Hemiphractus elioti, Hill & Martin & Stanley & Mendelson Iii, 2018

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 : 499-503

Hemiphractus elioti new species

( FIgS. 2–4 , 6 )

Hemiphractus panamensis Duellman 1970 [in part; for reference to specimens from western Panama.

Hemiphractus fasciatus Trueb 1974 [in part; for reference to specimens from western Panama]; Duellman 2001 [in part; for reference to specimens from western Panama]; Crawford et al. 2012 [in part, for samples from Río Blanco and El Copé, Prov. Coclé, and Altos de Maria, Prov. Panamá, Panamá]; Castroviejo-Fisher et al. 2015 [in part; for sample from El Copé Prov. Coclé, Panamá]; Köhler 2011 [fig. 467; in part, for map showing isolated populations in western Panama].

Holotype. UTACV A-64827 , A SMALL, ADULT MALE, OBTAINED AT PANAMá : PROV. PANAMá: RíO MARIA, ALTOS DE MARIA, N 0 8 o 38.585’, W 0 80o 04.357’, 875 M, ON 20 MAY 2005, BY EDgARDO GRIFFITh AND HEIDI ROSS .

Paratypes. FEMALES: UMMZ 152622, 152623, PANAMA: COCLé: 6 MI [9.7 kM] NNW EL COPé, 600–800 M; AMNH A124114, PANAMA: ChIRIqUI: CONTINENTAL DIVIDE ABOVE QUEBRADA DE ARENA, 1200–1300 M. MALES: AMNH A124113, A124115, PANAMA: ChIRIqUI: CONTINENTAL DIVIDE ABOVE QUEBRADA DE ARENA, 1200–1300 M; UMMZ 153118, PANAMA COCLé: 6 MI [9.7 kM] NNW EL COPé, ON CONTINENTAL DIVIDE AT SAwMILL, 600–800 M, USNM 572599, PANAMA: COCLé: N OF EL COPé.

Diagnosis. Hemiphractus elioti MAY BE DISTINgUIShED FROM ALL SPECIES OF Hemiphractus EXCEPT H. fasciatus , H. kaylockae , H. panamensis , AND H. scutatus BY NOT hAVINg A POSTORBITAL INDENTATION. Hemiphractus elioti MAY BE DISTINgUIShED FROM H. scutatus , AN AMAzONIAN SPECIES NOT OCCURRINg IN PANAMA, BY ITS SMALLER SIzE (MAXIMUM SVL IN MALES 52.5 MM VS. 57.4 MM IN H. scutatus ; FEMALES 64.7 MM VS. 80.5 MM IN H. scutatus ; DATA FOR H. scutatus FROM TRUEB 1974), AND BY hAVINg SMALL EXPANDED PADS ON ThE FINgERS AND TOES (ABSENT IN H. scutatus ). Hemiphractus elioti DIFFERS FROM H. fasciatus , H. kaylockae , AND H. panamensis BY hAVINg NEOPALATINE ODONTOIDS IN CONTACT wITh VOMERINE TEETh AT ThE CENTER OF ThE PALATAL REgION (NARROwLY SEPARATED IN H. kaylockae VS. wIDELY SEPARATED IN BOTh H. panamensis AND H. fasciatus ; CONDITION UNkNOwN IN H. scutatus ; FIg. 6 ). Hemiphractus elioti FURThER DIFFERS FROM H. panamensis AND H. fasciatus BY hAVINg APPROXIMATELY SEVEN VOMERINE ODONTOIDS (VS. APPROXIMATELY TwO; CONDITION UNkNOwN IN H. scutatus ; FIg. 6 ). Hemiphractus elioti MAY BE DISTINgUIShED FROM H. kaylockae BY hAVINg STRAIghT MARgIN OF ThE LATERAL MARgINS OF ThE PARAOCCIPITAL hORNS (wITh DISTINCT INDENTATION, FORMINg AN ANgULAR MARgIN IN H. kaylockae ; FIg. 2 ); SUPRAORBITAL RIDgES ABSENT (EVIDENT, DISTINCT IN H. fasciatus , wEAkLY DEVELOPED IN H. panamensis ; FIgS. 2 , 9 ); CANThAL RIDgES USUALLY ABSENT OR INDISTINCT (EVIDENT, DISTINCT IN H. fasciatus , H. kaylockae , AND SLIghTLY DEVELOPED IN H. panamensis ; FIgS. 2 , 9 ). Hemiphractus elioti hAS A SUBTEMPORAL FENESTRA VERY SMALL IN MALES, APPROXIMATELY ONE-hALF OF ORBIT IN FEMALES, whEREAS ThE FENESTRAE ARE APPROXIMATELY ONE-ThIRD DIAMETER OF ThE ORBIT IN BOTh MALES AND FEMALES OF H. kaylockae ( FIg. 2 ), APPROXIMATELY ½ DIAMETER OF ThE ORBIT OF FEMALES IN H. fasciatus ( FIg. 9 ; MALES UNkNOwN), VERY LARgE, APPROXIMATELY 1.5 TIMES SIzE OF ORBIT, IN FEMALES OF H. panamensis ( FIg. 9 ; MALES UNkNOwN); LATERAL MARgINS OF ThE qUADRATOjUgALS EXTEND TO LEVEL OF ThE LATERAL TIPS OF, OR VERY SLIghTLY BEYOND, ThE PARAOCCIPITAL hORNS IN DORSAL VIEw (EXTENDINg FAR BEYOND ThE LATERAL TIPS OF ThE PARAOCCIPITAL hORNS IN H. panamensis ; FIgS. 2 , 9 ).

Description of the holotype. SMALL, ADULT MALE, SNOUT–VENT LENgTh (SVL) 40.5 MM (NOT INCLUDINg PROBOSCIS); TIBIA LENgTh 22.3 MM (NOT INCLUDINg CALCAR); FOOT LENgTh 19.6 MM; hEAD LENgTh 20.3 MM (NOT INCLUDINg PROBOSCIS); hEAD wIDTh 20.2 MM; SNOUT–PARAOCCIPITAL PROCESS LENgTh 20.2 MM; TYMPANUM OVOID, 5.4 MM hIgh X 3.8 MM wIDE; EYE DIAMETER 5.5 MM; INTERORBITAL DISTANCE 6.9 MM; EYE–TYMPANUM DISTANCE 3.1 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 AND PARAOCCIPITAL RIDgES PRESENT, wEAkLY DEVELOPED; SUPRAORBITAL, zYgOMATIC RIDgES ABSENT; PREORBITAL, CANThAL RIDgES BARELY EVIDENT, LESS PROMINENT ThAN OCCIPITAL AND PARAOCCIPITAL RIDgES; CANThUS ROSTRALIS INDISTINCT; LOREAL REgION FLAT; PROBOSCIS ShORT, FLEShY, POINTED, TRIANgULAR IN CROSS SECTION; SNOUT ANgLINg POSTERIORLY AT LESS ThAN 45o FROM hORIzONTAL; NOSTRILS OVOID, SLIghTLY PROTUBERANT, PROjECTINg DORSALLY, SLIghTLY LATERALLY; LIPS NOT FLARED. TOP OF hEAD RELATIVELY FLAT; INTERORBITAL REgION 34 % hEAD wIDTh; EYE DIAMETER 27 % hEAD wIDTh; TYMPANUM VERTICALLY OVOID, ORIENTED POSTERIORLY AND SLIghTLY LATERALLY, VISIBLE BELOw PARAOCCIPITAL PROCESSES IN DORSAL VIEw, gREATEST DIAMETER 1.5 TIMES EYE DIAMETER; wIDTh OF TYMPANUM 1.2 TIMES EYE–TYMPANUM DISTANCE; UPPER EYELID BEARINg SINgLE ENLARgED POINTED PROjECTION AND SCATTERED SMALLER TUBERCLES; INTERORBITAL, POSTORBITAL TUBERCLES ABSENT; SkIN OVER POSTERIOR SURFACE OF PARS ARTICULARIS BEARINg ONE SMALL, FLEShY TUBERCLE. TONgUE LARgE, OVOID; VOCAL SLITS ABSENT; 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 ENLARgED DISTAL PADS; FINgER PADS 2.6 TIMES DIAMETER OUTER METACARPAL TUBERCLE LOw, TRIPARTITE, DIFFUSE; INNER METACARPAL TUBERCLE OVOID, RAISED; RELATIVE LENgThS OF FINgERS: III> IV> II> I; LATERAL FLANgES ABSENT ON FINgERS; FOREARM wITh SCATTERED TUBERCLES, LOw, ROUND, BEARINg wEAkLY DEVELOPED LATERAL FRINgE; wEBBINg ON hANDS ABSENT. HEELS OF ADPRESSED hIND LIMBS OVERLAP; TIBIOTARSAL ARTICULATION EXTENDINg TO SNOUT whEN hINDLIMB EXTENDED; TARSAL FOLD PRESENT, wEAkLY DEVELOPED; TIBIA LENgTh 55 % SVL; FOOT LENgTh 48 % SVL; OUTER METATARSAL TUBERCLE wEAkLY DEVELOPED, DIFFUSE; INNER METATARSAL TUBERCLE PRESENT, RAISED, OVOID; TOES II–IV wITh PROMINENT, LATERAL FRINgE; FRINgE OF TOE V EXTENDINg ONTO OUTER SURFACE OF BASE OF TARSUS; DORSAL SURFACE OF hIND LIMB RELATIVELY SMOOTh, BEARINg IRREgULAR SETS OF TUBERCLES CONCENTRATED ON ThREE TRANSVERSE BANDS OF DARk-BROwN COLORATION ON ShANk; CALCAR PRESENT, SMALL; RELATIVE LENgThS OF TOES: IV> V> III> II> I; TOE PADS 2 X wIDTh OF PENULTIMATE PhALANgES OF DIgITS; wEBBINg BASAL. CLOACAL OPENINg DIRECTED POSTEROVENTRALLY AT MIDLEVEL OF ThIgh. DORSUM AND FLANkS SMOOTh; VENTRAL SURFACES AND ThROAT AND ChIN IRREgULARLY gRANULAR; SACRAL DIAPOPhYSES PROjECTINg LATERALLY BEYOND TRANSVERSE PROCESSES OF ALL VERTEBRAE; LATERAL TERMINI OF TRANSVERSE PROCESSES OF PRESACRAL VERTEBRAE VI–VIII NOT CO-OSSIFIED wITh SkIN (NEARLY TO LEVEL OF SACRAL DIAPOPhYSES) AND VISIBLE EXTERNALLY; SIX NEURAL SPINES PROjECTINg DORSALLY, CO-OSSIFIED wITh SkIN, VISIBLE EXTERNALLY. PUPIL hORIzONTAL.

IN ALCOhOL, DORSUM BEIgE wITh MINUTE BLACk PUNCTATIONS ( FIg. 4 ). ThIgh AND ShANk BEARINg ThREE INDISTINCT TRANSVERSE BANDS OF IRREgULAR, BROwN MOTTLINg ON DORSAL SURFACES, CORRESPONDINg wITh ThREE BANDS OF TUBERCLES ON ShANk ONLY; ULTIMATE AND PENULTIMATE PhALANgES OF TOES IV–V DARk BROwN; FOREARM BEARINg BROAD, BROwN, TRANSVERSE BAND ON DORSAL SURFACE; DORSAL SURFACE OF MANUS MORE PALE ThAN FOREARM, ULTIMATE AND PENULTIMATE PhALANgES OF FINgERS V–IV DARk BROwN. UPPER AND LOwER LIPS wITh NARROw, TRANSVERSE, DARk BROwN BANDS, NOT EXTENDINg ONTO PROXIMAL SURFACES. FLANkS BEIgE wITh DIFFUSE BROwN PUNCTATIONS; VENTER BEIgE wITh SCATTERED DIFFUSE, FINE BROwN MARkINgS; ThROAT BEIgE wITh FINE BROwN MOTTLINg. VENTRAL SURFACE OF FOREARM BEIgE wITh FINE, BROwN MOTTLINg; VENTRAL SURFACE OF hAND BEIgE. VENTRAL SURFACES OF hIND LIMB AND FOOT BEIgE wITh FINE, PALE BROwN MOTTLINg; TOES I AND II ARE UNIFORMLY BEIgE, TOES III–V UNIFORMLY BROwN.

Variation. CONSISTENT SEXUAL DIMORPhISM IN COLORATION OR SkIN TEXTURE wAS NOT EVIDENT. GRANULATION OF ThE ThROAT IN Hemiphractus elioti IS VARIABLE AMONg SPECIMENS EXAMINED, VARYINg FROM SMOOThLY gRANULAR wITh NO TUBERCULATION (E.g., USNM 153118, 297543, 572600, 572601) TO SMOOThLY gRANULAR wITh DISCRETE TUBERCLES PRESENT (E.g., AMNH 107955, 124113, 124115, UMMZ 167358, 297542, 572599). TEXTURE OF ThE VENTER RANgES FROM SMOOTh (E.g., UMMZ 152622,152623) TO UNIFORMLY gRANULAR (E.g., UMMZ 297543); SOME SPECIMENS (E.g., AMNH 107955, 124113, 124114, UMMZ 153118, USNM 297541, 297542, 572599) wITh gRANULAR VENTERS ALSO hAD VARIABLE NUMBERS OF DISCRETE TUBERCLES PRESENT. ThROAT COLORATION IS hIghLY VARIABLE, INCLUDINg SPECIMENS wITh UNIFORM TAN (E.g., AMNH 107955, USNM 297542, 297543) TO UNIFORM DARk BROwN (E.g., AMNH 124113–15, UMMZ 152622, 152623, USNM 572599); EIThER PATTERN MAY INCLUDE A FAINT TO PROMINENT PALE-COLORED STRIPE EXTENDINg FROM ThE SYMPhYSIS OF ThE LOwER jAw ONTO ThE PECTORAL REgION, A SPECkLED PATTERN OF whITE AND BROwN (E.g., USNM 572600), OR A gENERALLY whITE gROUND COLORATION, wITh DISTINCT MARgINAL gRAY MARBLINg (E.g., USNM 572601). VENTRAL COLORATION IN MOST SPECIMENS TYPICALLY IS A CONTINUATION OF ThE gULAR COLORATION OF DARk BROwN OR TAN EXTENDINg TO APPROXIMATELY MID-ABDOMEN whERE IT ABRUPTLY ChANgES TO A TAN COLORATION (E.g., AMNH 107955, 124113–15, UMMZ 153118, 152622, 152623, USNM 572599). VENTRAL COLORATION OF SOME SPECIMENS (E.g., USNM 297541–43) IS UNIFORMLY PALE TAN, OR whITE (E.g., UMMZ 167358, 572600, USNM 572601); ONE SPECIMEN (USNM 572600) IS UNIqUE IN hAVINg A SPECkLED VENTRAL PATTERN OF BROwN AND whITE. ThE DORSUM IS UNIFORMLY SMOOThLY gRANULAR IN ALL SPECIMENS AND OVERALL A UNIFORM ShADE OF BROwN. MOST SPECIMENS ShOw A SINgLE DARk BAND ON ThE FOREARM (ABSENT OR BARELY DISCERNIBLE IN AMNH 107955, UMMZ 152623, USNM 297542, 297543, 572599, 572600); TwO OR ThREE DARk TRANSVERSE BANDS ON ThE FEMORAL AND TIBIAL REgIONS (ABSENT OR BARELY DISCERNIBLE ON AMNH 10795, UMMZ 152623, 297541–43, 572599–601), AND A DARk BAND OR ChEVRON PRESENT ON ThE SACRAL REgION (E.g., AMNH 124113–15, UMMZ 152623); IN ADDITION, SPECIMEN AMNH 124113 hAS A DARk IRREgULAR VERTEBRAL STRIPE EXTENDINg FROM ThE SACRAL BAND TO ThE OCCIPUT. ThE POSTERIOR SURFACE OF ThE ThIgh OF USNM 572599 BEARS A PATTERN OF TAN SPOTS OVER A BROwN BACkgROUND COLORATION, AND UMMZ 152623 hAS A DISTINCT whITE STRIPE ALONg ThIS SURFACE.

Etymology. EDgARDO GRIFFITh AND HEIDI ROSS hAVE gREATLY CONTRIBUTED TO ThE kNOwLEDgE BASE OF ThE NATURAL hISTORY, REPRODUCTIVE BIOLOgY, AND CONSERVATION OF PANAMANIAN AMPhIBIANS, INCLUDINg Hemiphractus . WE ARE PROUD TO ACkNOwLEDgE ThE EFFORTS OF OUR FRIENDS AND COLLEAgUES BY NAMINg ThIS NEw SPECIES OF Hemiphractus IN hONOR OF ThEIR SON, ELIOT. ThE SPECIFIC EPIThET IS ThE SINgULAR gENITIVE CASE OF ThE NAME ELIOT.

Distribution and ecology. SPECIMENS OF Hemiphractus elioti hAVE BEEN COLLECTED FROM VARIOUS LOCALITIES IN ThE CORDILLERA DE TALAMANCA, INCLUDINg SITES IN ThE PROVINCES OF BOCAS DEL TORO, ChIRIqUI, COCLé, AND PANAMá IN wESTERN PANAMA ( FIg. 5 ). HABITAT IS PRE-MONTANE MOIST PRIMARY FOREST RANgINg IN ELEVATION FROM 600–1600 M. LIVINg SPECIMENS hAVE BEEN OBSERVED FROM gROUND LEVEL TO <1 M IN VEgETATION AT NIghT. COLORATION AND MORPhOLOgICAL FEATURES PROVIDE EXCELLENT CAMOUFLAgE AMONgST LEAF LITTER ON ThE FOREST FLOOR. CAPTIVE SPECIMENS AT ThE EL VALLE AMPhIBIAN CONSERVATION CENTER FED READILY ON OThER ANURANS, gECkOS ( Hemidactylus SPP.), NEwBORN MICE ( Mus musculus ), AND EARThwORMS (H. ROSS AND E. GRIFFITh PERS. COMM., RLH PERS. OBS.). WhILE DISSECTINg SPECIMENS TO DETERMINE SEX, UNIDENTIFIED ANURANS wERE EVIDENT IN ThE STOMAChS OF SPECIMENS. NUMEROUS UNIDENTIFIED NEMATODE wORMS wERE ALSO OBSERVED IN ThE COELOM AND IMBEDDED IN ThE PERITONEUM AND TISSUES OF OThER ABDOMINAL ORgANS. MYERS (1966) DESCRIBED A DISTINCTIVE MOUTh gAPINg BEhAVIOR, YELLOw ORAL COLORATION, AND DEFENSIVE BITINg IN SPECIMENS REFERABLE TO H. panamensis ( FIg. 3B ) AND H. kaylockae ( FIg. 8C ). SIMILAR BEhAVIORS hAVE BEEN OBSERVED IN H. elioti . REPEATED OBSERVATIONS OF CAPTIVE REPRODUCTION OF ThIS SPECIES AT ThE EL VALLE AMPhIBIAN CONSERVATION CENTER hAVE ShOwN ThAT FEMALES CARRY EggS AND YOUNg ON ThEIR BACkS, AS IS kNOwN TO OCCUR IN ALL DESCRIBED SPECIES OF Hemiphractus .

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 8. Live specimens of Hemiphractus kaylockae. (A, B) Adult specimens from vicinity of Cana, Darién, Panama (B. Wilson, courtesy of El Valle Amphibian Conservation Center): probable males, SVL unknown. (C) Photograph (KUDA 003746) of specimen KU 93506 (C.W. Myers) from Cerro Pirre, Darién, Panama; adult female, SVL = 54.2 mm. (D) Photograph (KUDU 003744) of specimen KU 93505 (C.W. Myers) from Cerro Pirre, Darién, Panama; adult female, SVL = 40. 5 mm. (E) Adult specimen from vicinity of Cana, Darién, Panama (B. Wilson, courtesy of El Valle Amphibian Conservation Center): probably female, SVL unknown.

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.