1993 mccranie & wilson b dunni group

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A Review of the Bolitoglossa dunni Group (Amphibia: Caudata) from Honduras

with the Description of Three New Species

James R. McCranie; Larry David Wilson

Herpetologica, Vol. 49, No. 1. (Mar., 1993), pp. 1-15.

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HERPETOLOGICA VOL. 49 MARCH 1993 K O . 1Herpetologtca, 49(1) , 1993, 1-1562 1993 b\ The Herpetolog~sts'League, In c

A REVIEW OF THE BOLITOGLOSSA DUNNI GROUP(AMPHIBIA: CAUDATA) FROM HONDURAS WITH

THE DESCRIPTION OF THREE NEW SPECIESJAMES R. ~ ~ X ~ A N I E ' DAVIDWILSON~ND LARRY

'10770 SW 164th Street, Miami, FL 33157, U S A 2Department of Biology, South Campus, Miami-Dade Communi ty College, Miami, F L 33176, U S A

ABSTKACT: The type series of Bolitoglossa dunni Schmidt contains representatives of two distinctspecies of salamanders. Bolitoglossa dunni is redescribed and a description of the species heretoforesubsumed in dunni is provided. In addition, two new species of this group from cloud forestlocalities in southwestern Honduras are also described. Brief comments are also made on the ElSalvadorean populations previously assigned to B. dunni.Key words: Amphibia: Caudata; Bolitoglossa dunni group; Honduras; Plethodontidae

THE five described species in the Boli- Larson (1 9 8 3 ~) emonstrated that mor-toglossa d u n n i group occur in mountain- phological differences exist between theous regions from Chiapas, MAxico to Hon- two color phases as well. We herein pro-duras (Elias, 1984). All specimens of the vide a redescription of B . d u n n i (the ho-B . d u n n i group from Honduras and El lotype of which represents the unicolorSalvador have been included in the species species) and describe the patterned phaseB. d u n n i (Wake and Lynch, 1976), which as a new species. In recent )ears , we havewas described by Schmidt (1933) from the also collected several series of salamanders"mountains west of San Pedro [Sula], Hon- of this group (including color notes andduras." The type series of B . d u n n i con- photographs of living specimens) fromtains two color morphs, one with a unicolor other mountain ranges in western Hon-reddish dorsum and the other with a dor- duras. Comparison of these specimens tosum of some shade of brown that is spotted the redefined B . d u n n i , the other speciesor longitudinally banded with a paler color contained in its type series, and to the(see Dunn, 1926, for a description of the Chiapan and Guatemalan species of thetype series, as O e d i p u s m o r i o ) . Larson d u n n i group, reveals that our collections(1983a:95), working with material col- also contain two additional undescribedlected near the type locality of B . d u n n i , species.made an important discovery when he de-tected that the two color phases were ". . . ATERI RIALS AND METHODSdistinct at the molecular level and . . . All measurements are in millimeters:probably represent[ed] distinct species." made to the nearest tenth with dial calipersRecently, we have collected specimens of and a dissecting microscope. Standardboth color phases (including color notes length (SL) = snout to posterior edge ofand photographs of living specimens) also vent and body length (BL) = gular fold tofrom near the type locality of B . d u n n i posterior edge of vent. Other measure-and elsewhere. An examination of these ments employed as ratios in the subsequent


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2 HERPETOLOGICA [Vol.49, No . 1

width (HW).All tooth counts are both sidessummed. Specimens collected by us forthis study have been deposited in the FieldMuseum of Natural Hi story , Chicago(FMKH), Museum of ha tu ral History,University of Kansas, Lawrence (KC),andthe Museum of Vertebrate Zoology, Uni-versity of California, Berkeley (MVZ).Dataon three species extralimital to Honduras(cuchumatana, engelhardti, and helm-richi) were taken from the literature (Eli-as, 1984; Larson, 1983b; Schmidt, 1936;Stuart, 1943; Wake and Brame, 1969; Wakeand Lynch, 1976). The data included forthe fourth extralimital species (rostrata)were taken from 15 specimens examinedby us (MVZ 130181-82, 130184-86,130188, 130209, 130237-38, 130249,130256, 130260, 130267-68, 130272, De-partamento de San Marcos, Guatemala).We have not examined most of the typeseries of B . dunni (sensu lato) but haverelied largely upon recently collected ma-terial to characterize the two species rep-resented in that series. An analysis ofcovariance (SuperANOVA", Abacus Con-cepts, Inc., Berkeley, California) was usedto test the distinctiveness of Honduranmembers of the dun ni group based on sev-eral morphological and meristic charactersheretofore used in the taxonomy of thisgroup. The dorsal ground colors of the re-cently collected material have been com-pared to the color swatches in Smithe(1975). The color numbers used below re-fer to the swatches in that publication.

A REDESCRIPTIOXFBOLITOGLOSSAUNNI SCHMIDT(Fig. 1A)

Ho1otype.-FMNH 4550, adult male,from mountains west of San Pedro Sula,approximately 1375 m elevation, Depar-tamento d e Cortks, Honduras, 5 May 1923,Karl P. Schmidt.Description.-Large (SL in six adultmales, 51.1-59.6, 2 = 56.7; 50.1-73.2, f =64.6 in 10 adult females), robust memberof B. dunni group; snout truncate; labialprotuberances well developed in both sex-

broad in adults (HW/SL in six males, 16.6-17.596, = 17.1; 16.0-18.5%, f = 17.3 in10 females); distinct groove extending be-low eye, following the curvature of theeye, not extending to lip; eyes slightly pro-tuberant, narrowly visible beyond marginof jaw when viewed from below in males,not visible in females; postorbital grooveshallow, irregular, extending posteriorlyfrom eye, proceeding sharply ventrallyposterior to mandible, extending acrossthroat anterior to gular fold; maxillary teethabundant (six adult males, 71-93, f = 82.2;10 adult females, 67-91, f = 76.8), ex-tending beyond center of eye, increasingin number with increasing size; vomerineteeth abundant (six adult males, 30-37, f= 33.2; 10 adult females, 30-40, i?4.3),in long, single, arched series, extendingslightly beyond outer edge of internal na-res; premaxillary teeth (six adult males, 3-6 , f = 4.3; 10 adult females, 6-10, a = 7.9)enlarged, piercing lip in adult males, short,located behind lip in females; tail slightlycompressed laterally, constricted basally;tail long, about equal to SL in adult males(TL/SL in six, 90.9-104.396, R = 96.8),shorter than SL in adult females (TL/SLin nine, 80.1-88.496, 2 = 85.0); ail alwayslonger than BL in adults (TL/BL in fivemales, 121.5-142.9%, f = 131.5; 106.8-119.796, f = 113.1 in nine females); limbsslender, moderately long; limb intervalvarying from zero to one costal fold inadult males, one-half to two in adult fe-males; webbing moderate, with one-halfto one and one-half phalanges of third dig-it on forelimbs free of webbing, one to oneand one-half phalanges between digits 3and 4 on hindlimbs free of webbing (onespecimen, MVZ 186761, has two phalangeson hindlimbs free); digits bearing well-de-veloped subterminal pads; digits on fore-limbs in order of decreasing length, 3-4-2-1, on hindlimbs, 3-4-2-5-1.Coloration in life.-Very little varia-tion in color exists in this species. FMNH236515: uniform Mahogany Red (color132B) covering entire dorsal and lateralsurfaces of head, body, and tail; ventraland subcaudal surfaces slightly paler Ma-hogany Red; small gold spots on ventral


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March 19931 HERPETOLOGICA 3

FIG.1.-(A) Adult female (FMNH 36515) of B. dunni; (B) adult male (KU 219841) of banded phase ofB. conanti; (C ) adult male (KU 219854) of spotted phase of B. conanti; (D) subadult female (KU 219898)of B. card; (E) adult female (KU 219903) of B. celaque.


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4 HERPETOLOGICA [ V o l .49, No . 1

I I I89 88 87

FIG.2.-Map showing distribution of the membersof the Bolitoglossa dunni group in western Hondurasand adjacent El Salvador examined for this study.Open squares = B . d u n n i and B. conant i ; closedcircles = populations tentatively assigned to B. co-n a n t i ; closed square = B. carri; triangles = B. celaque;inverted triangle = population tentatively assigned toB. ce laque .feet; iris mottled brown and MahoganyRed. FMNH 236519-20 identical toFMNH 236515.Specimens examined for redescription(20).--FMNH 236515-16, 236518-19,236521, MVZ 186761,186765-66,186770,adult females, FMNH 236517, 236520,adult males, MVZ 186768, 186771, juve-niles or subadults, all from QuebradaGrande, 1370-1680 m, Departamento deCopin. MVZ 186759, adult female, MVZ186726, 186756, juveniles, all from CerroCusuco, 1550-1560 m, Departamento deCortks. MVZ 115319, 132944, 132946,UMMZ 80933 (formerly FMNH 4549),adult males, MVZ 132945, 132948, sub-adults, all from mountains west of San Pe-dro Sula, 1200-ca. 1375 m, Departamentode Cortks.

Distribution and ecology .-This spe-cies is known only from the sierras EspirituSanto and Omoa in northwestern Hon-duras (Fig. 2). Bolitoglossa d un ni has been

Premontane Wet Forest formation (= Sub-tropical Wet Forest formation of Hold-ridge, 1967) and in the Lower MontaneWet Forest formation (Holdridge, 1967)from 1200-1680 m elevation. Most of thespecimens that we collected were insidearboreal bromeliads (Tillandsia sp.), butseveral were found at night on tree stumpson a cleared hillside. A single specimenwas found during the day in the rottencenter of a tree stump on this same hillside.Schmidt (1942) provided an account of thebromeliad-inhabiting habits of the type se-ries of this species, which also includedspecimens of the patterned phase that wehere describe asBolitoglossa conanti sp. nov(Fig. 1B,C)

Holot ype.-KU 219840, adult female,from Quebrada Grande (15"05'N,88"55'W), 1370m elevation, Departamen-to de CopLn, Honduras, 6 May 1988,JamesR. McCranie and Larry David Wilson.Original number LDW 8756.Paratypes (42).-KU 219849, 219851,219853,219855,219858,219863,219864-69, MVZ 186762-63, 186767, 186769,186777, adult females, KU 219841-48,219850, 219852, 219854, 219856-57,219859-62, adult males, all from Quebra-da Grande, 1370-1680 m. MVZ 186757-58, adult females, MVZ 186760, adult male,all from Cerro Cusuco, 1540-1560 m, De-partamento de Cortks, Honduras. MVZ128271-72, adult females, MVZ 13249-50,UMMZ 80933 (formerly FhINH 4555),adult males, all from mountains west ofSan Pedro Sula, ca. 1500-1570 m, Depar-tamento de Cortks, Honduras.Diagnosis.-Bolitoglossa conanti dif-fers from B. dunni in dorsal coloration (abroad rufous-colored longitudinal bandfrom tip of snout onto tail or gold blotchingor spotting on a sepia ground color versusuniform Mahogany Red), subcaudal col-oration (tan to dark brown versus pale Ma-hogan) Red), coloration of the dorsal sur-faces of feet (absence versus presence ofconspicuous gold spots), size (SL f = 46.7in 24 adult males and 48.0 in 26 adultfemales versus 56.7 in six adult males and


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5arch 19931 HERPETOLOGICAT.~BLE.-Analysis of covariance for the effect ofthe four Honduran species of the d u n n i group andstandard length (SL) on number of maxillary teeth.The covariate was SL in a simultaneous Type 111 sumof squares model and a Fisher's Protected LSD post-hoc all pairs comparison at the significance level of0.05. There was no significant interaction betweenthe species and SL (F = 0.09, P = 0.9649) so the in-teraction was deleted from the model. Because of thelarge adult size of d u n n i , the analysis includes datafrom six subadults of that species.

SL 1 4089.39 4089.39 164.57 0.0001Species 3 5712.90 1904.30 76.64 0.0001Error 137 3404.31 24.85versus D CD P

celaque carri 3.06 2.51 0.0175celaque conan t i 11.30 1.99 0.0001celaque d u n n i 23.74 2.51 0.0001carri conanti 8.24 2.55 0.0001carri dunni 20.68 2.97 0.0001conan t i d u n n i 12.44 2.55 0.0001

TABLE .-Analysis of covariance for the effect ofthe four Honduran species of the d u n n i group andstandard length (SL) on number of vomerine teeth.The methods are the same as those used in Table 1.There was no significant interaction between the spe-cies and SL ( F = 1.17, P = 0.3244) so the interactionwas deleted from the model.

df SS MS F PSL 1 408.44 408.44 50.05 0.0001Species 3 1979.97 659.99 80.87 0,0001Error 137 1118.07 8.16

versus D CD Pcelaque carri 1.85 1.44 0.0122celaque conan t i 7.75 1.14 0.0001celaque dunni 10.81 1.44 0.0001carri conan t i 5.90 1.46 0.0001carri d u n n i 8.96 1.70 0.0001c o n a n t i d u n n i 3.06 1 .46 0.0001

number plotted against SL (Table 1; Fig.31, vomerine tooth number plotted againstSL (Table 2; Fig. 41, relative HW (Table3; Fig. 51, relative TL in both sexes (Tables4, 5; Figs. 6,7) ,and in amount of webbing(1%-2 phalanges of longest toes free versus(%-I% on forelimbs and 1-1s on hind-limbs). Larson (1983~) as demonstratedmolecular differences between the twospecies. Bolitoglossa conanti and B, cu-chumatana can be distinguished by dorsalcoloration (a broad, pale middorsal lon-gitudinal band from tip of snout onto tailor blotched or spotted versus pale dorso-lateral stripes), subcaudal coloration (tanto dark brown with gold or silver flecking

2030 35 40 45 50 55 60 65 70 75

SL (MM)FIG.3.-Comparison of standard length (SL) and

maxillary tooth number in the Honduran species ofthe Bolitoglossa dunni group. Species codes are soliddiamonds = carri, y = 0 . 8 4 9 ~+ 13.9, n = 22; opensquares = celaque, y = 0 . 7 4 4 ~+ 13.5,n = 51; open

1 0 ) , , , , , , , , ,30 35 40 45 50 55 60 65 70 75SL (MM)

FIG.4.-Comparison of standard length (SL) andvomerine tooth number in the Honduran species ofthe Bolitoglossa dunni group. Species coded as inFig. 3; carri, y = 0 . 1 7 2 ~+ 15.1, n = 22; celaque, y


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TABLE .-Analysis of covariance for the effect ofthe four Honduran species of the durzni group andstandard length (SL) on head width (in mm). Themethods are the same as those used in Table 1 exceptthat only five subadult dt~ntziwere included. The lowP value for species plus SL is a result of significantinteraction between the species carri, celaque, andconanti.< If SS \!IS F P

SL 1 90.14 90.14 754.41 0.0001Species 3 1.12 0.37 3.13 0.0278Species+ SL 3 1.75 0.58 4.89 0.0029Error 136 16.25 0.11versus D CD P

celaque carri 0.04 0.17 0.6524celaque conanti 0.04 0.14 0.5507celaque dunni 2.16 0.18 0.0001carri conanti 0.01 0.18 0.9867carri dunni 2.20 0.21 0.0001conanti dztnni 2.20 0.18 0.0001

or spotting versus tan or pale orange-yel-low with gray mottl ing), amount of web-bing on hindlimbs (1%-2 phalanges be-tween digits three and four free versus M-l ) ,and in TI, (TL/SL % = 91.6% in 36adults versus 77.3% in four). BolitogEo.ssaconant i differs from B, engelhardt i in dor-sal coloration (patterned versus usuallyuniform), size (SL % = 44.4 in 50 adultsversus 39.9 in l o) , maxillary tooth numberin adults ( f = 60.5 in 47 versus 49.5 in lo),

5 / I35 40 45 50 55 60 65 70 75SL (MM)

FIG. 5.-Comparison of standard length (SL) andhead width in the Honduran species of the Bolito-glossa dunni group. Species coded as in Fig. 3; carri,y = 0.117~+ 2.3, n = 22; celaque, y = 0 . 1 2 3 ~+ 1.7,

TABLE4.-Analysis of covariance for the effect ofthe four Honduran species of the dunni group andstandard length (SL) on female tail length (in mm).The methods are the same as those used in Table 1except that only two subadult drinni were included.There was no significant interaction between the spe-cies and SL jF = 2.06, P = 0,1221) so the interactionwas deleted from the model.df SS MS F P

SLSpeciesError1341

2177.69237.94233.292177.6979.315.($9

382.7213.94 0,00010.0001versus D CD P

celaquccelaquecelaquecarricarriconanti

carriconantidunniconantidunnidunni

5.460.649.296.1014.758.65

2.371.902.112.152.331.84

0,00010.50160.00010,00010.00010,0001

and amount of webbing (1112-2 phalangesof longest toes free versus one). Boli to-glossa conanti and B. helmrichi can bedistinguished by subcaudal coloration (tanto dark brown versus orange) and inamount of webbing (195-2 phalanges oflongest toes free versus only digital tips).Bolitoglossa conanti differs from B. ros-trata in dorsal coloration (a broad, palelongitudinal band from t ip of snout ontotail or gold blotching or spotting versuspale dorsolateral stripes or occasionally

35 40 45 50 55 60 6 5 70 75SL (MM)

FIG. 6.-Comparison of standard length (SL) andtail length in females of the Honduran species in theBolitoglossa dunni group. Species coded as in Fig. 3;carri, y = 0 . 7 0 1 ~+ 0.4, n = 7; celaque, y = 0.974~


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I-arch 19931 HERPETOLOGICA

TABLE5 -Analysis of covariance for the effect ofthe four Honduran species of the dunni group andstandard length (SL) on male tail length ( in mm).The methods are the same as those used in Table 1except that no subadults of dunni with complete tailswere available There was no significant interactionbetween the species and SL (F= 1.49,P = 0 2288) so

the interaction was deleted from the model.

SL 1 1422.85 1422.85 203.10 0.0001Species 3 432.27 144.09 20.57 0.0001Error 49 343.29 7.01

versus D CD Pcelaque carri 6.84 2.06 0.0001celaque conanti 5.96 1.73 0.0001celaque dunni 14.57 2.48 0.0001carri conanti 12.80 2.10 0.0001carri dunni 21.41 2.75 0.0001conanti dunni 8.61 2.51 0.0001

uniform dorsum),subcaudal coloration (tanto dark brown with gold or silver fleckingor spotting versus uniform pale beige todull yellow), size (SL f = 46.7 in 24 adultmales and 48.0 in 26 adult females versus53.2 in four adult'males and 59.3 in 11adult females), and in amount of webbing(1%-2 phalanges of longest toes free versus> 2 ) .

Description.--Medium-sized (SL in 24adult males, 38.3-50.5, f = 46.7; 41 .O-60.7,i= 48.0 in 26 adult females) member ofB . dunni group; snout truncate; adult fe-males more robust than relatively slendermales; labial protuberances well-devel-oped in both sexes, pronounced in adultmales; oval-shaped mental glands presentin adult males; head relatively narrow inadults (HW/SL in 24 males, 14.5-17.596,f = 15.7; 14.7-18.1%, 1 = 16.1 in 25 fe-males); distinct groove extending beloweye, following the curvature of the eye,not extending onto lip; eyes slightly pro-tuberant, narrowly visible beyond marginof jaw when viewed from below in males,not visible in females; postorbital grooveshallow, irregular, extending posteriorlyfrom eye, proceeding sharply ventrallyposterior to mandible, extending acrossthroat anterior to gular fold; maxillary teethabundant (21 adult males, 50-73, f f = 59.9;26 adult females, 50-74, f = 61.0), ex-tending beyond center of eye, increasing

35 40 45 50 55 60SL (MM)

FIG.7.-Comparison of standard length and taillength in males of the Honduran species in the Bolito-glossa dunni group. Species coded as in Fig. 3; carri,y = 0 . 7 9 7 ~- 0.2,n = 10; celaque, y = 1 . 3 8 8 ~- 24.9,n = 20; conanti, y = 1 . 5 2 6 ~- 25.3, n = 18; dunni,y = 1 . 1 4 9 ~- 10.2, n = 6.teeth abundant (21 adult males, 20-32, f= 28.0; 26 adult females, 24-33, f f = 29.5),in long, single, arched series, extendingslightly beyond outer edge of internal na-res; premaxillary teeth (23 adult males, 2-7, f = 4.0; 26 adult females, 5-9, i= 6.7)enlarged, piercing lip in adult males, short,located behind lip in females; tail slightlycompressed laterally, constricted basally;tail long, about equal to SL in adult males(TL/SL in 18, 83.8-110.496, ff = 98.4),shorter than SL in adult females (TL/SLin 18, 79.3-91.4%, f = 84.8); tail longerthan BL in adults (TL/BL in 16 males,113.2-147.796, f = 132.5; 98.9-119.09'~~= 112.5 n 18 females); limbs slender, mod-erately long; limb interval varying fromzero to one costal fold in adult males, one-half to two in adult females; webbing mod-erate, with one and one-half phalanges oninside and one and one-half to two on out-side of third digit on forelimbs free of web-bing (one specimen, MVZ 186777, was re-corded as having one phalanx f ree), oneand one-half to two phalanges betweendigits three and four on hindlimbs free ofwebbing; digits discrete, bearing well-de-veloped subterminal pads; digits on fore-limbs in order of decreasing length. 3-4-2-1, on hindlimbs, 3-4-2-5-1.Coloration in life.-Coloration is veryvariable in this species. There are two color


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mediacy. One ~ h a s e onsists of a broad,pale longitudinal band from the tip of thesnout onto the dorsal surface of the tail(Fig. 1B). The other phase consists of darkbrown dorsal surfaces with varyingamounts of gold spotting or blotching (Fig.1C). The subcaudal surface has gold (oc-casionally silver) flecking or spotting on atan to dark brown ground color. KU219841: all dorsal surfaces with broad Or-ange-Rufous (color 132C) longitudinalband with widely scattered brown spots;sides of head and body Amber (color 36);dorsal surface of legs Amber with Orange-Rufous spotting; venter of body brown withgold flecking and spotting; chin yellowish-tan; subcaudal surface tan with gold fleck-ing and spotting. KU 219855: dorsum ofhead, body, and basal section of tail withbroad Pratt's Rufous (color 140) longitu-dinal band with slight brown spotting onhead; sides of head and body Amber (color36); ventral and subcaudal surfaces darkbrown with scattered gold flecking; irisPratt's Rufous. KU 219842: all dorsal sur-faces Sepia (color 219) with heavy goldflecking, suggestive of banding; additionallarge gold spots on dorsal surface of basalsection of tail; sides of head and body Sepiawith light gold flecking; ventral and sub-caudal surfaces dark brown with scatteredgold flecking; iris with gold flecking. KU219870: same as KU 219842 except thatventral and subcaudal surfaces with silverflecking. KU 219844,219856: same as KU219842 except with considerably less goldflecking on dorsal surfaces. KU 219880:same as KU 219844, 219856 except forlarge Orange-Rufous (color 132C) blotch-ing on basal section of dorsal surface oftail. KU 219854: all dorsal surfaces Amber(color 36) with gold blotching and spot-ting; sides of head and body Amber; ven-tral and subcaudal surfaces brown withgold flecking and spotting; iris gold withblack reticulations. KU 219843: all dorsalsurfaces Kingfisher Rufous (color 240) withwidely scattered gold flecking; Orange-Rufous (color 132C) blotching on dorsalsurface of basal section of tail; ventral andsubcaudal surface brown with scatteredgold flecking; iris mottled gold and brown.

DLOGICA [Vol . 49 , No. 1

KU 219879: all dorsal surfaces with Or-ange-Rufous (color 132C) longitudinalband with dense gold flecking throughout;sides of head and body Amber (color 36)with heavy gold flecking; ventral and sub-caudal surfaces brown with gold fleckingand spotting.Measurements of ho1otype.-HW 9.6;head length 14.8; head depth at posteriorangle of jaw 4.8; eyelid length 3.5 ; eyelidwidth 1.5; anterior rim of orbit to snout4.0; horizontal orbital diameter 3.1; inter-orbital distance 3.5; distance between vo-merine teeth and parasphenoid tooth patch0.7; snout to forelimb 17.0; distance sep-arating internal nares 3.2; distance sepa-rating external nares 3.5; snout projectionbeyond mandible 1.5; SL 60.7; BL 45.9;snout to anterior angle of vent 56.0; axillato groin 32.6; TL 53.3; tail width at base3.7; tail depth at base 4.1; forelimb length13.4; hindlimb length 14.3; width of rightforefoot 4.5; width of right hindfoot 5.5

Referred specimens.-KU 219870-91,MVZ 186772, juveniles or subadults, allfrom Quebrada Grande. MVZ 186755,subadult, from Cerro Cusuco.Comm en ts. -Two specimens (MCZ21245-46) from Portillo Grande. Depar-tamento de Yoro, Honduras (Fig. 2) aretentatively assigned to B. conanti. In pre-servative, these specimens have dark browndorsal surfaces with lighter brown spots.MCZ 21246 has 87 maxillary teeth, whichis 13 higher than the highest count ob-tained for conanti, MCZ 21245 has 69which is near the upper count for conanti .This population may represent a distinctspecies. Three specimens (KC' 219892,LACM 46940, El Portillo, 1900 m; MVZ186727, El Chagiiitbn, 1920 m) from theDepartamento de Ocotepeque, Hondurasand one (MVZ 200535, Hacienda Monte-cristo, 2250 m) from the Departamento deSanta Ana, El Salvador (Fig. 2) are alsotentatively assigned to B. conanti. Thesespecimens either had small light spots onthe dorsal surfaces in life (KC' 219892) orlighter dorsal surfaces (at least anteriorly)somewhat suggestive of the longitudinallybanded phase of B , conant i (LACM 46940,MVZ 186727,200535). MVZ 186727 (a ju-


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March 19931 HERPET(

venile) also has a dark X mark on the neckand shoulder region similar to those seenin several young specimens of the bandedphase of B, conanti. Mertens (1952:Fig.30) illustrated a subadult from HaciendaMontecristo with this mark. New materialis needed to determine the taxonomic sta-tus of this series. The Yoro and Ocotepequespecimens were not used in the analysis ofB. conanti.

Distribution and ecology.-This spe-cies is known for certain (see Commentsabove) from the sierras Espiritu Santo andOmoa in northwestern Honduras (Fig. 2).Bolitoglossa conanti has been collected inthe Lower Montane Wet Forest formation(Holdridge 1967) from 1370-1680 m ele-vation. Most of the specimens that we col-lected were taken from arboreal brome-liads (Tillandsia sp.), but a few were alsotaken at night from vegetation alongsidea small stream. Schmidt (1942) also re-ported finding this species in bromeliads.Bolitoglossa conanti occurs sympatricallywith B. dunni in the mountains west ofSan Pedro Sula (Schmidt, 1942), at Que-brada Grande, and at Cerro Cusuco. Thespecies is also sympatric with Nototritonnasalis at the latter locality and B. rufes-cens at Quebrada Grande. All of thesesympatric species also occur in bromeliadsat these localities.Etymology.-We take pleasure in nam-ing this species in honor of Roger Conant,near the time he also has been honored forhis contributions to herpetology with asymposium by the Society for the Studyof Amphibians and Reptiles. With thenaming of Bolitoglossa conanti, we alsowish to take note of the influence Dr. Co-nant has had on the field of herpetology.His famous Field Guide, first published in1958 and now in its third edition, is a mod-el of the artful combination of scholarlydetail and popular appeal. His importancein the field of herpetological husbandry isunquestioned, as a result of his career atthe Philadelphia Zoo. His scholarly pub-lications have been many and influential.Finally, his latest major work on Agkistro-don and its allies is a testament to his ded-

his friendship with his now-deceased coau-thor, Dr. Howard K. Gloyd.

Bolitoglossa carri sp. nov.(Fig. 1D)Ho1otype.-FMNH 236502, adult fe-male, from Cerro Cantagallo, near Le-paterique (14"06'N, 8'i028'W), 1840 m el-evation, Departamento de FranciscoMorazhn, Honduras, 18 August 1986, JamesR. McCranie, Kenneth L. Williams, andLarry David Wilson. Original number

LDW 8361.Paratypes (21 ).-- FMN H 236470,236472, 236490, 236500, 236512, K U219893-94, adult females, FMNH 236452,236454, 236473-74, 236476-78, 236491-93, 236503, KU 219895-97, adult males,all from Cerro Cantagallo, 1840-2070 m.Diagnosis.-Bolitoglossa carri differsfrom B, dunni in dorsal coloration (palebrownish dorsolateral stripes on a darkerbrown ground color versus uniform Ma-hogany Red), subcaudal coloration (me-lanophores present on a pale yellow topinkish-cream ground color versus paleMahogany Red without melanophores),coloration of the dorsal surfaces of feet(absence versus presence of conspicuousgold spots), size (SL T = 42.6 in 14 adultmales and 49.9 in eight adult females ver-sus 56.7 in six adult males and 64.6 in 10adult females), maxillary tooth numberplotted against SL (Table 1; Fig. 3), vo-merine tooth number plotted against SL(Table 2; Fig. 4), relative HW (Table 3;Fig. 5), relative TL in both sexes (Tables4, 5; Figs. 6, 7) ,and in amount of webbingon hindlimbs (1%-2 phalanges betweendigits three and four free versus 1-1%).Bolitoglossa carri and B. conanti differ indorsal coloration (pale dorsolateral stripesbeginning behind eye versus a broad, palelongitudinal band from tip of snout ontotail or blotched or spotted with gold), sub-caudal coloration (melanophores presenton a pale yellow to pinkish-cream groundcolor versus tan to dark brown with gold


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10 HERPETOLOGICA [Vol. 49, No. I

plotted against SL (Table 1; Fig. 3), vo-merine tooth number plotted against SL(Table 2; Fig. 4), and in relative T L inboth sexes (Tables 4, 5; Figs. 6, 7). Boli-toglossa carri can be distinguished from3. cuchunzatana by maxillary tooth num-ber in adults (f = 52.3 in 22 versus 60.2in 17) and amount of webbing (1%-2 pha-langes between digits 3 arid 4 on hindlimhsfree versus Vz-1). BoEitoglossa carri and B.engelhardti can be distinguished by dorsalcoloration (pale dorsolateral stripes versususually uniform), size (SL R = 45.2 in 22adults versus 39.9 in lo) , amount of web-bing (1-2 phalanges of digit 3 on forelimbsand 1'h-nearly 2 between digits 3 and 4on hindlimbs free versus 1 \ ,and by TL(TLiSL f = 75 7% in 17 adults versus 0.88in nine) Bolitoglossa carri and B. helm-richi can be distinguished by subcaudalcoloration (pale yellow to pinkish-crearnversus orange), maxillary tooth number inadults (2 = 52 3 in 22 versus 61.9 in 22j.amount of webbing (1-2 ~hala ng esf digit3 on forelimbs and 195-2 between digits 3and 4 on hindlimbs free versus only digitaltips), and by T L (TL/SL f = 75.7% in 17adults versus 86.2% in four) Rolitoglossacarri differs from B. rostrata in maxillarytooth number in adults (R = 52.3 in 22versus 61 2 in 15),amount of webbing \twoor less phalanges of longest toes free versus> 2 ) , size (SL, R = 42.6 in 14 adult malesand 49.9 in eight adult females versus 53.2in four adult males and 59 3 in I 1 adultfemales), and in T L JTL/SL f = 75.7% in17 adults versus 87.0% in 13).

Description.-Medium-sized (SL in 14adult males, 37.4-48 0, R = 42.6; 42.4-58.1,R = 49.9 in eight adult females) memberof B, du nn i group; snout truncate; adultfemales more robust than relatively slen-der males; labial protuberances well-de-veloped in both sexes, pronounced in adultmales; oval-shaped mental glands presentin adult males; head broad in adults (HW/SL in 14 males, 15.9-17.796,X = 17.1; 15.0-17 0%,z = 16.3 in eight females); distinctgroove extending below eye, following thecurvature of the eye, not extending to lip;eyes slightly protuberant, narrowly visiblebeyond margin of jaw when viewed frorn

regular, extending posteriorly from eye,~roc eedin g harply ventrally posterior tomandible, extending across throat anteriorto gular fold, some specimens have one ortwo additional shallow, irregular groovesbetween anteriormost groove and gularfold; maxillary teeth relatively few innumber (14 adult males, 46-60, R = 50.4;eight adult females, 47-68, f = 55.6) , ex-tending beyond center of eye, increasingin number with increasing size; vomerineteeth relatively few in number (14 adultmales, 18-24, R = 22.1; eight adult females,22-28, R = 24.4), in long, single, archedseries, extending slightly beyond outer edgeof internal nares; premaxillary teeth (14adult males, 2-6, f = 3.4; eight adult fe-males, 4-7, f = 5.4) enlarged, piercing lipin adult males, short, located behind l ip infemales; tail lightly compressed laterally,constricted basally; tail short, always lessthan SL (TL/SL in 10 adult males, 73.9-83.0%,R = 79.2; 6'7.9-74.1%, P = 70.6 inseven adult females), slightly shorter thanBL in adult females (TL/BI, in seven, 91.5-100.9%.f = 95.1), slightly longer than BLin adult tnales (TL/BL in 10, 100.0-115.8%, f = 107.8); limbs slender, mod-erately long; limb interval varying from0-1 costal fold in adult males, from one-half to two and one-half in adult females;webbing moderate, with one to one andone-half phalanges on inside and one andone-half to two on outside of third digiton forelimbs free of webbing, one and one-half to two phalanges between digits 3 and4 on hindlimbs free of webbing: digits dis-crete, bearing well-developed subterminalpads; digits on forelimbs in order of de-creasing length. 3-4-2-1, on hindlimbs, 3-4---51.

Coloration in life.-Coloration is some-what variable in this species. All adultspecimens had pale brownish dorsolateralstripes from the shoulder region to the endof the body on a darker brown groundcolor. The area between the dorsolateralstripes varied in the extent of pale colorpresent. Some specimens had only a nar-row, incomplete pale nliddorsal stripe,whereas others had extensive pale areas onthe middorsum that were mottled with


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11arch 19931 HERPETOLOGICA

ken in those specimens with extensive palemiddorsal areas. The top of the head wasalways dark brown without paler mark-ings. The dorsal surface of the tail hadvarying amounts of pale brounish areasover its entire length. Subcaudal colorationvaried from pale yellow with small, scat-tered melanophores to pinkish-cream witha heavy sprinkling of melanophores withor without scattered rust red mottling. KC2 19895-96, 2 19898-99: dorsal surfaces ofheads, middorsal regions, and lateral areasRaw Umber (color 23); dorsolateral stripesKingfisher Rufous (color 240); dorsal sur-faces of tails Kingfisher Rufous. KU 219893:similar to KU 219895-96, 219898-99, ex-cept that dorsolateral stripes and dorsalsurface of tails Pratt 's Rufous (color 140).KC' 219894: dorsal surfaces of head, nar-row middorsal region, and lateral areasDark Brownish Olive (color 129); diffusedorsolateral stripes and dorsal surface oftail Raw Sienna (color 136) F3INH237502: ventral and subcaudal surfacespale yellow with scattered melanophores,additional scattered rust red mottling pres-ent on subcaudal surface. FMNH 236472-73: ventral and subcaudal surfaces pinkish-cream with heavy sprinkling of melano-phores. additional scattered rust red mot-tling on subcaudal surface.Measurements of ho1otype.-HW 8.7;head length 14.4; head depth at posteriorangle of jaw 5.2; eyelid length 3.7; eyelidwidth 1.9; anterior rim of orbit to snout4 0; horizontal orbital diameter 3.5; inter-orbital distance 3.6; distance between vo-merine teeth and parasphenoid tooth patch0.5; snout to forelimb 18.4; distance sep-arating internal nares 2.8; distance sepa-rating external nares 3.4; snout projectionbeyond mandible 1.5; SL 58.1; BL 43.7;snout to anterior angle of vent 53.5; axillato groin 31.1; TL 40.0; tail width at base3 2; tail depth at base 4.4; forelimb length13.4; hindlimb length 14.5; width of rightforefoot 4.3; width of right hindfoot 5.6.Referred specimens.-FMNH 236416-24, 236427-32, 236434-39, 236453,236455-58, 236475, 236499, KU 219898-901, all juveniles or subadults from CerroCantagallo.

(Fig. 2). Bolitoglossa carri has been col-lected from the Lower Montane Moist For-est formation (Holdridge, 1967) from1840-2070 m elevation. The former ele-vation was the lowest at which we workedin the area and the latter is at the summitof the highest peak in the region. Mostspecimens were collected inside bromeli-ads (Tillandsin sp.),but several were foundat night sitting on broad leaves growingon shrubs next to a small stream: and twowere underneath the bark of a large, fallentree in a clearing. The bromeliads con-taining salamanders were usually on trees,but a few salamanders were also found inbromeliads growing on the ground. Thesalamanders were usually found one to abromeliad: but in a few instances t\vo oc-curred in a single bromeliad; in two casesthree individuals were found in the samebromeliad. No other species of salamanderhas been collected on Cerro Cantagallo.

Etymology.-The species carri is namedin honor of the late Archie F. Carr. whotaught for five years at the Escuela Agrico-la Panamericana in El Zamorano, Hon-duras, not far from the type locality of thespecies. During his stay, Dr. Carr amassedan important collection of Honduran am-phibians and reptiles which is now housedat the American Museum of Natural His-tory. Dr. Carr was a naturalist and con-servationist extraordinaire with an envia-ble gift with words whose reputationextended far beyond the limits of herpe-tology.

Bolitoglossa celaque sp. nov.(Fig. 1E)Ho1otype.-FMNH 236504, adult fe-male, from near the Rio ArcAqual, easternside of Cerro Celaque, Cordillera de Ce-laque (14"32'11:, 88"40tW), 2480 m eleva-tion, Departamento de Lempira, Hondu-ras, 1 August 1985, James R . McCranie,Kenneth L. Williams, and Larry DavidWilson. Original number LDW 6577.Paratypes ( 5 1 ) . - FMNH 236485,236488,236506-07,236509, KU 219902-03, h4VZ 186728, 186734, 186737, adultfemales, FMNH 236482, 236487, 236489,236505,236508,236510-11, MVZ 186729-


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12 HERPETOLOGICA [Vol.49, N o. 1from the eastern side of Cerro Celaque,1930-2620 m. KU 219905-06,219910-12,219914, adult females, KU 219904,219907-09, 219913, adult males, all fromSan Pedro, 1990 m, Departamento d e In-tibucii, Honduras. FMNH 236479,236496,236513, adult females, FMNH 236480,236494-95, 236497-98, 236514, KU219915, MVZ 186751, adult males, all fromZacate Blanco, 2070-2150 m, Departa-mento d e Intibuch, Honduras.Diagnosis.-Bolitoglossa celaque dif-fers from B. d u n n i in dorsal coloration(russet with minute gold flecking versusuniform Mahogany Red), subcaudal col-oration (pale yellow to dark orange versuspale Mahogany Red), coloration of thedorsal surfaces of feet (absence versus pres-ence of conspicuous gold spots), size (SL f= 47.7 in 32 adult males and 48.3 in 20adult females versus 56.7 in six adult malesand 64.6 in 10 adult females), maxillarytooth number plotted against SL (Table 1;Fig. 3 ), vomerine tooth number plottedagainst SL (Table 2; Fig. 4) , relative HW(Table 3; Fig. 5) , relative TL in both sexes(Tables 4, 5; Figs. 6, 7), and in amount ofwebbing (usually two phalanges of longesttoes free versus %-I% phalanges of digit3 on forelimbs and 1-1% between digits 3and 4 on hindlimbs). Bolitoglossa celaqueand B . co n a n t i differ in dorsal coloration(russet with minute gold flecking versus abroad, pale longitudinal band from tip ofsnout onto tail or with gold spotting orblotching on a sepia ground color), sub-caudal coloration (gray or orange punc-tations on a pale yellow to dark orangeground color versus gold flecking or spot-ting on a tan to dark brown ground color),maxillary tooth number plotted against SL(Table 1; Fig. 3 ), vomerine tooth numberplotted against SL (Table 2; Fig. 4 ), rela-tive TL in males (Table 5; Fig. 7), and inamount of webbing (usually two phalangesof longest toes free versus 1%-2). Bolito-glossa celaque differs from B . carri in dor-sal coloration (russet with minute goldflecking versus pale dorsolateral stripes),maxillary tooth number plotted against SL(Table 1;Fig. 3), vomerine tooth numberplotted against SL (Table 2; Fig. 4) , rela-

7 ) ,and in amount of webbing (usually twophalanges of longest toes free versus 1-2on digit 3 of forelimbs and 1Yz-2 betweendigits 3 and 4 on hindlimbs). Bolitoglossacelaque and B . cu ch u ma t a n a can be dis-tinguished by dorsal coloration (russet withminute gold flecking versus pale dorsolat-era1 stripes), maxillary tooth number inadults (f = 49.2 in 51 versus 60.2 in 17),and by amount of webbing (usually twophalanges between digits 3 and 4 on hind-limbs free versus %-1). Bolitoglossa ce-laque can be distinguished from B. enge l -hard t i by size (SL f = 47.9 in 52 adultsversus 39.9 in 10) and amount of webbing(usually two phalanges of longest toes freeversus one). Bolitoglossa celaque and B .he lmr ich i can be distinguished by dorsalcoloration (russet with minute gold fleck-ing versus patterned with stripes, spots, orblotches), maxillary tooth number in adults(f = 49.2 in 51 versus 61.9 in 22), andamount of webbing (usually two phalangesof longest toes free versus only digital tips).Bolitoglossa celaque differs from B. ros-trata in dorsal coloration (russet with mi-nute gold flecking versus usually with paledorsolateral stripes), size (SL f = 47.7 in32 adult males and 48.3 in 20 adult femalesversus 53.2 in four adult males and 59.3in 11adult females), maxillary tooth num-ber in adults (f = 49.2 in 51 versus 61.2in 13), and amount of webbing (usuallytwo phalanges of longest toes free versus>2) .Description.-Medium-sized (SL in 32adult males, 37.2-55.1, f = 47.7; 36.6-62.2,f = 48.3 in 20 adult females) member ofB . d u n n i group; snout truncate; adult fe-males more robust than relatively slendermales; labial protuberances well-devel-oped in both sexes, pronounced in adultmales; oval-shaped mental glands presentin adult males; head relatively narrow inadults (HW/SL in 32 males, 14.7-17.2%,f = 15.9; 14.4-18.0%, f = 16.0 in 20 adultfemales); distinct groove extending beloweye, following the curvature of the eye,not extending to lip; eyes slightly protu-berant, narrowly visible beyond margin ofjaw when viewed from below in males, notor only barely visible in females; postor-


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posteriorly from eye, proceeding sharplyventrally posterior to mandible, extendingacross throat anterior to gular fold; max-illary teeth relatively few in number (32adult males, 40-58, f = 50.1; 19 adult fe-males, 25-62, f = 47.8), extending beyondcenter of eye, increasing in number withincreasing size; vomerine teeth relativelyfew in number (32 adult males, 15-26, f= 20.8; 19 adult females, 14-30, f = 21.6),in long, single, arched series, extendingslightly beyond medial border of internalnares; premaxillary teeth (32 adult males,1-5, f = 3.6; 19 adult females, 3-8, f =6.0) enlarged, piercing lip in adult males,short, located behind lip in females; tailslightly compressed laterally, constrictedbasally; tail relatively long, always less thanSL (TL/SL in 20 adult males, 70.4-98.792,f = 85.4; 62.9-86.892, f = 79.0 in 10 adultfemales), usually more than BL (TL/BLin 20 adult males, 97.0-132.8%, f = 115.1;83.1-112.4%, f = 103.7 in 10 adult fe-males); limbs slender, moderately long;limb interval varying from zero to onecostal fold in males, zero to two and one-half in females; webbing slightly reduced,with one and one-half to two (most oftentwo) phalanges on both sides of digit 3 onforelimbs free of webbing, one and one-half to little over two (most often two)phalanges between digits 3 and 4 on hind-limbs free of webbing; digits discrete,bearing well-developed subterminal pads,digits on forelimbs in order of decreasinglength, 3-4-2-1, on hindlimb, 3-4-2-5-1.

Coloration in life.-Only minor vari-ation in dorsal coloration, consisting ofamount and color of flecking, has beenrecorded in this species. The subcaudalcoloration varied from pale yellow to darkorange. KU 219902: all dorsal surfacesRusset (color 34), with minute gold flecks;dorsal surface of tail with heavier goldflecking; lateral portion of body Russet,with copper flecking; ventral and subcau-dal surfaces pale yellow, with ocherblotching and scattered minute gold fleck-ing; iris irridescent copper. KU 219903:similar to KU 219902, but with less goldand copper flecking. KU 219915 and

over bodies, feet, and basal section of tails;ventral surfaces of heads and bodies paleorange; subcaudal surfaces pale yellow withgray and orange punctations. KU 219904:similar to KU 219915 and FMNH 236494,except that gold flecking especially heavyon eyelids and snout; iris also heavilyflecked with gold. FMNH 236480: similarto KU 219915 and FMNH 236494, exceptthat ventral surfaces of head and body paleyellow and subcaudal surface pale orange.FMNH 236459: all dorsal surfaces Russet(color 34), with minute chocolate brownflecking dorsally, silver flecking laterally;subcaudal surface dark orange, with grayand silver flecking. The minute gold fleck-ing on the dorsum quickly disappears inpreservative. All specimens have uniformdorsal surfaces in preservative exceptFMNH 236488 which has incomplete, paledorsolateral stripes.Measurements of ho1otype.-HW 9.5;head length 13.8; head depth at posteriorangle of jaw 5.8; eyelid length 4.2; eyelidwidth 2.1; anterior rim of orbit to snout4.3; horizontal orbital diameter 3.9; inter-orbital distance 3.5; distance between vo-merine teeth and parasphenoid tooth patch0.5; snout to forelimb 17.4; distance sep-arating internal nares 2.6; distance sepa-rating external nares 3.3 ; snout projectionbeyond mandible 1.5 ; SL 62.2; BL 48.4;snout to anterior angle of vent 58.7; axillato groin 34.0; TL 50.9; tail width at base4.6; tail depth at base 5.1; forelimb length16.0; hindlimb length 17.0; width of rightforefoot 6.3; width of right hindfoot 8.1.

Referred specimens.-FMNH 236425-26, 236440-51, 236469, 236471, 236481,236483-84,236486,236501, MVZ 186732,186740, 186743-44, 186747-50, juvenilesor subadults from Cerro Celaque. FMNH236459-67, KU 219916 juveniles or sub-adults from Zacate Blanco. FMNH 236433,236468, MVZ 186752-54, juveniles or sub-adults from the Sierra de Montecillos southof Tutule, 2100-2160 m, Departamento deLa Paz. MVZ 39733-34,39738,39742-43,39745,39748,39752,39765,39767,39772,adult females, MVZ 39735-36, 39739,39744, 39759, 39761-62, 39766, 39768,


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14 HERPETOLOGICA [Vol.49, No. 1

39760,39763-64,39769,39773,39775, ju-veniles or subadults, all from Los Eses-miles, 2440-2720 m, Departamento deChalatenango, El Salvador (see Commentbe lo^ ) .Comment.-Wake and Lynch (1976)assigned all El Salvadorean salamanders ofthe dunni group to B , dunni. The seriesfrom Los Esesmiles (Fig. 2) that we ex-amined are similar to B. celaque in size,in being unpatterned in preserkatibe, andin maxillary and vomerine tooth numbers.Howeker, they differ from celaque in hav-ing less webbing (over two phalanges oflongest toes free versus ~lsually wo) andhaving broader heads (pair u ise AN-COYJ\,F =9 47, P =0.0030).These spec-imens are tentative]!, assigned to B , ce-laque, but were not used in the analysis ofthat species.Distribution and ecology.-This spe-cies is known in southwestern Hondurasfrom four isolated mountain ranges in thecordilleras Celaque, Opalaca, and klon-tecillos (Fig. 2). Bolitoglossa celaque hasbeen collected from the Lower MontaneMoist Forest formation (Holdridge, 1967)from 1930-2620 m elevation. The speciesis both arboreal and terrestrial. -4t CerroCelaque, specimens were collected b j dayunder moss mats both on the ground andon tree trunks and in both terrestrial andarboreal bromeliads (the latter Tillandsiusp , the former an unidentified spiny spe-cies) -4single specimen was collected onCerro Celaque as it walked across the wetsurface of leaf litter in the ear ly afternoon.At night, B. celaque was commonly seenon leaves and rocks alongside streams onCerro Celaque All specimens, except forone that was under a log, from the otherthree localities for this species were takenby day from arboreal bromeliads (Til-landsia sp ) . No other species of salaman-der has been found with B. celaque.Etymology.-The name celaque is usedas a noun in apposition. It refers to thename of the highest mountain in Hondu-ras, Cerro Celaque, which is also the typelocality of the species The mountain'sname, according to Cruz (1986), s derivedfrom the Aztec word celac or ceelac,

may be in reference to the frigid Rio Ar-ciqual , which cascades off the eastern faceof Cerro Celaque.

Elias (1984) made an important contri-bution to our knowledge of the systematicsof the Middle American members of theBolitoglossa beta group when he recog-nized that two of the species groups (thehelmrichi and rostrata groups) of thesesalamanders as recognized by Wake andLynch (1976) were actually " . . . large,heterogeneous and united by no specificcharacters" (p . 14). He then subdividedand reorganized these t\vo groups into vrhathe felt were ". . . tight clusters of pheno-typically similar species" (Elias, 1984:14).One such assemblage (with five describedspecies, plus one undescribed Guatemalanspecies) was the dunni group ". . . char-acterized by blunt rounded toe tips, fullydeveloped subdigital pads, and a darkbrown ground color, frequently markedwith a lighter brown dorsal swath or pairedshoulder stripes . . ." (Elias, 1984:16).Lar-son (1983a), using protein comparisons,concluded that phylogenetic inferenceswere compatible with the species groupsrecognized by Elias (1984) and stated thatElias' dunni group ". . . appear[s] to forma monophyletic lineage whose primary di-chotomy is approximately 12 million yearsold" (Larson, 1983a:97).The three new species described in thispaper clearly are members of the dunn igroup as defined by Elias (1984).However,because of the uncertain taxonomic statusof the Honduran and El Salvadorean pop-ulations commented upon above. we feelthat any discussion about the relationshipsof B. carri, celaque, conanti, and dunniwithin the dunni group would be pre-mature at this time. In addition, a bio-chemical analysis of carri and celaque isneeded before the genetic distances of thesespecies from conanti and dunni can bedetermined.

Acknowledgments.-Valuable field assistance \+,asprovided by G. A . Cruz, M . R . Espinal, K . Hogan, L.Porras, J . C. Rindfleish, and K. L. Williams. Cruz alsodonated his Cerro Celaque collection of salamanders,


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15a r c h 19931 HERPETOLOGICA

Aguilar, G. A. Cruz, and S. Midence for their helpin securing collecting permits from the Departamen-to de Recursos Naturales, Tegucigalpa. We thank thefollowing people and their museums for the loan ofpertinent material; Museum of Comparative Zoology,Harvard University (MCZ)-J. P. Rosado and P. Al-berch; Museum of Vertebrate Zoology, University ofCalifornia, Berkeley (MVZ)-D. Wake, B. Stein, andK. Autumn; Natural History Museum of Los AngelesCounty (LACM)-J. W. Wright; University of Mich-igan, Museum of Zoology (UMMZ)-A. G. Kluge andG. Schneider.

CRCZ,G. A. 1986. Guia de 10s parques nacionales,refugios de vida silvestre, reservas biologicas y mon-umentos naturales de Honduras. Asociaci6n Hon-durefia de Ecologia. Tegucigalpa.

Dux s, E. R. 1926 . The Salamanders of the FamilyPlethodontidae. Smith College, Northampton,h.iassachusetts.ELIAS, . 1983. Salamanders of the northwesternhighlands of Guatemala. Nat. Hist. Mus. Los An-geles. Co. Contrib. Sci. 348: l - 20 .

HOLDRIDGE,. R. 1967. Life Zone Ecology. Re-vised edition. Tropical Science Center, San Josi.,Costa Rica.

L.\RSON,4. 1983a. A molecular phylogenetic per-spective on the origins of a lowland tropical sala-mander fauna. I. Phylogenetic inferences from pro-tein comparisons. Herpetologica 39:85-99.. 1983b. A molecular phylogenetic perspec-tive on the origins of a lowland tropical salamander

fauna. 11. Patterns of morphological evolution. Evo-lution 37:1141-1153.MERTENS,R. 1952. Die amphibien und reptilienvon El Salvador, auf grund der reisen von R. Mer-tens und A. Zilch. Abh. Senckenberg, h'atuf. Ges.487:1-120.

SCHMIDT, . P. 1933. New reptiles and amphibiansfrom Honduras. Zool. Ser. Field hlus. Nat. Hist. 2 0 :15-22.. 1936. Guatemalan salamanders of the ge-nus O e d i p u s . Zool. Ser. Field Mus. Nat. Hist. 2 0 :135-166.1942. A cloud forest camp in Honduras.Chicago Naturalist 3:23-30.

SMITHE,F. B. 1975. Naturalist's Color Guide.American Museum of Natural History, New York.STUART,L. C. 1943. Taxonomic and geographiccomments on Guatemalan salamanders of the ge-nus O e d i p u s . Misc. Publ. Mus. Zool. Univ. Michi-gan 5 6 : l - 3 3 .WAKE,D. B., AND A . H. BR.\ME,R . 1969. System-atics and evolution of neotropical salamanders ofthe Bolitoglossa helmrichi group. Contrib. Sci. Los.4ngeles Co. Mus. Nat. Hist. 175: l - 40 .W.AKE,D. B., AND J. F. LYSCH. 1976 . The distri-bution, ecology, and evolutionary history of pleth-odontid salamanders in tropical America. Nat. Hist.Mus. Los Angeles Co. Sci. Bull. 25: l - 65 .

Accep ted : 30 March 1992Associate Editor: David Cannatella


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You have printed the following article:

A Review of the Bolitoglossa dunni Group (Amphibia: Caudata) from Honduras with theDescription of Three New Species

James R. McCranie; Larry David Wilson

Herpetologica, Vol. 49, No. 1. (Mar., 1993), pp. 1-15.

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Literature Cited

A Molecular Phylogenetic Perspective on the Origins of a Lowland Tropical SalamanderFauna. II. Patterns of Morphological Evolution

Allan Larson

Evolution, Vol. 37, No. 6. (Nov., 1983), pp. 1141-1153.

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1993 mccranie & wilson b dunni group

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