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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
Herpetologtca, 49(1), 1993, 1-15 6 1993 b\ The Herpetolog~sts'League, Inc 2

MARCH 1993

KO. 1

A REVIEW OF THE BOLITOGLOSSA DUNNI GROUP

(AMPHIBIA: CAUDATA) FROM HONDURAS WITH THE DESCRIPTION OF THREE NEW SPECIES
JAMES

R. ~ ~ X ~ A N I ELARRY AND ' DAVIDWILSON~

'10770 SW 164th Street, Miami, FL 33157, U S A


2Department of Biology, South Campus,
Miami-Dade Community College, Miami, F L 33176, U S A

ABSTKACT: The type series of Bolitoglossa dunni Schmidt contains representatives of two distinct species of salamanders. Bolitoglossa dunni is redescribed and a description of the species heretofore subsumed in dunni is provided. In addition, two new species of this group from cloud forest localities in southwestern Honduras are also described. Brief comments are also made on the El Salvadorean populations previously assigned to B. dunni.

Key words:

Amphibia: Caudata; Bolitoglossa dunni group; Honduras; Plethodontidae

THE five described species in the Bolitoglossa d u n n i group occur in mountainous regions from Chiapas, MAxico to Honduras (Elias, 1984). All specimens of the B. d u n n i group from Honduras and El Salvador have been included in the species B. d u n n i (Wake and Lynch, 1976), which was described by Schmidt (1933) from the "mountains west of San Pedro [Sula], Honduras." The type series of B. d u n n i contains two color morphs, one with a unicolor reddish dorsum and the other with a dorsum of some shade of brown that is spotted or longitudinally banded with a paler color (see Dunn, 1926, for a description of the type series, as Oedipus morio). Larson (1983a:95), working with material collected near the type locality of B. d u n n i , made an important discovery when he detected that the two color phases were ". . . distinct at the molecular level and . . . probably represent[ed] distinct species." Recently, we have collected specimens of both color phases (including color notes and photographs of living specimens) also from near the type locality of B. d u n n i and elsewhere. An examination of these specimens plus the material studied by

Larson ( 1 9 8 3 ~ ) demonstrated that morphological differences exist between the two color phases as well. We herein provide a redescription of B. d u n n i (the holotype of which represents the unicolor species) and describe the patterned phase as a new species. In recent )ears, we have also collected several series of salamanders of this group (including color notes and photographs of living specimens) from other mountain ranges in western Honduras. Comparison of these specimens to the redefined B. d u n n i , the other species contained in its type series, and to the Chiapan and Guatemalan species of the d u n n i group, reveals that our collections also contain two additional undescribed species.
ATERI RIALS AND METHODS

All measurements are in millimeters: made to the nearest tenth with dial calipers and a dissecting microscope. Standard length (SL) = snout to posterior edge of vent and body length (BL) = gular fold to posterior edge of vent. Other measurements employed as ratios in the subsequent discussion are tail length (TL) and head

HERPETOLOGICA

[Vol. 49, No. 1

width (HW).All tooth counts are both sides summed. Specimens collected by us for this study have been deposited in the Field Museum of Natural History, Chicago (FMKH), Museum of hatural History, University of Kansas, Lawrence (KC),and the Museum of Vertebrate Zoology, University of California, Berkeley (MVZ).Data on three species extralimital to Honduras (cuchumatana, engelhardti, and helmrichi) were taken from the literature (Elias, 1984; Larson, 1983b; Schmidt, 1936; Stuart, 1943; Wake and Brame, 1969;Wake and Lynch, 1976). The data included for the fourth extralimital species (rostrata) were taken from 15 specimens examined by us (MVZ 130181-82, 130184-86, 130188, 130209, 130237-38, 130249, 130256, 130260, 130267-68, 130272, Departamento de San Marcos, Guatemala). We have not examined most of the type series of B. dunni (sensu lato) but have relied largely upon recently collected material to characterize the two species represented in that series. An analysis of covariance (SuperANOVA", Abacus Concepts, Inc., Berkeley, California) was used to test the distinctiveness of Honduran members of the dunni group based on several morphological and meristic characters heretofore used in the taxonomy of this group. The dorsal ground colors of the recently collected material have been compared to the color swatches in Smithe (1975). The color numbers used below refer to the swatches in that publication. A REDESCRIPTIOX OF BOLITOGLOSSAU N N I SCHMIDT D (Fig. 1A) Ho1otype.-FMNH 4550, adult male, from mountains west of San Pedro Sula, approximately 1375 m elevation, Departamento de Cortks, Honduras, 5 May 1923, Karl P. Schmidt. Description.-Large (SL in six adult males, 51.1-59.6, 2 = 56.7; 50.1-73.2, f = 64.6 in 10 adult females), robust member of B. dunni group; snout truncate; labial protuberances well developed in both sexes, pronounced in adult males; oval-shaped mental glands present in adult males; head

broad in adults (HW/SL in six males, 16.617.596, = 17.1; 16.0-18.5%, f = 17.3 in 10 females); distinct groove extending below eye, following the curvature of the eye, not extending to lip; eyes slightly protuberant, narrowly visible beyond margin of jaw when viewed from below in males, not visible in females; postorbital groove shallow, irregular, extending posteriorly from eye, proceeding sharply ventrally posterior to mandible, extending across throat anterior to gular fold; maxillary teeth abundant (six adult males, 71-93, f = 82.2; 10 adult females, 67-91, f = 76.8), extending beyond center of eye, increasing in number with increasing size; vomerine teeth abundant (six adult males, 30-37, f = 33.2; 10 adult females, 30-40, i= 34.3), ? in long, single, arched series, extending slightly beyond outer edge of internal nares; premaxillary teeth (six adult males, 36 , f = 4.3; 10 adult females, 6-10, a = 7.9) enlarged, piercing lip in adult males, short, located behind lip in females; tail slightly compressed 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/SL in nine, 80.1-88.496, 2 = 85.0);tail always longer than BL in adults (TL/BL in five males, 121.5-142.9%, f = 131.5; 106.8119.796,f = 113.1 in nine females); limbs slender, moderately long; limb interval varying from zero to one costal fold in adult males, one-half to two in adult females; webbing moderate, with one-half to one and one-half phalanges of third digit on forelimbs free of webbing, one to one and one-half phalanges between digits 3 and 4 on hindlimbs free of webbing (one specimen, MVZ 186761, has two phalanges on hindlimbs free); digits bearing well-developed subterminal pads; digits on forelimbs in order of decreasing length, 3-42-1, on hindlimbs, 3-4-2-5-1. Coloration in life.-Very little variation in color exists in this species. FMNH 236515: uniform Mahogany Red (color 132B) covering entire dorsal and lateral surfaces of head, body, and tail; ventral and subcaudal surfaces slightly paler Mahogany Red; small gold spots on ventral surface; limbs Mahogany Red, with conspicuous gold spots on dorsal surfaces of

March 19931

HERPETOLOGICA

FIG.1.-(A) Adult female (FMNH 236515) of B. dunni; (B) adult male (KU 219841) of banded phase of B. 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.

HERPETOLOGICA

[Vol. 49, No. 1

Premontane Wet Forest formation (= Subtropical Wet Forest formation of Holdridge, 1967) and in the Lower Montane Wet Forest formation (Holdridge, 1967) from 1200-1680 m elevation. Most of the specimens that we collected were inside arboreal bromeliads (Tillandsia sp.), but several were found at night on tree stumps on a cleared hillside. A single specimen was found during the day in the rotten center of a tree stump on this same hillside. Schmidt (1942) provided an account of the bromeliad-inhabiting habits of the type series of this species, which also included specimens of the patterned phase that we here describe as Bolitoglossa conanti sp. nov (Fig. 1B,C) Holot ype.-KU 219840, adult female, from Quebrada Grande (15"05'N, 88"55'W), 1370 m elevation, Departamento de CopLn, Honduras, 6 May 1988,James R. McCranie and Larry David Wilson. Original number LDW 8756. Paratypes (42).-KU 219849, 219851, 219853,219855,219858,219863,21986469, MVZ 186762-63, 186767, 186769, 186777, adult females, KU 219841-48, 219850, 219852, 219854, 219856-57, 219859-62, adult males, all from Quebrada Grande, 1370-1680 m. MVZ 18675758, adult females, MVZ 186760, adult male, all from Cerro Cusuco, 1540-1560 m , Departamento de Cortks, Honduras. MVZ 128271-72, adult females, MVZ 13249-50, UMMZ 80933 (formerly FhINH 4555), adult males, all from mountains west of San Pedro Sula, ca. 1500-1570 m, Departamento de Cortks, Honduras. Diagnosis.-Bolitoglossa conanti differs from B. dunni in dorsal coloration (a broad rufous-colored longitudinal band from tip of snout onto tail or gold blotching or spotting on a sepia ground color versus uniform Mahogany Red), subcaudal coloration (tan to dark brown versus pale Mahogan) Red), coloration of the dorsal surfaces of feet (absence versus presence of conspicuous gold spots), size (SL f = 46.7 in 24 adult males and 48.0 in 26 adult females versus 56.7 in six adult males and 64.6 in 10 adult females), maxillary tooth

I 89

I
88

I
87

FIG.2.-Map showing distribution of the members of the Bolitoglossa d u n n i group in western Honduras and adjacent El Salvador examined for this study. Open squares = B. d u n n i and B. conanti; closed circles = populations tentatively assigned to B. conanti; closed square = B. carri; triangles = B. celaque; inverted triangle = population tentatively assigned to

B. celaque.

feet; iris mottled brown and Mahogany Red. FMNH 236519-20 identical to FMNH 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, juveniles or subadults, all from Quebrada Grande, 1370-1680 m , Departamento de Copin. MVZ 186759, adult female, MVZ 186726, 186756, juveniles, all from Cerro Cusuco, 1550-1560 m, Departamento de Cortks. MVZ 115319, 132944, 132946, UMMZ 80933 (formerly FMNH 4549), adult males, MVZ 132945, 132948, subadults, all from mountains west of San Pedro Sula, 1200-ca. 1375 m, Departamento de Cortks. Distribution a n d ecology .-This species is known only from the sierras Espiritu Santo and Omoa in northwestern Honduras (Fig. 2). Bolitoglossa dunni has been collected in the upper elevations of the

March 19931

HERPETOLOGICA

T.~BLE 1.-Analysis of covariance for the effect of the four Honduran species of the d u n n i group and standard length (SL) on number of maxillary teeth. The covariate was SL in a simultaneous Type 111 sum of squares model and a Fisher's Protected LSD posthoc all pairs comparison at the significance level of 0.05. There was no significant interaction between the species and SL ( F = 0.09, P = 0.9649) so the interaction was deleted from the model. Because of the large adult size of dunni, the analysis includes data from six subadults of that species.

TABLE 2.-Analysis of covariance for the effect of the four Honduran species of the d u n n i group and standard 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 species and SL ( F = 1.17, P = 0.3244) so the interaction was deleted from the model.
df
SS
MS

SL Species Error

1 3 137

408.44 408.44 1979.97 659.99 1118.07 8.16


D

50.05 80.87
CD

0.0001 0,0001
P

SL Species Error

versus

1 4089.39 4089.39 164.57 0.0001 3 5712.90 1904.30 76.64 0.0001 137 3404.31 24.85
versus

CD

celaque celaque celaque carri carri conanti

carri conanti dunni conanti dunni dunni

3.06 11.30 23.74 8.24 20.68 12.44

2.51 1.99 2.51 2.55 2.97 2.55

0.0175 0.0001 0.0001 0.0001 0.0001 0.0001

celaque celaque celaque carri carri conanti

carri conanti dunni conanti dunni dunni

1.85 7.75 10.81 5.90 8.96 3.06

1.44 1.14 1.44 1.46 1.70 1.46

0.0122 0.0001 0.0001 0.0001 0.0001 0.0001

number plotted against SL (Table 1; Fig. 31, vomerine tooth number plotted against SL (Table 2; Fig. 41, relative HW (Table 3; Fig. 51, relative TL in both sexes (Tables 4 , 5 ; Figs. 6 , 7 ) , and in amount of webbing (1%-2 phalanges of longest toes free versus (%-I% on forelimbs and 1 1 on hind- s limbs). Larson ( 1 9 8 3 ~ has demonstrated ) molecular differences between the two species. Bolitoglossa conanti and B, cuchumatana can be distinguished by dorsal coloration (a broad, pale middorsal longitudinal band from tip of snout onto tail or blotched or spotted versus pale dorsolateral stripes), subcaudal coloration (tan to dark brown with gold or silver flecking

20 30 35 40 45 50 55 60 65 70 75

SL (MM)

10) 30

,
35

,
40

,
45

,
60

,
65

,
70

,
75

FIG.3.-Comparison of standard length (SL) and maxillary tooth number in the Honduran species of the Bolitoglossa d u n n i group. Species codes are solid + diamonds = carri, y = 0 . 8 4 9 ~ 13.9, n = 22; open squares = celaque, y = 0 . 7 4 4 ~ 13.5, n = 51; open + circles = conanti, y = 0 . 8 2 9 ~ 20.9, n = 47; solid + circles = dunni, y = 0 . 7 9 5 ~ 29.1, n = 22. +

50 55 SL (MM)

FIG.4.-Comparison of standard length (SL) and vomerine tooth number in the Honduran species of the Bolitoglossa dunni group. Species coded as in Fig. 3; carri, y = 0 . 1 7 2 ~ 15.1, n = 22; celaque, y + = 0 . 3 5 9 ~ 3.9, n = 51; conanti, y = 0 . 1 7 4 ~ 20.5, + n = 47; dunni, y = 0 . 2 3 9 ~ 18.7, n = 22. +

TABLE 3.-Analysis of covariance for the effect of the four Honduran species of the durzni group and standard length (SL) on head width (in mm). The methods are the same as those used in Table 1 except that only five subadult dt~ntzi ere included. The low w P value for species plus SL is a result of significant interaction between the species carri, celaque, and conanti.
<If
SS \!IS F

TABLE 4.-Analysis of covariance for the effect of the four Honduran species of the dunni group and standard length (SL) on female tail length (in mm). The methods are the same as those used in Table 1 except that only two subadult drinni were included. There was no significant interaction between the species and SL jF = 2.06, P = 0,1221) so the interaction was deleted from the model.
df
SS
MS

SL Species Species + SL Error celaque celaque celaque carri carri conanti

1 3

90.14 1.12 1.75 16.25

90.14 0.37 0.58 0.11


D

754.41 3.13 4.89


CD

0.0001 0.0278 0.0029


P

SL Species Error celaquc celaque celaque carri carri conanti

1 2177.69 2177.69 382.72 0,0001 79.31 13.94 0.0001 3 237.94 5.($9 41 233.29
versus

3 136

CD

versus

carri conanti dunni conanti dunni dztnni

0.04 0.04 2.16 0.01 2.20 2.20

0.17 0.14 0.18 0.18 0.21 0.18

0.6524 0.5507 0.0001 0.9867 0.0001 0.0001

carri conanti dunni conanti dunni dunni

5.46 0.64 9.29 6.10 14.75 8.65

2.37 1.90 2.11 2.15 2.33 1.84

0,0001 0.5016 0.0001 0,0001 0.0001 0,0001

or spotting versus tan or pale orange-yellow with gray mottling), amount of webbing on hindlimbs (1%-2 phalanges between digits three and four free versus Ml ) ,and in TI, (TL/SL % = 91.6% in 36 adults versus 77.3% in four). BolitogEo.ssa conanti differs from B, engelhardti in dorsal coloration (patterned versus usually uniform), size (SL % = 44.4 in 50 adults versus 39.9 in lo), maxillary tooth number in adults (f = 60.5 in 47 versus 49.5 in lo),

and amount of webbing (111'2-2 phalanges of longest toes free versus one). Bolitoglossa conanti and B. helmrichi can be distinguished by subcaudal coloration (tan to dark brown versus orange) and in amount of webbing (195-2 phalanges of longest toes free versus only digital tips). Bolitoglossa conanti differs from B. rostrata in dorsal coloration (a broad, pale longitudinal band from tip of snout onto tail or gold blotching or spotting versus pale dorsolateral stripes or occasionally

35

40

45

50

55 SL (MM)

60

65

70

75

35

40

45

50

55 60 SL (MM)

65

70

75

FIG. 5.-Comparison of standard length (SL) and head width in the Honduran species of the Bolitoglossa dunni group. Species coded as in Fig. 3; carri, y = 0 . 1 1 7 ~ 2.3, n = 22; celaque, y = 0 . 1 2 3 ~ 1.7, + n = 52; conanti; y = 0 . 1 5 6 ~ 0.2, n = 49; dunni, y = 0.156~ 1.0, n = 21.

FIG. 6.-Comparison of standard length (SL) and tail length in females of the Honduran species in the Bolitoglossa dunni group. Species coded as in Fig. 3; carri, y = 0 . 7 0 1 ~ 0.4, n = 7; celaque, y = 0 . 9 7 4 ~ + - 9.3, n = 10; conanti, y = 0 . 7 4 4 ~ 4.8, n = 18; dunni, y = 0 . 9 6 3 ~ 7.4, n = 11. -

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TABLE -Analysis of covariance for the effect of 5 the four Honduran species of the dunni group and standard length (SL) on male tail length (in mm). The methods are the same as those used in Table 1 except that no subadults of dunni with complete tails were available There was no significant interaction between the species and SL (F = 1.49,P = 0 2288) so the interaction was deleted from the model.

SL Species Error

1 3 49

1422.85 1422.85 203.10 0.0001 432.27 144.09 20.57 0.0001 343.29 7.01
versus

CD

celaque celaque celaque carri carri conanti

carri conanti dunni conanti dunni dunni

6.84 5.96 14.57 12.80 21.41 8.61

2.06 1.73 2.48 2.10 2.75 2.51

0.0001 0.0001 0.0001 0.0001 0.0001 0.0001

35

40

45

50

55

60

SL (MM)

FIG.7.-Comparison of standard length and tail length in males of the Honduran species in the Bolitoglossa 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. -

uniform dorsum),subcaudal coloration (tan to dark brown with gold or silver flecking or spotting versus uniform pale beige to dull yellow), size (SL f = 46.7 in 24 adult males and 48.0 in 26 adult females versus 53.2 in four adult'males and 59.3 in 11 adult females), and in amount of webbing (1%-2 phalanges of longest toes free versus >2). Description.--Medium-sized (SL in 24 adult males, 38.3-50.5, f = 46.7; 41 .O-60.7, i= 48.0 in 26 adult females) member of B. dunni group; snout truncate; adult females more robust than relatively slender males; labial protuberances well-developed in both sexes, pronounced in adult males; oval-shaped mental glands present in adult males; head relatively narrow in adults (HW/SL in 24 males, 14.5-17.596, f = 15.7; 14.7-18.1%, 1 = 16.1 in 25 females); distinct groove extending below eye, following the curvature of the eye, not extending onto lip; eyes slightly protuberant, narrowly visible beyond margin of jaw when viewed from below in males, not visible in females; postorbital groove shallow, irregular, extending posteriorly from eye, proceeding sharply ventrally posterior to mandible, extending across throat anterior to gular fold; maxillary teeth abundant (21 adult males, 50-73, ff = 59.9; 26 adult females, 50-74, f = 61.0), extending beyond center of eye, increasing in number with increasing size; vomerine

teeth abundant (21 adult males, 20-32, f = 28.0; 26 adult females, 24-33, ff = 29.5), in long, single, arched series, extending slightly beyond outer edge of internal nares; premaxillary teeth (23 adult males, 27, f = 4.0; 26 adult females, 5-9, i= 6.7) enlarged, piercing lip in adult males, short, located behind lip in females; tail slightly compressed 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/SL in 18, 79.3-91.4%, f = 84.8); tail longer than BL in adults (TL/BL in 16 males, 113.2-147.796, f = 132.5; 98.9-119.09'~~f = 112.5in 18 females); limbs slender, moderately long; limb interval varying from zero to one costal fold in adult males, onehalf to two in adult females; webbing moderate, with one and one-half phalanges on inside and one and one-half to two on outside of third digit on forelimbs free of webbing (one specimen, MVZ 186777, was recorded as having one phalanx free), one and one-half to two phalanges between digits three and four on hindlimbs free of webbing; digits discrete, bearing well-developed subterminal pads; digits on forelimbs in order of decreasing length. 3-42-1, on hindlimbs, 3-4-2-5-1. Coloration in life.-Coloration is very variable in this species. There are two color phases, as well as various stages of inter-

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mediacy. One ~ h a s e consists of a broad, pale longitudinal band from the tip of the snout onto the dorsal surface of the tail (Fig. 1B). The other phase consists of dark brown dorsal surfaces with varying amounts of gold spotting or blotching (Fig. 1C). The subcaudal surface has gold (occasionally silver) flecking or spotting on a tan to dark brown ground color. KU 219841: all dorsal surfaces with broad Orange-Rufous (color 132C) longitudinal band with widely scattered brown spots; sides of head and body Amber (color 36); dorsal surface of legs Amber with OrangeRufous spotting; venter of body brown with gold flecking and spotting; chin yellowishtan; subcaudal surface tan with gold flecking and spotting. KU 219855: dorsum of head, body, and basal section of tail with broad Pratt's Rufous (color 140) longitudinal band with slight brown spotting on head; sides of head and body Amber (color 36); ventral and subcaudal surfaces dark brown with scattered gold flecking; iris Pratt's Rufous. KU 219842: all dorsal surfaces Sepia (color 219) with heavy gold flecking, suggestive of banding; additional large gold spots on dorsal surface of basal section of tail; sides of head and body Sepia with light gold flecking; ventral and subcaudal surfaces dark brown with scattered gold flecking; iris with gold flecking. KU 219870: same as KU 219842 except that ventral and subcaudal surfaces with silver flecking. KU 219844,219856: same as KU 219842 except with considerably less gold flecking on dorsal surfaces. KU 219880: same as KU 219844, 219856 except for large Orange-Rufous (color 132C) blotching on basal section of dorsal surface of tail. KU 219854: all dorsal surfaces Amber (color 36) with gold blotching and spotting; sides of head and body Amber; ventral and subcaudal surfaces brown with gold flecking and spotting; iris gold with black reticulations. KU 219843: all dorsal surfaces Kingfisher Rufous (color 240) with widely scattered gold flecking; OrangeRufous (color 132C) blotching on dorsal surface of basal section of tail; ventral and subcaudal surface brown with scattered gold flecking; iris mottled gold and brown.

KU 219879: all dorsal surfaces with Orange-Rufous (color 132C) longitudinal band with dense gold flecking throughout; sides of head and body Amber (color 36) with heavy gold flecking; ventral and subcaudal surfaces brown with gold flecking and spotting. Measurements of ho1otype.-HW 9.6; head length 14.8; head depth at posterior angle of jaw 4.8; eyelid length 3.5; eyelid width 1.5; anterior rim of orbit to snout 4.0; horizontal orbital diameter 3.1; interorbital distance 3.5; distance between vomerine teeth and parasphenoid tooth patch 0.7; snout to forelimb 17.0; distance separating internal nares 3.2; distance separating external nares 3.5; snout projection beyond mandible 1.5; SL 60.7; BL 45.9; snout to anterior angle of vent 56.0; axilla to groin 32.6; TL 53.3; tail width at base 3.7; tail depth at base 4.1; forelimb length 13.4; hindlimb length 14.3; width of right forefoot 4.5; width of right hindfoot 5.5 Referred specimens.-KU 219870-91, MVZ 186772, juveniles or subadults, all from Quebrada Grande. MVZ 186755, subadult, from Cerro Cusuco. Comments. -Two specimens (MCZ 21245-46) from Portillo Grande. Departamento de Yoro, Honduras (Fig. 2) are tentatively assigned to B. conanti. In preservative, these specimens have dark brown dorsal surfaces with lighter brown spots. MCZ 21246 has 87 maxillary teeth, which is 13 higher than the highest count obtained for conanti, MCZ 21245 has 69 which is near the upper count for conanti. This population may represent a distinct species. Three specimens (KC' 219892, LACM 46940, El Portillo, 1900 m; MVZ 186727, El Chagiiitbn, 1920 m) from the Departamento de Ocotepeque, Honduras and one (MVZ 200535, Hacienda Montecristo, 2250 m) from the Departamento de Santa Ana, El Salvador (Fig. 2) are also tentatively assigned to B. conanti. These specimens either had small light spots on the dorsal surfaces in life (KC' 219892) or lighter dorsal surfaces (at least anteriorly) somewhat suggestive of the longitudinally banded phase of B, conanti (LACM 46940, MVZ 186727,200535). MVZ 186727 (a ju-

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venile) also has a dark X mark on the neck and shoulder region similar to those seen in several young specimens of the banded phase of B, conanti. Mertens (1952:Fig. 30) illustrated a subadult from Hacienda Montecristo with this mark. New material is needed to determine the taxonomic status of this series. The Yoro and Ocotepeque specimens were not used in the analysis of B. conanti. Distribution a n d ecology.-This species is known for certain (see Comments above) from the sierras Espiritu Santo and Omoa in northwestern Honduras (Fig. 2). Bolitoglossa conanti has been collected in the Lower Montane Wet Forest formation (Holdridge 1967) from 1370-1680 m elevation. Most of the specimens that we collected were taken from arboreal bromeliads (Tillandsia sp.), but a few were also taken at night from vegetation alongside a small stream. Schmidt (1942) also reported finding this species in bromeliads. Bolitoglossa conanti occurs sympatrically with B. dunni in the mountains west of San Pedro Sula (Schmidt, 1942), at Quebrada Grande, and at Cerro Cusuco. The species is also sympatric with Nototriton nasalis at the latter locality and B. rufescens at Quebrada Grande. All of these sympatric species also occur in bromeliads at these localities. Etymology.-We take pleasure in naming this species in honor of Roger Conant, near the time he also has been honored for his contributions to herpetology with a symposium by the Society for the Study of Amphibians and Reptiles. With the naming of Bolitoglossa conanti, we also wish to take note of the influence Dr. Conant has had on the field of herpetology. His famous Field Guide, first published in 1958 and now in its third edition, is a model of the artful combination of scholarly detail and popular appeal. His importance in the field of herpetological husbandry is unquestioned, as a result of his career at the Philadelphia Zoo. His scholarly publications have been many and influential. Finally, his latest major work on Agkistrodon and its allies is a testament to his dedication to his field and to the strength of

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

Bolitoglossa carri sp. nov. (Fig. 1D) Ho1otype.-FMNH 236502, adult female, from Cerro Cantagallo, near Lepaterique (14"06'N, 8'i028'W), 1840 m elevation, Departamento de Francisco Morazhn, Honduras, 18 August 1986, James R. McCranie, Kenneth L. Williams, and Larry David Wilson. Original number LDW 8361. Paratypes (21).--FMNH 236470, 236472, 236490, 236500, 236512, K U 219893-94, adult females, FMNH 236452, 236454, 236473-74, 236476-78, 23649193, 236503, KU 219895-97, adult males, all from Cerro Cantagallo, 1840-2070 m. Diagnosis.-Bolitoglossa carri differs from B, dunni in dorsal coloration (pale brownish dorsolateral stripes on a darker brown ground color versus uniform Mahogany Red), subcaudal coloration (melanophores present on a pale yellow to pinkish-cream ground color versus pale Mahogany Red without melanophores), coloration of the dorsal surfaces of feet (absence versus presence of conspicuous gold spots), size (SL T = 42.6 in 14 adult males and 49.9 in eight adult females versus 56.7 in six adult males and 64.6 in 10 adult females), maxillary tooth number plotted against SL (Table 1; Fig. 3), vomerine tooth number plotted against 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 of webbing on hindlimbs (1%-2 phalanges between digits three and four free versus 1-1%). Bolitoglossa carri and B. conanti differ in dorsal coloration (pale dorsolateral stripes beginning behind eye versus a broad, pale longitudinal band from tip of snout onto tail or blotched or spotted with gold), subcaudal coloration (melanophores present on a pale yellow to pinkish-cream ground color versus tan to dark brown with gold or silver flecking), maxillary tooth number

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plotted against SL (Table 1; Fig. 3), vomerine tooth number plotted against SL (Table 2; Fig. 4), and in relative TL in both sexes (Tables 4, 5; Figs. 6, 7). Bolitoglossa carri can be distinguished from 3. cuchunzatana by maxillary tooth number in adults (f = 52.3 in 22 versus 60.2 in 17) and amount of webbing (1%-2 phalanges between digits 3 arid 4 on hindlimhs free versus Vz-1). BoEitoglossa carri and B. engelhardti can be distinguished by dorsal coloration (pale dorsolateral stripes versus usually uniform), size (SL R = 45.2 in 22 adults versus 39.9 in lo), amount of webbing (1-2 phalanges of digit 3 on forelimbs and 1'h-nearly 2 between digits 3 and 4 on hindlimbs free versus 1\, and by T L (TLiSL f = 75 7% in 17 adults versus 0.88 in nine) Bolitoglossa carri and B. helmrichi can be distinguished by subcaudal coloration (pale yellow to pinkish-crearn versus orange), maxillary tooth number in adults (2 = 52 3 in 22 versus 61.9 in 22j. amount of webbing (1-2 ~halanges digit of 3 on forelimbs and 195-2 between digits 3 and 4 on hindlimbs free versus only digital tips), and by TL (TL/SL f = 75.7% in 17 adults versus 86.2% in four) Rolitoglossa carri differs from B. rostrata in maxillary tooth number in adults (R = 52.3 in 22 versus 61 2 in 15),amount of webbing \two or less phalanges of longest toes free versus > 2 ) , size (SL, R = 42.6 in 14 adult males and 49.9 in eight adult females versus 53.2 in four adult males and 59 3 in I 1 adult females), and in T L JTL/SL f = 75.7% in 17 adults versus 87.0% in 13). Description.-Medium-sized (SL in 14 adult males, 37.4-48 0, R = 42.6; 42.4-58.1, R = 49.9 in eight adult females) member of B, dunni group; snout truncate; adult females more robust than relatively slender males; labial protuberances well-developed in both sexes, pronounced in adult males; oval-shaped mental glands present in adult males; head broad in adults (HW/ SL in 14 males, 15.9-17.796,X = 17.1; 15.017 0%, z = 16.3 in eight females); distinct groove extending below eye, following the curvature of the eye, not extending to lip; eyes slightly protuberant, narrowly visible beyond margin of jaw when viewed frorn below in males, not or only barely visible in females; postorbital groove shallo\v, ir-

regular, extending posteriorly from eye, ~roceedingsharply ventrally posterior to mandible, extending across throat anterior to gular fold, some specimens have one or two additional shallow, irregular grooves between anteriormost groove and gular fold; maxillary teeth relatively few in number (14 adult males, 46-60, R = 50.4; eight adult females, 47-68, f = 55.6), extending beyond center of eye, increasing in number with increasing size; vomerine teeth relatively few in number (14 adult males, 18-24, R = 22.1; eight adult females, 22-28, R = 24.4), in long, single, arched series, extending slightly beyond outer edge of internal nares; premaxillary teeth (14 adult males, 2-6, f = 3.4; eight adult females, 4-7, f = 5.4) enlarged, piercing lip in adult males, short, located behind lip in females; tail lightly compressed laterally, constricted basally; tail short, always less than SL (TL/SL in 10 adult males, 73.983.0%, R = 79.2; 6'7.9-74.1%, P = 70.6 in seven adult females), slightly shorter than BL in adult females (TL/BI, in seven, 91.5100.9%.f = 95.1), slightly longer than BL in adult tnales (TL/BL in 10, 100.0115.8%, f = 107.8); limbs slender, moderately long; limb interval varying from 0-1 costal fold in adult males, from onehalf to two and one-half in adult females; webbing moderate, with one to one and one-half phalanges on inside and one and one-half to two on outside of third digit on forelimbs free of webbing, one and onehalf to two phalanges between digits 3 and 4 on hindlimbs free of webbing: digits discrete, bearing well-developed subterminal pads; digits on forelimbs in order of decreasing length. 3-4-2-1, on hindlimbs, 3-49 -51. Coloration in life.-Coloration is somewhat variable in this species. All adult specimens had pale brownish dorsolateral stripes from the shoulder region to the end of the body on a darker brown ground color. The area between the dorsolateral stripes varied in the extent of pale color present. Some specimens had only a narrow, incomplete pale nliddorsal stripe, whereas others had extensive pale areas on the middorsum that were mottled with darker brown. The inner borders of the dorsolateral stripes may be partially bro-

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ken in those specimens with extensive pale middorsal areas. The top of the head was always dark brown without paler markings. The dorsal surface of the tail had varying amounts of pale brounish areas over its entire length. Subcaudal coloration varied from pale yellow with small, scattered melanophores to pinkish-cream with a heavy sprinkling of melanophores with or without scattered rust red mottling. KC 2 19895-96, 2 19898-99: dorsal surfaces of heads, middorsal regions, and lateral areas Raw Umber (color 23); dorsolateral stripes Kingfisher Rufous (color 240); dorsal surfaces of tails Kingfisher Rufous. KU 219893: similar to KU 219895-96, 219898-99, except that dorsolateral stripes and dorsal surface of tails Pratt's Rufous (color 140). KC' 219894: dorsal surfaces of head, narrow middorsal region, and lateral areas Dark Brownish Olive (color 129); diffuse dorsolateral stripes and dorsal surface of tail Raw Sienna (color 136) F3INH 237502: ventral and subcaudal surfaces pale yellow with scattered melanophores, additional scattered rust red mottling present on subcaudal surface. FMNH 23647273: ventral and subcaudal surfaces pinkishcream with heavy sprinkling of melanophores. additional scattered rust red mottling on subcaudal surface. Measurements of ho1otype.-HW 8.7; head length 14.4; head depth at posterior angle of jaw 5.2; eyelid length 3.7; eyelid width 1.9; anterior rim of orbit to snout 4 0; horizontal orbital diameter 3.5; interorbital distance 3.6; distance between vomerine teeth and parasphenoid tooth patch 0.5; snout to forelimb 18.4; distance separating internal nares 2.8; distance separating external nares 3.4; snout projection beyond mandible 1.5; SL 58.1; BL 43.7; snout to anterior angle of vent 53.5; axilla to groin 31.1; TL 40.0; tail width at base 3 2; tail depth at base 4.4; forelimb length 13.4; hindlimb length 14.5; width of right forefoot 4.3; width of right hindfoot 5.6. Referred specimens.-FMNH 23641624, 236427-32, 236434-39, 236453, 236455-58, 236475, 236499, KU 219898901, all juveniles or subadults from Cerro Cantagallo. Distribution a n d ecology.--This species is known only from the type locality

(Fig. 2). Bolitoglossa carri has been collected from the Lower Montane Moist Forest formation (Holdridge, 1967) from 1840-2070 m elevation. The former elevation was the lowest at which we worked in the area and the latter is at the summit of the highest peak in the region. Most specimens were collected inside bromeliads (Tillandsin sp.),but several were found at night sitting on broad leaves growing on shrubs next to a small stream: and two were underneath the bark of a large, fallen tree in a clearing. The bromeliads containing salamanders were usually on trees, but a few salamanders were also found in bromeliads growing on the ground. The salamanders were usually found one to a bromeliad: but in a few instances t\vo occurred in a single bromeliad; in two cases three individuals were found in the same bromeliad. No other species of salamander has been collected on Cerro Cantagallo. Etymology.-The species carri is named in honor of the late Archie F. Carr. who taught for five years at the Escuela Agricola Panamericana in El Zamorano, Honduras, not far from the type locality of the species. During his stay, Dr. Carr amassed an important collection of Honduran amphibians and reptiles which is now housed at the American Museum of Natural History. Dr. Carr was a naturalist and conservationist extraordinaire with an enviable gift with words whose reputation extended far beyond the limits of herpetology. Bolitoglossa celaque sp. nov. (Fig. 1E) Ho1otype.-FMNH 236504, adult female, from near the Rio ArcAqual, eastern side of Cerro Celaque, Cordillera de Celaque (14"32'11:, 88"40tW), 2480 m elevation, Departamento de Lempira, Honduras, 1 August 1985, James R . McCranie, Kenneth L. Williams, and Larry David Wilson. Original number LDW 6577. Paratypes (51).-FMNH 236485, 236488,236506-07,236509, K U 21990203, h4VZ 186728, 186734, 186737, adult females, FMNH 236482, 236487, 236489, 236505,236508,236510-11, MVZ 18672931, 186733, 186735-36, 186738-39, 186741-42, 186745-46, adult males, all

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from the eastern side of Cerro Celaque, 1930-2620 m. K U 219905-06,219910-12, 219914, adult females, KU 219904, 219907-09, 219913, adult males, all from San Pedro, 1990 m, Departamento de Intibucii, Honduras. FMNH 236479,236496, 236513, adult females, FMNH 236480, 236494-95, 236497-98, 236514, K U 219915, MVZ 186751, adult males, all from Zacate Blanco, 2070-2150 m, Departamento de Intibuch, Honduras. Diagnosis.-Bolitoglossa celaque differs from B. d u n n i in dorsal coloration (russet with minute gold flecking versus uniform Mahogany Red), subcaudal coloration (pale yellow to dark orange versus pale Mahogany Red), coloration of the dorsal surfaces of feet (absence versus presence of conspicuous gold spots), size (SL f = 47.7 in 32 adult males and 48.3 in 20 adult females versus 56.7 in six adult males and 64.6 in 10 adult females), maxillary tooth number plotted against SL (Table 1; Fig. 3), vomerine tooth number plotted against 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 of webbing (usually two phalanges of longest toes free versus %-I% phalanges of digit 3 on forelimbs and 1-1% between digits 3 and 4 on hindlimbs). Bolitoglossa celaque and B. conanti differ in dorsal coloration (russet with minute gold flecking versus a broad, pale longitudinal band from tip of snout onto tail or with gold spotting or blotching on a sepia ground color), subcaudal coloration (gray or orange punctations on a pale yellow to dark orange ground color versus gold flecking or spotting on a tan to dark brown ground color), maxillary tooth number plotted against SL (Table 1; Fig. 3), vomerine tooth number plotted against SL (Table 2; Fig. 4), relative TL in males (Table 5; Fig. 7), and in amount of webbing (usually two phalanges of longest toes free versus 1%-2). Bolitoglossa celaque differs from B. carri in dorsal coloration (russet with minute gold flecking versus pale dorsolateral stripes), maxillary tooth number plotted against SL (Table 1; Fig. 3), vomerine tooth number plotted against SL (Table 2; Fig. 4), relative TL in both sexes (Tables 4, 5; Figs. 6,

7 ) ,and in amount of webbing (usually two


phalanges of longest toes free versus 1-2 on digit 3 of forelimbs and 1Yz-2 between digits 3 and 4 on hindlimbs). Bolitoglossa celaque and B. cuchumatana can be distinguished by dorsal coloration (russet with minute gold flecking versus pale dorsolatera1 stripes), maxillary tooth number in adults (f = 49.2 in 51 versus 60.2 in 17), and by amount of webbing (usually two phalanges between digits 3 and 4 on hindlimbs free versus %-1). Bolitoglossa celaque can be distinguished from B. engelhardti by size (SL f = 47.9 in 52 adults versus 39.9 in 10) and amount of webbing (usually two phalanges of longest toes free versus one). Bolitoglossa celaque and B. helmrichi can be distinguished by dorsal coloration (russet with minute gold flecking versus patterned with stripes, spots, or blotches), maxillary tooth number in adults (f = 49.2 in 51 versus 61.9 in 22), and amount of webbing (usually two phalanges of longest toes free versus only digital tips). Bolitoglossa celaque differs from B. rostrata in dorsal coloration (russet with minute gold flecking versus usually with pale dorsolateral stripes), size (SL f = 47.7 in 32 adult males and 48.3 in 20 adult females versus 53.2 in four adult males and 59.3 in 11 adult females), maxillary tooth number in adults (f = 49.2 in 51 versus 61.2 in 13), and amount of webbing (usually two phalanges of longest toes free versus >2). Description.-Medium-sized (SL in 32 adult males, 37.2-55.1, f = 47.7; 36.6-62.2, f = 48.3 in 20 adult females) member of B. d u n n i group; snout truncate; adult females more robust than relatively slender males; labial protuberances well-developed in both sexes, pronounced in adult males; oval-shaped mental glands present in adult males; head relatively narrow in adults (HW/SL in 32 males, 14.7-17.2%, f = 15.9; 14.4-18.0%, f = 16.0 in 20 adult females); distinct groove extending below eye, following the curvature of the eye, not extending to lip; eyes slightly protuberant, narrowly visible beyond margin of jaw when viewed from below in males, not or only barely visible in females; postorbital groove shallow, irregular, extending

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posteriorly from eye, proceeding sharply ventrally posterior to mandible, extending across throat anterior to gular fold; maxillary teeth relatively few in number (32 adult males, 40-58, f = 50.1; 19 adult females, 25-62, f = 47.8), extending beyond center of eye, increasing in number with increasing size; vomerine teeth relatively few in number (32 adult males, 15-26, f = 20.8; 19 adult females, 14-30, f = 21.6), in long, single, arched series, extending slightly beyond medial border of internal nares; 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; tail slightly compressed laterally, constricted basally; tail relatively long, always less than SL (TL/SL in 20 adult males, 70.4-98.792, f = 85.4; 62.9-86.892, f = 79.0 in 10 adult females), usually more than BL (TL/BL in 20 adult males, 97.0-132.8%, f = 115.1; 83.1-112.4%, f = 103.7 in 10 adult females); limbs slender, moderately long; limb interval varying from zero to one costal fold in males, zero to two and onehalf in females; webbing slightly reduced, with one and one-half to two (most often two) phalanges on both sides of digit 3 on forelimbs free of webbing, one and onehalf to little over two (most often two) phalanges between digits 3 and 4 on hindlimbs free of webbing; digits discrete, bearing well-developed subterminal pads, digits on forelimbs in order of decreasing length, 3-4-2-1, on hindlimb, 3-4-2-5-1. Coloration in life.-Only minor variation in dorsal coloration, consisting of amount and color of flecking, has been recorded in this species. The subcaudal coloration varied from pale yellow to dark orange. KU 219902: all dorsal surfaces Russet (color 34), with minute gold flecks; dorsal surface of tail with heavier gold flecking; lateral portion of body Russet, with copper flecking; ventral and subcaudal surfaces pale yellow, with ocher blotching and scattered minute gold flecking; iris irridescent copper. KU 219903: similar to KU 219902, but with less gold and copper flecking. KU 219915 and FMNH 236494: dorsal surfaces Russet (color 34) with minute gold flecks scattered

over bodies, feet, and basal section of tails; ventral surfaces of heads and bodies pale orange; subcaudal surfaces pale yellow with gray and orange punctations. KU 219904: similar to KU 219915 and FMNH 236494, except that gold flecking especially heavy on eyelids and snout; iris also heavily flecked with gold. FMNH 236480: similar to KU 219915 and FMNH 236494, except that ventral surfaces of head and body pale yellow and subcaudal surface pale orange. FMNH 236459: all dorsal surfaces Russet (color 34), with minute chocolate brown flecking dorsally, silver flecking laterally; subcaudal surface dark orange, with gray and silver flecking. The minute gold flecking on the dorsum quickly disappears in preservative. All specimens have uniform dorsal surfaces in preservative except FMNH 236488 which has incomplete, pale dorsolateral stripes. Measurements of ho1otype.-HW 9.5; head length 13.8; head depth at posterior angle of jaw 5.8; eyelid length 4.2; eyelid width 2.1; anterior rim of orbit to snout 4.3; horizontal orbital diameter 3.9; interorbital distance 3.5; distance between vomerine teeth and parasphenoid tooth patch 0.5; snout to forelimb 17.4; distance separating internal nares 2.6; distance separating external nares 3.3; snout projection beyond mandible 1.5; SL 62.2; BL 48.4; snout to anterior angle of vent 58.7; axilla to groin 34.0; TL 50.9; tail width at base 4.6; tail depth at base 5.1; forelimb length 16.0; hindlimb length 17.0; width of right forefoot 6.3; width of right hindfoot 8.1. Referred specimens.-FMNH 23642526, 236440-51, 236469, 236471, 236481, 236483-84,236486,236501, MVZ 186732, 186740, 186743-44, 186747-50, juveniles or subadults from Cerro Celaque. FMNH 236459-67, KU 219916 juveniles or subadults from Zacate Blanco. FMNH 236433, 236468, MVZ 186752-54, juveniles or subadults from the Sierra de Montecillos south of Tutule, 2100-2160 m, Departamento de La 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, 39770-71,39774, adult males, MVZ 39737, 39740-41,39746-47,39749-51,39753-58,

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39760,39763-64,39769,39773,39775, juveniles or subadults, all from Los Esesmiles, 2440-2720 m , Departamento de Chalatenango, El Salvador (see Comment be lo^ ). Comment.-Wake and Lynch (1976) assigned all El Salvadorean salamanders of the dunni group to B , dunni. The series from Los Esesmiles (Fig. 2) that we examined are similar to B. celaque in size, in being unpatterned in preserkatibe, and in maxillary and vomerine tooth numbers. Howeker, they differ from celaque in having less webbing (over two phalanges of longest toes free versus ~lsuallytwo) and having broader heads (pairu ise ANCOYJ\, F = 9 47, P = 0.0030).These specimens are tentative]!, assigned to B , celaque, but were not used in the analysis of that species. Distribution and ecology.-This species is known in southwestern Honduras from four isolated mountain ranges in the cordilleras Celaque, Opalaca, and klontecillos (Fig. 2). Bolitoglossa celaque has been collected from the Lower Montane Moist Forest formation (Holdridge, 1967) from 1930-2620 m elevation. The species is both arboreal and terrestrial. -4t Cerro Celaque, specimens were collected b j day under moss mats both on the ground and on tree trunks and in both terrestrial and arboreal bromeliads (the latter Tillandsiu sp , the former an unidentified spiny species) -4single specimen was collected on Cerro Celaque as it walked across the wet surface of leaf litter in the early afternoon. At night, B. celaque was commonly seen on leaves and rocks alongside streams on Cerro Celaque All specimens, except for one that was under a log, from the other three localities for this species were taken by day from arboreal bromeliads (Tillandsia sp ). No other species of salamander has been found with B. celaque. Etymology.-The name celaque is used as a noun in apposition. It refers to the name of the highest mountain in Honduras, Cerro Celaque, which is also the type locality of the species The mountain's name, according to Cruz (1986),is derived from the Aztec word celac or ceelac, meaning "in the cold waters". The name

may be in reference to the frigid Rio Arciqual, which cascades off the eastern face of Cerro Celaque. Elias (1984) made an important contribution to our knowledge of the systematics of the Middle American members of the Bolitoglossa beta group when he recognized that two of the species groups (the helmrichi and rostrata groups) of these salamanders as recognized by Wake and Lynch (1976) were actually ". . . large, heterogeneous and united by no specific characters" (p. 14). He then subdivided and reorganized these t\vo groups into vrhat he felt were ". . . tight clusters of phenotypically similar species" (Elias, 1984:14). One such assemblage (with five described species, plus one undescribed Guatemalan species) was the dunni group ". . . characterized by blunt rounded toe tips, fully developed subdigital pads, and a dark brown ground color, frequently marked with a lighter brown dorsal swath or paired shoulder stripes . . ." (Elias, 1984:16).Larson (1983a), using protein comparisons, concluded that phylogenetic inferences were compatible with the species groups recognized by Elias (1984) and stated that Elias' dunni group ". . . appear[s] to form a monophyletic lineage whose primary dichotomy is approximately 12 million years old" (Larson, 1983a:97). The three new species described in this paper clearly are members of the dunni group as defined by Elias (1984).However, because of the uncertain taxonomic status of the Honduran and El Salvadorean populations commented upon above. we feel that any discussion about the relationships of B. carri, celaque, conanti, and dunni within the dunni group would be premature at this time. In addition, a biochemical analysis of carri and celaque is needed before the genetic distances of these species from conanti and dunni can be determined.
Acknowledgments.-Valuable field assistance \+,as provided by G. A. Cruz, M. R. Espinal, K. Hogan, L. Porras, J. C. Rindfleish, and K. L. Williams. Cruz also donated his Cerro Celaque collection of salamanders, for which we are grateful. We also want to thank U'.

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Aguilar, G. A. Cruz, and S. Midence for their help in securing collecting permits from the Departamento de Recursos Naturales, Tegucigalpa. We thank the following people and their museums for the loan of pertinent material; Museum of Comparative Zoology, Harvard University (MCZ)-J. P. Rosado and P. Alberch; Museum of Vertebrate Zoology, University of California, Berkeley (MVZ)-D. Wake, B. Stein, and K. Autumn; Natural History Museum of Los Angeles County (LACM)-J. W. Wright; University of Michigan, Museum of Zoology (UMMZ)-A. G. Kluge and G. Schneider.

CRCZ,G. A. 1986. Guia de 10s parques nacionales, refugios de vida silvestre, reservas biologicas y monumentos naturales de Honduras. Asociaci6n Hondurefia de Ecologia. Tegucigalpa. D u x s , E. R. 1926. The Salamanders of the Family Plethodontidae. Smith College, Northampton, h.iassachusetts. ELIAS, P. 1983. Salamanders of the northwestern highlands of Guatemala. Nat. Hist. Mus. Los Angeles. Co. Contrib. Sci. 348:l-20. HOLDRIDGE, R. 1967. Life Zone Ecology. ReL. vised edition. Tropical Science Center, San Josi., Costa Rica. L.\RSON, 1983a. A molecular phylogenetic per'4. spective on the origins of a lowland tropical salamander fauna. I. Phylogenetic inferences from protein comparisons. Herpetologica 39:85-99. . 1983b. A molecular phylogenetic perspective on the origins of a lowland tropical salamander

fauna. 11. Patterns of morphological evolution. Evolution 37:1141-1153. MERTENS, 1952. Die amphibien und reptilien R. von El Salvador, auf grund der reisen von R. Mertens und A. Zilch. Abh. Senckenberg, h'atuf. Ges. 487:1-120. SCHMIDT, . P. 1933. New reptiles and amphibians K from Honduras. Zool. Ser. Field hlus. Nat. Hist. 20: 15-22. . 1936. Guatemalan salamanders of the genus Oedipus. Zool. Ser. Field Mus. Nat. Hist. 20: 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. L. STUART, C. 1943. Taxonomic and geographic comments on Guatemalan salamanders of the genus Oedipus. Misc. Publ. Mus. Zool. Univ. Michigan 56:l-33. WAKE, B., AND A. H. BR.\ME, R . 1969. SystemD. J atics and evolution of neotropical salamanders of the Bolitoglossa helmrichi group. Contrib. Sci. Los .4ngeles Co. Mus. Nat. Hist. 175:l-40. W.AKE, B., A N D J. F. LYSCH. 1976. The distriD. bution, ecology, and evolutionary history of plethodontid salamanders in tropical America. Nat. Hist. Mus. Los Angeles Co. Sci. Bull. 25:l-65.
Accepted: 30 March 1992 Associate 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 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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Literature Cited
A Molecular Phylogenetic Perspective on the Origins of a Lowland Tropical Salamander Fauna. II. Patterns of Morphological Evolution Allan Larson Evolution, Vol. 37, No. 6. (Nov., 1983), pp. 1141-1153.
Stable URL:
http://links.jstor.org/sici?sici=0014-3820%28198311%2937%3A6%3C1141%3AAMPPOT%3E2.0.CO%3B2-9