult males 19.4-21.3 mm SVL (n = 4, x= 20.33 ± 0.41 mm); testes unpig- mented, white, large (~1.8-2.2 mm long), roughly one-half of kidney length. Adult female (CH 4512) 22.6 mm SVL, with large, convoluted oviducts and large brown ova. A low median lin¬ gual process (fig. 2) discernible on tongue except that it is seemingly retracted and concealed in a pit in CH 4552. See Discussion. Skin in preservative smoothly rugose to faintly granular dorsally, smooth ventrally; no postric- tal or cloacal tubercles. Greatest head width (between angles of jaws) 35%-39% of SVL. Snout sloping, acutely rounded (nearly pointed) in profile, broadly rounded to nearly truncate in dorsal and ventral view. Nares visible from in front, barely visible from above or below. Canthus rostralis rounded to weakly angulate; loreal region slightly concave, sloping slightly outward to lip. Inter¬ orbital region not well defined but nonetheless noticeably wider than upper eyelid. Length of snout in lateral view much shorter (64%-79%) than eye length; center naris to edge of eye 50%-64% of eye length. Tympanic annulus and membrane virtually concealed in a few specimens; otherwise 2012 MYERS ET AL.: ANOMALOGLOSSUS IN PANAMA 7 weakly indicated antero- ventrally and concealed by diffuse supratympanic bulge posterodorsally; tympanum small, its esti¬ mated diameter less than one-half of eye length. Teeth present on maxilla. Hand (fig. 4) relatively small, its length in four adult males 26%-27% of SVL, 68%-74% of greatest head width; smaller hand in one adult female 23% SVL, 59% head width. Relative lengths of appressed fingers III > IV > II > I; tip of finger I either reaching or failing to reach disc of finger II; finger III not swollen. Discs of all fingers moder¬ ately expanded; third fin- FIG. 3. Anomaloglossus isthminus, new species. Dorsal and ventral views of ger disc 14-17 times ac^t ma^e holotype (MVUP 2345). Scale = 10 mm. wider than distal end of adjacent phalanx. Base of palm with a large, irregularly rounded, median metacarpal tuber¬ cle, a smaller elliptical inner metacarpal tuber¬ cle on base of first finger; one to three subarticular tubercles (one each on fingers I, II; two or three on fingers III, IV; an inconspicu¬ ous small, distal third subarticular tubercle sometimes distinguishable on fingers III and IV); all tubercles low, with slightly rounded surfaces. Fringes along sides of fingers weakly developed (virtually absent); proximal part of fringe on median side of first finger extending faintly to inner metacarpal tubercle; fringe on lateral side of fourth finger faintly continuous with a weak outer metacarpal fold that extends to the large palmar tubercle. AMERICAN MUSEUM NOVITATES NO. 3763 Hind limbs relatively long, with heel of forward-pressed limb reaching to snout tip or between eye and snout tip; tibia 48%-52% of SVL. Relative lengths of appressed toes IV > V ~ III > II > I; first toe reaching middle of subarticular tubercle of second. Toe discs noticeably expanded, those on third and fourth toes 1.3-1.8 times wider than adjacent phalanges. Feet “webbed” to base of each toe disc, although the web is distally narrowed to a fringe on medial sides of toes II and III and on both sides of toe IV, yielding an approximate webbing formula of I 1-2 II 1-2 III 1-2 IV 2-1 V (fig. 4). The fringes fold downward along each side of toe IV, forming “folded flaplike fringes.”6 A well-developed fringe along the outer free edges of toes I and V. One to three nonprotuberant subarticular tubercles; a small round to oval outer meta¬ tarsal tubercle and a slightly longer elliptical inner metatarsal tubercle. An oblique tarsal fold extending proximolateral on distal half of tarsus is continuous with fringe on free edge of first toe; tarsal fold varying from a weak ridge to a strong keel, lacking tubercle or pronounced elevation at its proximal end. Color in Life: Based on a few color photographs and the collectors’ color notes abstracted below, the dorsal body varies from brown with irregular dark brown or cream blotching to blackish-brown with greenish-gold reticulum. The aforesaid “greenish” dorsal reticulum—doc¬ umented in a color photo of the holotype—is more bronzy in another view of the same speci¬ men (fig. 5). Sides brown without conspicuous pale spots or oblique lateral stripe. A narrow, sometimes broken postocular line from below eye to arm insertion varies from “blue-gold” to bronze (fig. 5). There are either bright yellow or pale blue-gold marks atop base of the upper arm and atop thigh near groin (these markings appear very pale yellowish in color photos of the holotype). Venter light blue (fig. 6), with blackish reticulations on throat and scattered over abdominal area and limbs as described for the adult female below.] CH 4512 (adult female): Dorsum brown with some irregular darker blotches. A bright yellow spot in [near] the groin and at the dorsal insertion of upper arm. Venter light blue with a black reticulum This black reticulum extends to the belly, throat, arms and legs. CH 4552 (adult male): Iris dark brown. Two light blue-gold spots behind the eye, towards the insertion of arm. Dorsum brown with some irregular cream blotches. Anterior area of throat light blue with brown blotches, rest of throat brown. Chest, belly and anterior ventral surface of thighs light blue. Posterior ventral surface of thighs brown. Concealed surfaces of shanks and feet light blue. Dorsal surface of hind legs with light and dark brown bars. A light blue-gold spot on anterior portion of hind leg near the groin and at the dorsal insertion of upper arm. CH 4557 (adult male): A fine light blue-gold line posterior to eye. Dorsum brown with some irregular black blotches. Throat and belly light blue. The throat with a central brown blotch. Dorsal surface of thighs with dark brown and black bars. Anterior ventral surface of thighs light blue. A light blue-gold spot on anterosuperior portion of thigh near the groin and at the dorsal insertion of upper arm. 6 As defined by Myers and Donnelly (2008: 42). 2012 MYERS ET AL.: ANOMALOGLOSSUS IN PANAMA 9 FIG. 5. Anomaloglossus isthminus, new species. The adult male holotype. in life (MVUP 2345, 20.6 mm SVL) (photographed by Marcos Guerra at Smithsonian Tropical Research Institute). CH 4566 (juvenile female): Similar in color to CH 4557. Venter light blue with small brown blotches in the throat. MVUP 2345 (CH 4568, adult male): Iris light brown. A fine light blue-gold line posterior to eye, towards the insertion of arm. Dorsum dark brown (almost black) with a greenish gold reticulum. Throat brown with light blue blotches on anterior area of throat. Belly and anterior ventral surface of thighs light blue. Dorsal surface of thighs with brown and golden brown bars. Concealed surfaces of shanks light blue. A light blue-gold spot at the base (anterosuperior por¬ tion) of the hind leg and at the dorsal insertion of upper arm. Pericloacal area orange. KU 172790 (adult male): “chocolate brown above, with a yellow flash mark atop base of upper arm, and with a tinge of metallic blue on the venter” as given in Myers (1991: 5). 10 AMERICAN MUSEUM NOVITATES NO. 3763 FIG. 6. Anomaloglossus isthminus, new species. Ventral view of the adult male holotype in life (photographed by Marcos Guerra at Smithsonian Tropical Research Institute). Color Pattern in Preservative (figs. iA-D, 3): Dorsum pale to light brown, with irreg¬ ular spots and blotches of dark brown. Loreal region and upper lip suffused with brown. A distinct to indistinct, narrow whitish postocular line, which starts below lower rear edge of eye and extends obliquely (nearly horizontally in one) between tympanic area and corner of mouth to the arm. Flanks brown, lacking pale stripes; axilla and groin either brown or whitish, without pale flash marks. Limbs irregularly banded with dark and light brown; fingers I and II partly whitish. Rear of thighs brown without definite pattern. Ventral surfaces whitish, with brown blotching or brown reticulum under head, this pattern extending weakly to strongly onto chest; some inconspicuous small suffusions of pale brown on belly. Palms and soles brown. The golden marks atop the base of the upper arm and atop the thigh near the groin are essentially lost in preservative (fig. 1; Myers, 1991: fig. 3C), although a few specimens retain diffuse pale yellowish spots on the upper arm above axilla (fig. IB, ID). This is in contrast to many dendrobatoids that have bright axillary and/or inguinal flash marks that remain con¬ spicuous even in preservative. VOCALIZATION Anomaloglossus isthminus is diurnal and has been heard calling only by day. Tape record¬ ings were made by Jaramillo along a headwater branch of the Rio Piedras at the type locality, at a daytime air temperature of 21° C on February 25, 1997. Part of the 14-minute tape is of a specimen of A. isthminus calling from a rock in the stream, with consequential loud back¬ ground noise of flowing water; some other calling individuals were concealed. The 1-14 note vocalizations of another dendrobatoid, Colostethus pratti, also can be heard on the first part of