Hemiphractus panamensis, Duellman, 1970

Hill, Robert L., Martin, Kathryn G., Stanley, Edward & Mendelson Iii, Joseph R., 2018, A taxonomic review of the genus Hemiphractus (Anura: Hemiphractidae) in Panama: Description of Two New Species, Resurrection of Hemiphractus panamensis (Stejneger, 1917), and Discussion of Hemiphractus fasciatus Peters, 1862, Zootaxa 4429 (3), pp. 495-512 : 506-509

Hemiphractus panamensis

( FIgS. 3 , 4 , 6 , 7 , 9 )

Cerathyla panamensis Stejneger 1917 . Holotype: USNM 55320 . Type locality: Signal Loma ( Loma Peak ) on the north coast of Panama, three miles [4.8 km] south of Santa Isabel, Prov . Colón, Panama .

Hemiphractus fasciatus Trueb 1974 [in part; for reference to specimens from Chagres Highlands, Panama]; Duellman, 2001 [in part; for reference to specimens from Chagres Highlands, Panama]; Crawford et al. 2012 [in part, for samples from Cerro Brewster, Prov. Panama, Panama]; Castroviejo-Fisher et al. 2015 [in part, for samples from Cerro Bruja and Cerro Brewster, Prov. Panama, Panama]; Köhler, 2011 [figs. 472, 473; in part; for map showing isolated populations in the Chagres Highlands, Panama].

Re-description of the Holotype. MEDIUM-SIzED ADULT FEMALE, SVL 59.3 MM (NOT INCLUDINg PROBOSCIS); TIBIA LENgTh 31.2 MM (NOT INCLUDINg CALCAR); FOOT LENgTh 27.3 MM; hEAD LENgTh 26.6 MM (NOT INCLUDINg PROBOSCIS); hEAD wIDTh 31.4 MM; SNOUT–PARAOCCIPITAL PROCESS LENgTh 25.8 MM; TYMPANUM OVOID, 5.2 MM hIgh X 2.6 MM wIDE; EYE DIAMETER 7.4 MM; INTERORBITAL DISTANCE 12.0 MM; EYE–TYMPANUM DISTANCE 3.3 MM; BODY MODERATE; hEAD LARgE, wIDER ThAN BODY, gREATEST wIDTh AT LEVEL OF PARS ARTICULARIS, BEARINg PROMINENT PARAOCCIPITAL PROCESSES; PARAOCCIPITAL PROCESSES PROjECTINg POSTEROLATERALLY IN DORSAL VIEw, EXTENDINg POSTERIOR TO LEVEL OF PARS ARTICULARIS; SkIN CO-OSSIFIED OVER DORSAL SURFACE OF SkULL; OCCIPITAL RIDgES INDISTINCT, PARAOCCIPITAL RIDgES PRESENT, wEAkLY DEVELOPED; SUPRAORBITAL AND CANThAL RIDgES SLIghTLY DEVELOPED; zYgOMATIC AND PREORBITAL RIDgES ABSENT; CANThUS ROSTRALIS INDISTINCT; LOREAL REgION FLAT; PROBOSCIS ShORT, FLEShY, POINTED, TRIANgULAR IN CROSS SECTION; SNOUT ANgLINg POSTERIORLY AT APPROXIMATELY 45o FROM hORIzONTAL; NOSTRILS OVOID, SLIghTLY PROTUBERANT, PROjECTINg DORSALLY, SLIghTLY LATERALLY; LIPS NOT FLARED. TOP OF hEAD RELATIVELY FLAT; INTERORBITAL REgION 38 % hEAD wIDTh; EYE DIAMETER 24 % hEAD wIDTh; gREATEST DIAMETER 70 % TIMES EYE DIAMETER; wIDTh OF TYMPANUM 84 % TIMES EYE–TYMPANUM DISTANCE; UPPER EYELID APPEARINg TO BEAR ONLY A SINgLE ENLARgED POINTED PROjECTION, SCATTERED SMALLER TUBERCLES ABSENT; INTERORBITAL, POSTORBITAL TUBERCLES ABSENT; SkIN OVER POSTERIOR SURFACE OF PARS ARTICULARIS BEARINg ONE LARgE, FLEShY TUBERCLE. TONgUE LARgE, OVOID; MANDIBLE BEARINg SERRATIONS ALONg ENTIRE LENgTh OF LOwER jAw, wITh TwO PROMINENT, TUSk-LIkE ODONTOIDS PROjECTINg FROM MENTOMECkELIAN BONES. AXILLARY MEMBRANE ABSENT; ThORACIC FOLD ABSENT; DERMAL FOLDS ON wRISTS ABSENT; FINgERS LONg, SLENDER, BEARINg SLIghTLY ENLARgED DISTAL PADS; INNER OUTER METACARPAL TUBERCLES PRESENT, BUT CONDITION OF SPECIMEN DOES NOw ALLOw FOR ACCURATE DESCRIPTION; RELATIVE LENgThS OF FINgERS: III> IV> II> I; LATERAL FLANgES APPEAR TO BE ABSENT ON FINgERS; CONDITION OF SPECIMEN DOES NOT PERMIT DESCRIPTION, OR DETERMINATION OF PRESENCE/ ABSENCES OF TUBERCLES ON FOREARMS; wEBBINg ON hANDS ABSENT. HEELS OF ADPRESSED hIND LIMBS OVERLAP; TIBIOTARSAL ARTICULATION NOT EXTENDINg TO SNOUT whEN hINDLIMB EXTENDED; CONDITION OF SPECIMEN DOES NOT PERMIT DESCRIPTION OF TARSAL FOLDS; TIBIA LENgTh 53 % SVL; FOOT LENgTh 46 % SVL; INNER AND OUTER METATARSAL TUBERCLES PRESENT, BUT CONDITION IS UNCLEAR; PRESENCE OF CONDITION OF LATERAL FRINgE ON TOES UNCLEAR; CALCAR PRESENT, SMALL; RELATIVE LENgThS OF TOES: IV> V> III> II> I; wEBBINg BASAL. CLOACAL OPENINg DIRECTED POSTEROVENTRALLY AT MIDLEVEL OF ThIgh.

TEXTURE AND COLORATION OF ThE SkIN, AS wELL AS ThE FINER DETAILS OF ThE hANDS AND FEET ARE UNCLEAR, AS ThE TYPE SPECIMEN IS qUITE DESICCATED. SIMILARLY, ThE DESICCATED NATURE OF ThE hOLOTYPE MAkES IT UNCLEAR ThE LEVEL TO whICh ThE TRANSVERSE PROCESSES OF ThE VERTEBRAE ARE CO-OSSIFIED wITh ThE SkIN. ThE PUPIL IS NOT VISIBLE. IN ALL MAjOR MORPhOLOgICAL FEATURES AND MEASUREMENTS, STEjNEgER’S (1917) DESCRIPTION CLOSELY MATChES ThE CURRENT CONDITION OF ThE hOLOTYPE (USNM 55330). EXCEPTIONS INCLUDE COLORATION NOTES AND TEXTURAL DETAILS OF ThE SkIN, VARIOUS TUBERCLES, AND DIgITAL PADS ( STEjNEgER 1917: 32), ALL OF whICh ARE NOw FADED OR PhYSICALLY DISTORTED—EVIDENTLY AS A RESULT OF DESICCATION AT SOME POINT IN ITS hISTORY. LIkEwISE, ThE INDENTATIONS OF EggS ON ThE DORSAL SURFACE OF ThIS FEMALE ThAT STEjNEgER (1917: 33) DESCRIBED NO LONgER ARE EVIDENT.

Diagnosis. Hemiphractus panamensis MAY BE DISTINgUIShED FROM ALL SPECIES OF Hemiphractus EXCEPT H. elioti , H. fasciatus , H. kaylockae , AND H. scutatus BY NOT hAVINg A POSTORBITAL INDENTATION (PRESENT IN OThER SPECIES). Hemiphractus panamensis MAY BE DISTINgUIShED FROM H. scutatus , AN AMAzONIAN SPECIES NOT OCCURRINg IN PANAMA, BY ITS SMALLER SIzE (MAXIMUM SVL IN MALES 52.5 VS. 57.4, FEMALES 64.7 VS. 80.5; DATA FOR H. scutatus FROM TRUEB 1974), AND BY hAVINg SMALL EXPANDED PADS ON ThE FINgERS AND TOES (ABSENT IN H. scutatus ). Hemiphractus panamensis DIFFERS FROM H. elioti AND H. kaylockae BY hAVINg NEOPALATINE AND VOMERINE ODONTOIDS wIDELY SEPARATED (VS. IN CONTACT IN H. elioti AND NARROwLY SEPARATED IN H. kaylockae ; FIg. 6 ). Hemiphractus panamensis APPEARS TO hAVE ONLY ABOUT TwO ODONTOIDS ON EACh VOMER, whEREAS H. elioti AND H. kaylockae BOTh hAVE ABOUT SEVEN (CONDITION UNkNOwN IN H. scutatus ; FIg. 6 ). Hemiphractus panamensis MAY BE DISTINgUIShED FROM H. kaylockae BY hAVINg A STRAIghT LATERAL MARgIN OF ThE PARAOCCIPITAL hORNS ( FIgS. 2 , 9 ); SUPRAORBITAL RIDgES EVIDENT, wEAkLY DEVELOPED (ABSENT OR wEAkLY DEVELOPED IN H. elioti AND H. kaylockae ; FIgS. 2 , 9 ); CANThAL RIDgES EVIDENT, DISTINCT (ABSENT OR wEAkLY DEVELOPED IN H. elioti ; FIgS. 2 , 9 ). Hemiphractus panamensis hAS A VERY LARgE SUBTEMPORAL FENESTRA APPROXIMATELY 1.5 X SIzE OF ThE ORBIT IN FEMALES (MALES UNkNOwN), SUBTEMPORAL FENESTRA APPROXIMATELY ONE-ThIRD DIAMETER OF ThE ORBIT IN BOTh MALES AND FEMALES IN H. kaylockae , APPROXIMATELY ONE-hALF DIAMETER OF ORBIT IN FEMALES OF H. elioti (VERY SMALL IN MALES), APPROXIMATELY ONE-hALF DIAMETER OF ThE ORBIT IN FEMALES OF H. fasciatus (MALES UNkNOwN); LATERAL MARgINS OF ThE qUADRATOjUgAL EXTENDINg FAR BEYOND ThE PROFILE OF ThE PARAOCCIPITAL hORNS IN DORSAL VIEw (ABOUT TO LEVEL OF PARAOCCIPITAL hORNS, OR SLIghTLY BEYOND, H. elioti , H. fasciatus , AND H. kaylockae ).

Distribution and ecology. Hemiphractus panamensis IS kNOwN FROM SPECIMENS COLLECTED FROM CENTRAL PANAMA IN ThE PROVINCES OF COLON AND SAN BLAS wIThIN OR IN PROXIMITY TO whAT IS NOw ThE ChAgRES NATIONAL PARk ( FIg. 5 ). SPECIMENS hAVE BEEN COLLECTED AT ~750 M ELEVATION IN LOwER MONTANE FORESTS wITh hIgh RAINFALL ( MYERS 1966). TRUEB (1974) REPORTED A LARgE SPECIMEN (KU 93503) ThAT CONTAINED “…AN UNIDENTIFIED gASTROPOD, A LIzARD ( Ptychoglossus ), AND A DENDROBATID FROg ( Colostethus ) IN ThE STOMACh. MYERS (1966) REPORTED H. panamensis TO DISPLAY MOUTh gAPINg BEhAVIOR AND BITINg whEN ACCOSTED ( FIg. 3B ). AN INDIVIDUAL COLLECTED BY MYERS IN 1964 AT

ALTOS DE PACORA ( PANAMA PROVINCE) wAS “ON A STICk LYINg ON BARE SOIL, IN A ROADSIDE DITCh AT NIghT. TwO LIVE SPECIMENS OBSERVED AT NIghT BY T. HERMAN AND COLLEAgUES (PERS. COMM.) AT CERRO BREwSTER (PANAM PROVINCE) wERE OBSERVED ON VEgETATION <1 M ABOVE gROUND LEVEL ( FIg. 3A, C ).

Remarks. WhILE OSTEOLOgICAL ChARACTERS OF ThE SkULL wOULD NOT SEEM TO BE USEFUL DIAgNOSTIC ChARACTERS IN ThE ABSENCE OF SkELETAL SPECIMENS OR RADIOgRAPhY, wE NOTE ThAT ThE DISTINCTLY ENLARgED SUBTEMPORAL FENESTRA OF H. panamensis CAN BE DETECTED BY MANUAL PALPATION AND IS ALSO VISIBLE EVEN IN PhOTOgRAPhS OF LIVINg SPECIMENS. DESPITE ThE RELATIVE PROXIMITY OF ThE RANgE OF ThIS SPECIES TO PANAMA CITY, AND INFRASTRUCTURAL DEVELOPMENTS SUCh AS ThE PANAMA CANAL AND VARIOUS SMIThSONIAN FIELD STATIONS, RELATIVELY FEw SPECIMENS OF ThIS SPECIES APPEAR TO EXIST IN MUSEUM COLLECTIONS. AS A RESULT, DATA REgARDINg VARIATION IN COLORATION AND MORPhOMETRY ARE UNAVAILABLE.

Figures

FIGURE 2. High-resolution Computed Tomography reconstructions of the skulls of Hemiphractus elioti new species (female: UMMZ 152622; male: AMNH 124113) and H. kaylockae new species (female: KU 93506; male: UMMZ 152624).

FIGURE 3. Photographs of live specimens of Hemiphractus from Panama. (A) Specimen of H. panamensis photographed in situ (T. Herman); sex and SVL unknown. (B) Specimen of H. panamensis displaying mouth gaping behavior while being handled (E. Baitchman); sex and SVL unknown. (C) Live specimen of H. panamensis photographed in situ (T. Herman); sex and SVL unknown. Specimens of H. panamensis all found at Cerro Brewster, Panama, Panama. (D, E) Adults of H. elioti, from Altos de María, near El Valle de Antón, Panama (B. Wilson, courtesy of El Valle Amphibian Conservation Center); probably males, SVL unknown. (F) Captive adult female of H. elioti with eggs (B. Wilson, courtesy of El Valle Amphibian Conservation Center); SVL unknown.

FIGURE 4. Dorsal, ventral, and lateral views of the heads and dorsal and ventral views of preserved specimens of Hemiphractus spp. from Panama. (A) Hemiphractus elioti (adult male holotype, SVL = 25.4 mm, head length = 12.0 mm). (B) H. kaylockae (adult female holotype, SVL = 54.2 mm, head length = 24.3 mm). (C) H. panamensis (adult female KU 116353, SVL = 40.4 mm, head length = 20.0 mm). The outline of the subtemporal fossae of the skulls typically are visible (and palpable) in preserved specimens; here they are delineated by dashed lines. Photographs by A. Thompson, courtesy of Zoo Atlanta.

FIGURE 5. A map of the known distributions of the three species of Hemiphractus recognized herein, from Panama. Type localities are white symbols, black symbols represent localities for museum vouchers and photographic records (see Fig. 3). Symbols represent H. elioti (triangles); H. panamensis (circles), and H. kaylockae (square). At this time, we know of no confirmed records of these species from neighboring Costa Rica or Colombia, and we posit that H. fasciatus Peters, 1862, does not occur in Panama, but more likely is restricted to Ecuador and perhaps part of Colombia.

FIGURE 6. Condition of the vomer and neopalatine relationships of the Hemiphractus spp. considered in this paper. (A) adult male of H. kaylockae (UMMZ 152624), showing narrow separation between odontoids on vomers and neopalatines. (B) Adult male H. elioti AMNH 124113), showing contact between odontoids on vomers and neopalatines. (C, D) adult females of H. panamensis (KU 116353) and H. fasciatus (holotype), showing wide separation between odontoids on the vomers and palatines, as well as the reduced number of vomerine odontoids compared to H. elioti and H. kaylockae. Figures are presented in similar, but not fully equivalent scales.

FIGURE 7. Preserved holotype of Cerathyla (= Hemiphractus) panamensis Stejneger, 1917. Specimen (USNM 55320) is an adult female, SVL = 59.3 mm. Photographs courtesy of USNM curatorial staff.

FIGURE 9. High-resolution Computed Tomography reconstructions of the skulls of Hemiphractus panamensis, the holotype of H. fasciatus, and an unidentified specimen of Hemiphractus from Prov. Nariño, Colombia (KU 200261). This latter specimen was initially identified and catalogued as H. fasciatus; we are unsure of its actual taxonomic identity because it resembles neither the holotype of H. fasciatus nor any of the material from Panama.