Hemiphractus kaylockae, 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 : 503-506

Hemiphractus kaylockae new species

( FIgS. 2 , 4 , 6 , 8 )

Hemiphractus panamensis Duellman 1970 [in part; for reference to specimens from eastern Prov. Darién, Panamá.

Hemiphractus fasciatus Trueb 1974 [in part; for reference to specimens from eastern Prov. Darién, Panamá]; Duellman 2001 [in part; for reference to specimens from eastern Prov. Darién, Panamá]; Crawford et al. 2012 and Castroviejo-Fisher et al. 2015 [in part, for samples from Cana, Prov. Darién, Panamá]; Köhler 2011 [in part; for map showing isolated populations in extreme eastern Panama].

Holotype. KU 107409, ADULT FEMALE, OBTAINED AT PANAMá : PROV. DARIéN: N RIDgE OF CERRO CITURO, SIERRA DE PIRRE, N 0 7 o 56’ , W 77o 42’ (APPROX.), 900–1000 M ON 17 JANUARY 1966, BY ChARLES W. MYERS.

Paratypes, MALES: KU 107411, 107414 PANAMA: DARIEN: N RIDE CERRO CITURO, SERRANIA DE PIRRE, 1000 M; KU 107419 PANAMA: DARIEN: N RIDE CERRO CITURO, SERRANIA DE PIRRE, 900 M; KU 154992, PANAMA: DARIEN: S SLOPE CERRO CITURO, SERRANIA DE PIRRE, 1100 M. FEMALES: AMNH A92668, PANAMA: DARIEN: N RIDgE CERRO CITURO, SERRANIA DE PIRRE, 1100 M; KU 93506, PANAMA: DARIEN: SE SLOPE CERRO PIRRE, 1440 M; KU 107413, PANAMA: DARIEN: N RIDgE CERRO CITURO, SERRANIA DE PIRRE, 1000 M; KU 93507, PANAMA: DARIEN, SE SLOPE CERRO PIRRE, 1440 M.

Diagnosis. Hemiphractus kaylockae MAY BE DISTINgUIShED FROM ALL SPECIES OF Hemiphractus EXCEPT H. elioti , H. fasciatus , H. panamensis AND H. scutatus BY NOT hAVINg A POSTORBITAL INDENTATION ( FIg. 2 ). Hemiphractus kaylockae 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 kaylockae DIFFERS FROM H. elioti BY hAVINg NEOPALATINE AND VOMERINE ODONTOIDS NARROwLY SEPARATED (IN CONTACT IN H. elioti ; CONDITION UNkNOwN IN H. scutatus ), whEREAS ThESE ROwS OF ODONTOIDS ARE wIDELY SEPARATED IN H. panamensis AND H. fasciatus ( FIg. 6 ). Hemiphractus kaylockae FURThER DIFFERS FROM H. panamensis AND H. fasciatus BY hAVINg APPROXIMATELY SEVEN VOMERINE ODONTOIDS (VS. APPROXIMATELY TwO IN BOTh H. panamensis AND H. fasciatus ). Hemiphractus kaylockae MAY BE DISTINgUIShED FROM H. elioti , H. fasciatus , AND H. panamensis BY hAVINg A DISTINCT INDENTATION ON ThE paraoccipital hORMS, FORMINg AN ANgULAR MARgIN OF ThE PARAOCCIPITAL hORNS ( FIgS. 2 , 9 ); SUPRAORBITAL RIDgES ABSENT (EVIDENT, DISTINCT IN H. fasciatus AND wEAkLY DEVELOPED IN H. panamensis ; FIgS. 2 , 9 ); CANThAL RIDgES EVIDENT, DISTINCT (ABSENT OR wEAkLY DEVELOPED IN H. elioti ; FIg. 2 ). Hemiphractus kaylockae hAS A SUBTEMPORAL FENESTRA APPROXIMATELY ONE-ThIRD DIAMETER OF ThE ORBIT IN BOTh MALES AND FEMALES, APPROXIMATELY ONE-hALF DIAMETER OF ORBIT IN FEMALES IN H. elioti (VERY SMALL IN MALES; FIg. 2 ), APPROXIMATELY ONEhALF DIAMETER OF ThE ORBIT IN FEMALES OF H. fasciatus ( FIg. 9 ; MALES UNkNOwN), VERY LARgE, APPROXIMATELY 1.5 X SIzE OF ORBIT, IN FEMALES OF H. panamensis ( FIg. 9 ; MALES UNkNOwN); LATERAL MARgINS OF ThE qUADRATOjUgALS EXTEND TO LEVEL OF LATERAL TIPS OF, OR SLIghTLY LESS ThAN, BEYOND ThE PARAOCCIPITAL hORNS IN DORSAL VIEw (EXTENDINg FAR BEYOND ThE PROFILE OF ThE PARAOCCIPITAL hORNS IN H. panamensis ; FIgS. 2 , 9 ).

Description of the holotype. AN ADULT FEMALE: SVL 54.2 MM (NOT INCLUDINg PROBOSCIS); TIBIA LENgTh 28.4 MM (NOT INCLUDINg CALCAR); FOOT LENgTh 25.3 MM; hEAD LENgTh 24.3 MM (NOT INCLUDINg PROBOSCIS); hEAD wIDTh 26.5 MM; SNOUT-PARAOCCIPITAL PROCESS LENgTh 25.1 MM; TYMPANUM OVOID, 6.6 MM hIgh X 3.6 MM wIDE; EYE DIAMETER 6.5 MM; INTERORBITAL DISTANCE 10.2 MM; EYE – TYMPANUM DISTANCE 5.4 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 RIDgE PRESENT, wELL-DEVELOPED; PARAOCCIPITAL RIDgES PRESENT, wEAkLY DEVELOPED; SUPRAORBITAL, zYgOMATIC, PREORBITAL, AND CANThAL RIDgES ABSENT; CANThUS ROSTRALIS INDISTINCT; LOREAL REgION FLAT; PROBOSCIS ShORT, FLEShY, POINTED, LATERALLY FLATTENED IN CROSS SECTION; SNOUT ANgLINg POSTERIORLY AT LESS ThAN 45o FROM hORIzONTAL; NOSTRILS OVOID, SLIghTLY PROTUBERANT, PROjECTINg DORSALLY; LIPS NOT FLARED. TOP OF hEAD RELATIVELY FLAT; INTERORBITAL REgION 38 % OF hEAD wIDTh; EYE DIAMETER 25 % hEAD wIDTh; TYMPANUM VERTICALLY OVOID, ORIENTED POSTERIORLY AND SLIghTLY LATERALLY, VISIBLE BELOw PARAOCCIPITAL PROCESSES IN DORSAL VIEw, gREATEST DIAMETER EqUIVALENT TO EYE DIAMETER; wIDTh OF TYMPANUM 1.6 TIMES EYE–TYMPANUM DISTANCE; UPPER EYELID wITh ONE ENLARgED POINTED PROjECTION AND SCATTERED SMALLER TUBERCLES; INTERORBITAL TUBERCLE ABSENT; POSTORBITAL TUBERCLE 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 ABOUT 1.8 TIMES wIDTh OF DIgITS AT PENULTIMATE PhALANgE; RELATIVE LENgThS OF FINgERS: III> IV> II> I; LATERAL FRINgE ON FINgERS ABSENT; OUTER METACARPAL TUBERCLE LOw, OVOID; INNER METACARPAL TUBERCLE OVOID, DIFFUSE; FOREARM wITh SCATTERED TUBERCLES, LARgE, POINTED, VARIOUSLY FUSED TO PRODUCE AN EVIDENT LATERAL FRINgE; wEBBINg ON hANDS ABSENT. HEELS OF ADPRESSED hIND LIMBS OVERLAP; TIBIOTARSAL ARTICULATION EXTENDINg TO SNOUT whEN hIND LIMB EXTENDED; TARSAL FOLD PRESENT, wEAkLY DEVELOPED; TIBIA LENgTh 52 % SVL; FOOT LENgTh 47 % SVL; OUTER METATARSAL TUBERCLE ABSENT; INNER METATARSAL TUBERCLE PRESENT, LARgE, OVOID; TOES IV AND V wITh LATERAL FRINgE, PROMINENT ON TOE V; FRINgE OF TOE V EXTENDINg ONTO OUTER SURFACE OF BASE OF TARSUS; DORSAL SURFACE OF hIND LIMB RELATIVELY SMOOTh, BEARINg IRREgULAR SETS OF TUBERCLES ALONg ThREE TRANSVERSE BANDS ON ShANk; CALCAR PRESENT, SMALL; RELATIVE LENgThS OF TOES: IV> V> III> II> I; TOE PADS ABOUT 1.5 TIMES wIDTh OF PENULTIMATE PhALANgE; wEBBINg BASAL. CLOACAL OPENINg DIRECTED POSTEROVENTRALLY AT MIDLEVEL OF ThIgh. DORSUM SMOOTh wITh SCATTERED, LOw, DIFFUSE TUBERCLES; FLANkS SMOOTh; VENTRAL SURFACES AND ThROAT SMOOTh wITh SCATTERED LOw, DIFFUSE TUBERCLES; SACRAL DIAPOPhYSES PROjECTINg LATERALLY, SLIghTLY NARROwER ThAN POSTERIORMOST TRANSVERSE PROCESSES OF ALL VERTEBRAE; LATERAL TERMINI OF TRANSVERSE PROCESSES OF PRESACRAL VERTEBRAE VI–VIII NOT CO-OSSIFIED wITh SkIN; FIVE NEURAL SPINES PROjECTINg DORSALLY, CO-OSSIFIED wITh SkIN, VISIBLE EXTERNALLY. PUPIL hORIzONTAL.

ALL DORSAL SURFACES UNIFORMLY BROwN ( FIg. 4 ). ThIgh BEARINg TwO, ShANk BEARINg ThREE INDISTINCT TRANSVERSE BANDS OF IRREgULAR, BROwN MOTTLINg ON DORSAL SURFACES, CORRESPONDINg wITh ThREE BANDS OF TUBERCLES ON ThESE BANDS; TOES I AND II ARE UNIFORMLY BEIgE, TOES III–V UNIFORMLY BROwN; FOREARM DARk BROwN, BEARINg BROAD, BROwN, TRANSVERSE BAND ON DORSAL SURFACE; DORSAL SURFACE OF MANUS MORE PALE ThAN FOREARM, ULTIMATE AND PENULTIMATE PhALANgES OF FINgERS I AND II BEIgE, FINgERS III AND 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 DARk BROwN, EXTENDINg ONTO PECTORAL REgION. VENTRAL SURFACES OF FOREARMS BEIgE wITh FINE, BROwN MOTTLINg; VENTRAL SURFACES OF hANDS BEIgE. VENTRAL SURFACES OF hIND LIMB AND FOOT BEIgE wITh FINE, PALE BROwN MOTTLINg.

Variation. CONSISTENT SEXUAL DIMORPhISM IN COLORATION OR SkIN TEXTURE wAS NOT EVIDENT ( FIgS. 4 , 8 ). GRANULATION ON ThE ThROAT IN Hemiphractus kaylockae IS VARIABLE AMONg SPECIMENS EXAMINED, VARYINg FROM SMOOTh (E.g., KU 93506, 93507, 107411, 107417, 154992) TO SMOOTh wITh TUBERCLES (E.g., AMNH 92668, KU 107419), SMOOThLY gRANULAR wITh NO TUBERCLES (E.g., KU 107412, 107413), TO SMOOThLY gRANULAR wITh TUBERCLES PRESENT (E.g., KU 107408, 107414, 107422, 107423, 154991, UMMZ 152624). GRANULATION OF ThE VENTER RANgES FROM SMOOTh TO UNIFORMLY gRANULAR (E.g., KU, 107411, 107422, 154992, UMMZ 152624); SOME SPECIMENS wITh SMOOTh OR UNIFORMLY gRANULAR VENTERS ALSO hAD VARIABLE NUMBERS OF DISCRETE TUBERCLES PRESENT (E.g., AMNH 92668, KU 93506, 93507, 107412, 107413, 107414, 107417, 107419, 107423, 154991). ThROAT COLORATION IN MOST SPECIMENS IS TAN (E.g. KU 107417) TO BROwN OR DARk BROwN (E.g., AMNH 92668, KU 93506, 107408, 107411, 107412, 107419, 107422, 154992) wITh SOME ShOwINg PROMINENT whITE MOTTLINg, STRIPINg, AND/OR PROMINENT LABIAL BARRINg (E.g. KU 93506, 107412, 107422, 154992) AND KU 93507, 107413, 107414, 107417, 107423, 154991, UMMZ 152624 DISPLAYINg STIPPLINg OR SPECkLINg OF BROwN AND whITE. 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 BEIgE COLORATION ThAT TYPICALLY CONTAINS VARIABLE AMOUNTS OF DARkER BROwN STIPPLINg (E.g., AMNH 92668, KU 93507, 107408, 107411, 107414, 107417, 107419, 107423, 154992); KU 93506 AND KU 107413 ShOw A SIMILAR PATTERN hOwEVER ALSO DISPLAY whITE MOTTLINg IN ThE PECTORAL REgION whILE KU 107422 DISPLAYS whITE SPOTTINg POSTERIORLY. ThE DORSUM OF H. kaylockae IS SMOOTh OR UNIFORMLY SMOOThLY gRANULAR IN ALL SPECIMENS wITh KU 93506, 107423, 154991, 154992, AND UMMZ 152624 hAVINg SOME TUBERCLES. DORSAL COLORATION IN MOST IS AN OVERALL TAN OR BEIgE ThAT MAY OR MAY NOT INCLUDE SOME LEVEL OF DARkER BROwN STIPPLINg OR MOTTLINg (E.g. KU 107408, 107412, 107413, 107417, 107419, 107422, 107423, 154991, 154992) whILE SOME SPECIMENS DISPLAY AN OVERALL UNIFORM BROwN COLOR (E.g. AMNH 92668, KU 93506, 93507, 107411, 107414). SOME SPECIMENS ShOw A PROMINENT DARk BAND ON ThE FOREARM (E.g. KU 107408, 107412, 107422, 107423, 154992). HOwEVER, ThIS BAND IS DIFFUSE OR wEAkLY DEVELOPED IN ALL OThER SPECIMENS. TwO OR ThREE DARk TRANSVERSE BANDS ARE PRESENT ON ThE FEMORAL AND TIBIAL REgIONS (ABSENT OR BARELY DISCERNIBLE ON AMNH 92668, KU 93506, 107411, 107419, 154991), AND A DIFFUSE OR VERY EVIDENT DARk BAND OR ChEVRON PRESENT ON ThE SACRAL REgION (E.g., KU 107408 [wEAkLY DEVELOPED], 107412, 107414, 107417, 107422, 154992, UMMZ 152624); IN SPECIMENS KU 107422 AND 154992 ThERE ARE TwO ChEVRONS PRESENT; IN ADDITION, SPECIMENS KU 107414, 107417, 107422, AND 154992 hAVE A DARk IRREgULAR VERTEBRAL STRIPE EXTENDINg FROM ThE SACRAL BAND CONNECTINg TO A DARk BLOTCh AT ThE OCCIPUT. ThE POSTERIOR SURFACE OF ThE ThIgh OF KU 107412–14 AND UMMZ 152624 BEARS A PATTERN OF TAN OR BROwN MOTTLED wITh whITE OR BEIgE whILE ALL OThER SPECIMENS BEAR A DIFFUSE (E.g., KU 93506, 93507, 107411, 107422, 107423, 154991, 154992) OR DISTINCT whITE STRIPE ALONg ThIS SURFACE (E.g., AMNH 92668, KU 107408, 107417, 107419).

Etymology. ThIS SPECIES IS NAMED IN hONOR AND MEMORY OF JULIA KAYLOCk. ShE wAS A COLLEAgUE AND FRIEND, BELOVED BY MANY, whO wAS PASSIONATE ABOUT ThE CONSERVATION OF ThE wORLD’S BIODIVERSITY, ESPECIALLY ThAT OF AMPhIBIANS. ShE SPENT ThE LATTER PART OF hER LIFE wORkINg TIRELESSLY TO IMPROVE AND DOCUMENT hUSBANDRY TEChNIqUES FOR ThREATENED PANAMANIAN AMPhIBIANS (E.g., HILL et al. 2012) MAINTAINED IN ex situ FACILITIES. ShE LOST A LIFELONg BATTLE wITh TYPE I DIABETES AT ThE AgE OF 28 ON 12 JUNE 2009. WE gRACIOUSLY ACkNOwLEDgE hER PASSION FOR AMPhIBIAN CONSERVATION IN ThE NAMINg OF ThIS SPECIES OF Hemiphractus . ThE SPECIFIC EPIThET IS ThE SINgULAR gENITIVE CASE OF ThE NAME KAYLOCk.

Distribution and ecology. SPECIMENS OF Hemiphractus kaylockae hAVE BEEN COLLECTED ONLY IN ONE REgION OF FAR EASTERN PANAMA, IN ThE SERRANíA DE PIRRE, PROV. DARIEN ( FIg. 5 ). ThIS REgION REMAINS POORLY DOCUMENTED BIOLOgICALLY, AND ThIS SPECIES IS EXPECTED TO OCCUR SOMEwhAT MORE BROADLY—AT LEAST UNTIL ThE RECENT INVASION OF ThE AMPhIBIAN PAThOgEN Batrachochytrium dendrobatidis. ThIS FROg hAS BEEN FOUND ONLY IN MOIST, LOwER MONTANE FORESTS AT ELEVATIONS FROM 900 TO 1550 M ( MYERS 1966). SPECIMENS COLLECTED AT NIghT AT CERRO PIRRE (PROV. DARIEN) BY MYERS (1966: 70) wERE OBSERVED “…ON LOw PERChES (LOg, STICkS, DEAD PALM FRONDS) ONLY ONE-hALF TO 2 FEET ABOVE gROUND, AND TwO wERE ON PALM FRONDS 3 TO 4 FEET ABOVE ThE FOREST FLOOR. MYERS (1966) ALSO PROVIDED IN ThIS ACCOUNT A DESCRIPTION OF A MOUTh gAPINg BEhAVIOR IN whICh ThE YELLOwISh ORANgE TONgUE IS DISPLAYED FOLLOwED BY A PAINFUL BITE wITh CONTINUED AgITATION AS wELL AS A “LOUD CATLIkE SqUALL EMITTED whEN REMOVINg AN INDIVIDUAL FROM A BAg ( FIg. 8C ). IT IS LIkELY ThAT FEMALES OF ThIS SPECIES CARRY EggS ON ThE BACk AS hAS BEEN DOCUMENTED FOR ALL OThER DESCRIBED SPECIES OF Hemiphractus . AS ALL DOCUMENTED OBSERVATIONS hAVE OCCURRED AT NIghT, IT IS INFERRED ThAT ThIS SPECIES IS MOST LIkELY NOCTURNAL.

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 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.