The Frog Genus Atelopus in Ecuador

(Anura: Bufonidae)

JAMES A. PETERS

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 145
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 145

The Frog Genus Atelopus in Ecuador

(Anura: Bufonidae)

James A. Peters

SMITHSONIAN INSTITUTION PRESS

City of Washington

1973
ABSTRACT

Peters, James A. The Frog Genus Atelopus in Ecuador. Smithsonian Contributions to Zoology, number 145, 49 pages, 31 figures, 1973.—The genus

Atelopus is represented in Ecuador by 15 species, of which 6 are here described

as new taxa. A key for identification, descriptions, and figures are presented for

all species, and a detailed analysis of variation in proportions attempts to

quantify certain subjective characteristics used by earlier frog systematists.

The genus includes two nonphylogenetic groups of species, the first, with

slender bodies and long limbs, usually found in the lowlands and lower mountain slopes, and the second, with stocky bodies and short limbs, usually found

at higher altitudes in the Andes. The species appear to be highly responsive to

environmental changes, and successful invasions of the mountains by the lowland species are accompanied by differentiation, evolution, and speciation.

Invasions are likely to be along major river drainages. The invaders are effectively isolated from adjacent drainages by factors such as the highly dissected

terrain, rain shadow, barren areas, and so on. High altitude species are restricted

in lateral distribution, but often have considerable altitudinal range, while low

altitude species have wide latitudinal range, but are restricted altitudinally.

The mountains in Ecuador are "continental islands," showing diversity and

adaptive radiation similar to that of the Hawaiian Islands, with the major difference that the Andes are not dependent upon "waifs" or "drifters" for invaders,

but are constantly supplied by the abundant life of the surrounding tropical

forests.

OFFICIAL PUBLICATION DATF is handstampcd in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 4775. SERIES COVER

DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Peters,' James Arthur, 1922-1972

The frog genus Atelopus in Ecuador.

(Smithsonian contributions to zoology, no. 145)

Bibliography: p.

1. Atelopus. 2. Frogs—Ecuador. I. Title. II. Series: Smithsonian Institution. Smithsonian

contributions to zoology, no. H.">.

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Contents

Page

Introduction 1

Acknowledgments 1

Species Groups within the Genus 2

Comparative Measurements 2

Ratio 3

Knee to Knee/Standard Distance 3

Tibiofibula/Standard Distance 4

Head Length/Standard Distance 5

Head Width/Head Length 6

Tibiofibula/Head Length 7

Tibiofibula/Knee to Knee 7

Sexual Dimorphism 8

Key to the Atelopus of Ecuador 9

Atelopus arthuri, new species 10

Atelopus balios, new species . 12

Atelopus bomolochos, new species 14

Atelopus boulengeri Peracca 16

Atelopus bufoniformis Peracca 18

Atelopus elegans (Boulenger) 20

Atelopus halihelos, new species 22

Atelopus ignescens (Cornalia) 24

Atelopus longirostris Cope 27

Atelopus mindoensis, new species 30

Atelopus nepiozomus, new species 32

Atelopus pachydermus (Schmidt) 34

Atelopus palmatus Andersson 38

Atelopus planispinus Espada 39

Atelopus pulcher pulcher (Boulenger) 41

Present Position of Taxa Formerly Assigned to Atelopus in Ecuador 43

Incertae Sedis . 43

Atelopus longirostris marmorata Werner 43

Zoogeography and Evolution 44

Literature Cited 48

i n

The Frog Genus Atelopus in Ecuador

(Anura: Bufonidae)

James A. Peters

Introduction

It has been my good fortune to be able to spend

considerable time in the field in Ecuador, where I

collected representatives of most of the species of

Atelopus now known from that country. There

are three new species in the material I obtained,

and three more new species have been identified in

the collections of other museums. In addition, the

taxonomic position of several of the previously

described forms has been revised as a consequence

both of the new material available and of reexamination of the type specimens or series. The preparation of this paper has been stimulated by the

recent work on the genus Atelopus by Juan Rivero.

My collections strongly supplement his material,

and I have found his discussions of the Ecuadorian

species cogent and stimulating, even with regard

to those points on which we are not in agreement.

The synonymies presented for each species are

limited to only those references pertaining to Ecuadorian records and specimens, except where a

change in status of the taxon is involved. I have

made no attempt to document use of a taxonomic

name in an inclusive manner.

ACKNOWLEDGMENTS.—My field work in Ecuador

was supported in 1954 by the American Philosophical Society and Brown University, in 1962 by Grant

No. G21010 of the National Science Foundation,

and in 1965-1966 by the Smithsonian Institution.

I am indebted to each for this aid. Many people

James A. Peters, Department of Vertebrate Zoology, National

Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. {Deceased 18 December 1972.)

have helped me to amass the material on which

this study is primarily based, and I wish particularly to thank Manuel Olalla, Peter Spoecker, Robert Mullen, and Gonzalo Herrera, who have helped

in the field. Gustavo OrceW. continues as an

invaluable friend and critic, and his collections,

now at the National Museum of Natural History,

Smithsonian Institution, were extremely useful in

this study. The curators at the following institutions have loaned specimens to me and have provided me with space to work during visits to their

institutions, and I appreciate both kindnesses (abbreviations used in the text follow the name):

American Museum of Natural History (AMNH);

British Museum (Natural History) (BMNH);

California Academy of Sciences (CAS, CAS-SU

refers to numbers assigned to specimens from the

collections of Stanford University, now housed at

the Academy); Field Museum of Natural History

(FMNH); Museo ed Instituto di Zoologia Systematica, Universita di Torino, Italy (UT); Museo Nacional de Ciencias Naturales, Madrid, Spain (MM);

Museum of Comparative Zoology, Harvard University (MCZ); Museum National d'Histoire Naturelle, Paris, France (PM); University of Kansas

Museum of Natural History (UKMNH); University of Michigan Museum of Zoology (UMMZ);

and University of Puerto Rico, Mayaguez

(UPR-M). The abbreviation used for the collections in the National Museum of Natural History

is USNM (United States National Museum). Uncataloged specimens from the James A. Peters

(JAP) and the Gustavo OrceW. (GOV) collections

are deposited at the National Museum of Natural

History.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

SPECIES GROUPS WITHIN THE GENUS.—Although

one should not attempt the definition of species

groups unless his study encompasses the entire

range of variation in the genus, it has been most

convenient for me to recognize that there are two

different groups of species in the Ecuadorian Atelopus. I am not here according these groups any

kind of taxonomic status, and I regard them as

phenetic rather than phylogenetic, so I would not

like to have any taxonomic or phylogenetic interpretation placed on my treatment of them. But

the groups do provide an evident basis for discussion and organization of data, as well as a primary

step in definition of the species and in the preparation of a key for identification.

The longirostris group includes the slim-bodied,

long-legged taxa, while the ignescens group includes the taxa with short, heavy-set bodies and

short limbs. Brighter colors and flash patterns tend

to occur more commonly in the longirostris group,

while the members of the ignescens group tend

to have darker, more sombre colors. Coloration is

not invariable, however, and is not a good basis

for group recognition. The longirostris group is

found at lower altitudes on both sides of the

Andes in Ecuador, while most of the taxa in the

ignescens group are high altitude forms.

The species in the ignescens group include

nrthuri, bomolochos, bufoniformis, halihelos, ignescens, nepiozomus, and pachydermus. The members of the longirostris group are balios, elegans,

longirostris, palmatus, planispinus, and pulcher.

The largest species of all, bonlengeri, and one of

the smallest, mindoensis, are intermediate between

the two groups in many respects, and are difficult

to assign in any unequivocal way. This intermediacy is referred to repeatedly in the subsequent

sections, and reinforces my statement above that

these are not taxonomic groupings, but simply

represent convenient groupings. The species are

arranged on the graphs in the following sections

by groups, with the two "intermediate" species between them.

While groups were not emphasized as such by

McDiarmid (1971) in his recent review of the

genus, and he repeatedly pointed out the uniformity of characters within Atelopus in contrast

to its differences when compared with other genera,

lie did recognize the existence of the two groups

discussed here. He referred in several places to

the long-legged and the short-legged species, and

pointed out several anatomical characteristics

which differentiate the two groups. Unfortunately,

his samples were not large enough to permit full

review of all the Ecuadorian species. Where he

had material it was usually only a single specimen,

and he had no material at all for several of the

taxa.

COMPARATIVE MEASUREMENTS.—All measurements used in this study were made with a Helios

vernier caliper, measuring to the nearest tenth of

a millimeter. The "standard distance" is the

straight-line distance between the tip of the snout

and the posterior end of the coccyx. Although

many previous authors have defined the body

length as from the snout to the anus, preservation

techniques have an effect on the position of the

anus, and the tissues in that region can be easily

compressed, so the measurement is more variable

and not as accurate or consistent as the one defined

here. All other measurements are taken from a

bony surface to another bony surface, insofar as

possible. The measurement from knee to knee is

made with the femora at right angles to the body,

forming a straight line. The calipers are set solidly

against the distal ends of the femora to record the

measurement. The tibiofibula measurement is

from end to end of the bone. The head width is

taken at the widest point of the head, with the

points of the caliper set to pass snugly over the

head. The head length is from the snout tip to

the posteriormost margin of the skull at the dorsal

midline.

As mentioned elsewhere, the two species groups

are clearly distinctive in their general appearance.

The differences are well reflected by calculations of

proportional values, using the various measurements discussed above. In some proportions there

is a considerable sexual dimorphism, and the sexes

are separated in all calculations, even where the

differences were not significant.

The figures show the mean as a vertical line, the

total range of variation as the largest rectangle, the

mean it twice the standard error of the mean as

the smallest rectangle, and the mean ± one standard deviation as the third rectangle. Only the

mean and range are given for very small samples

(fewer than 6). The colors of the rectangles are

reversed from one sex to the other to permit rapid

recognition. The value found as a proportion is
NUMBER 145

always plotted in these figures as a percentage, and

it should be clear that all references to ratios, proportions, and percentages are to these figures.

All references in this paper to significant differences between the means of any two samples is

based on non-overlap between the rectangles representing the mean ± twice its standard error.

This spread of values includes slightly more than

95 percent of the possible values of the mean

itself, and nonoverlap between the two populations

indicates a very high likelihood that the two samples as drawn are different.

Ratio

KNEE TO KNEE/STANDARD DISTANCE.—In this ratio, seen in Figures 1 and 2, higher values indicate

a longer leg in proportion to body length, or standard distance. Although the two species groups are

quite distinct from each other both in body shape

and leg length, as pointed out elsewhere, this is

not strongly reflected by proportional differences.

This is because both body and legs are elongated

in the longirostris group, while in the ignescens

group both body and legs are short and stubby.

IGNESCENS

BOMOLOCHOS

ARTHURI

HALIHELOS

PACHYDERMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PALMATUS

LONGIROSTRIS

ELEGANS

75 80 85 90 95 100 105

FIGURE 1 Species comparison, males, for the ratio knee to knee/standard distance x 100.

Numbers in this and Figure 2 refer to sample size, which are same for other figures.

BUFONIFORMIS

IGNESCENS

BOMOLOCHOS

ARTHURI

PACHYDERMUS

NEPIOZOMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PALMATUS

PLANISPINUS

LONGIROSTRIS

ELEGANS

65 70 75 80 85 90 95 100 105

FIGURE 2. Species comparison, females, for the ratio knee to knee/standard distance x 100.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Even so, the longirostris group is characterized by

values greater than 89 for the proportion, while

individual proportions in the ignescens group are

almost always below 90. A. pachydermus males,

of the ignescens group, range up to values of 94,

and A. balios, of the longirostris group, has values

for most of the females and many of the males below a ratio of 90. A. mindoensis and A. boulengeri

are fairly typical of the ignescens group, with only

a few males falling above a value of 90.

Sexual dimorphism is fairly strong in this characteristic. There is a significant difference between

the means for the sexes in every species except A.

elegans, when samples are large enough for comparison, although there is considerable overlap in

the range of variation of the sexes. The males have

longer legs in proportion to the body length than

the females in every species.

TIBIOFIBULA/STANDARD DISTANCE.—This character is similar to the previous one (KK/SD) in that

it is an expression of a primary difference between

the two groups of species, but it expresses the contrast much more strongly (Figures 3 and 4). Although the body and femora have changed in such

a fashion that the proportion between them is similar in the two groups, the region of the tibiofibula

is much more elongated, proportionately, than the

femoral region in the longirostris group, and this

has been reflected in this ratio.

Individuals in the ignescens group are characterized by values below 45 for this ratio, while

IGNESCENS

B0M0L0CH0S

ARTHURI

HALIHELOS

PACHYDERMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PALMATUS

LONGIROSTRIS

ELEGANS

— • • ••• --•• • • » — • —i » > — i i • • • i i i i i \ •

35 4 0 45 5 0 55

FICURE 3.—Species comparison, males, for the ratio tibiofibula/standard distance x 100.

BUFONIFORMIS

IGNESCENS

B0M0L0CH0S

ARTHURI

PACHYDERMUS

NEPIOZOMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PLANISPINUS

PALMATUS

LONGIROSTRIS

ELEGANS

30 35 40 45 50

FIGUKE 4.—Species comparison, females, for the ratio tibiofibula/standard distance x 100-

55
NUMBER 145

those in the longirostris group almost invariably

have values above 45. In the latter group only

males and females of A. pulcher give lower figures,

while the remaining species have practically no

individuals with values below 46. In the ignescens

group males of A. pachydermus and A. halihelos

include individuals with values above 45, none of

which reach 46, and females of A. nepiozomus may

have values up to 47. No males of A. nepiozomus

have yet been taken, but it might be predicted

that they would give higher values, as well.

The two species A. mindoensis and A. boulengeri

are completely intermediate in this character. Most

of the females of both species lie below 46, most

of the males of both fall above 45. Within the

sexes, the mean for each of these species is significantly higher than that of all others within the

ignescens group, except for females of A. nepiozomus, but the same is true of a comparison with the

members of the longirostris group, where the values

for these two species are significantly lower.

Sexual dimorphism is apparent in all species of

the ignescens group for this character, with the

males having means significantly higher than the

females. The opposite is true of the longirostris

group, for in no case are the means of the two

sexes significantly different. In this regard, A. mindoensis and A. boulengeri show similarities to the

ignescens group, since the means for the sexes are

significantly different in both species.

HEAD LENGTH/STANDARD DISTANCE.—This ratio

(Figures 5 and 6) produces differences between

species within the two major groups, rather than

IGNESCENS

BOMOLOCHOS

ARTHURI

HALIHELOS

PACHYDERMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PALMATUS

LONGIROSTRIS

ELEGANS

26 28 30 32 34 36 38 40

FIGURE 5 Species comparison, males, for the ratio head

length/standard distance x 100.

BUFONIFORMIS

IGNESCENS

ARTHURI

BOMOLOCHOS

PACHYDERMUS

NEPIOZOMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PALMATUS

PLANISPINUS

LONGIROSTRIS

ELEGANS

26 28 30 32 34 36 38 40

FIGURE 6 Species comparison, females, for the ratio head length/standard

distance X 100-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

following group lines, and thus provides a basis

for separation of species within groups. Thus, all

individuals of A. balios and A. elegans, both of the

longirostris group, fall below a proportional value

of 33, as do the following species from the ignescens group: all females of A. bomolochos and A.

ignescens and all males of boulengeri. In the

longirostris group this is also true of most individuals of both sexes of A. pulcher as well as females

of A. planispinis and A. longirostris, and in the

ignescens group, most of the males of A. ignescens

as well as the females of A. arthuri, A. boulengeri,

and A. mindoensis. On the other hand, values of

greater than 33 are found, in the ignescens group,

in all A. bufoniformis as well as the males of A.

arthuri, A. mindoensis, A. halihelos, and A. bomolochos, while, in the longirostris group, values

above 33 are characteristic of both sexes of A. palmatus, and the females of A. longirostris. A pachydermus, of the ignescens group, is completely

intermediate in this character, with as many individuals in both sexes with values above 33 as

those below. The mean for females in this species

is 31.7, for males it is 33.2.

It is apparent from the figures above that sexual

dimorphism occurs in some of the species, but

again it is a mixture between groups. In the ignescens group significant differences exist between the

means for the sexes in A. bomolochos, A. arthuri,

and A. mindoensis, while there are no sexual differences within A. boulengeri or A. pachydermus,

and the means for the sexes in A. ignescens are

very narrowly different at the 95 percent level. In

the longirostris group, significant sexual differences

in this character exist in A. longirostris at a high

level of significance, and in A. pulcher with only

marginal significance, but not at all in A. elegans.

It appears likely that sexual dimorphism exists in

A. balios, and perhaps not in A. palmatus, but the

samples are insufficient to be certain of either.

HEAD WIDTH/HEAD LENGTH.—As seen in Figures

7 and 8, the members of the ignescens group are

characterized by a broad head in relation to its

length (measured from the tip of the snout to the

posteriormost point on the dorsal midline). This

is most exaggerated in A. ignescens itself. The

distinction between the two groups is not sharp,

and several species show an intermediate condition.

Within the ignescens group, A. bufoniformis, A.

ignescens, A. arthuri, A. pachydermus, and A. boulengeri are quite distinct, with all the individuals

showing a ratio of 95 or more. Also, within the

longirostris group, A. planispinus, A palmatus, and

A. longirostris are equally distinct, with ratios generally below 92. Three species in the ignescens

group also show such low values, however, including A. bomolochos, A. halihelos, and A. nepiozomus, with the means for the last two significantly

different from all other species in the group, and

with the latter two having mean values below 90.

IGNESCENS

BOMOLOCHOS

ARTHURI

HALIHELOS

PACHYDERMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PALMATUS

LONGIROSTRIS

ELEGANS

115

FIGURE 7.—Species comparison, males, for the ratio head width/head length x 100.
NUMBER 145

BUFONIFORMIS

IGNESCENS

BOMOLOCHOS

ARTHURI

PACHYDERMUS

NEPIOZOMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PLANISPINUS

PALMATUS

LONGIROSTRIS

ELEGANS

70 80 90

FIGURE 8.—Species comparison, females, for the ratio head

width/head length x 100.

In the longirostris group, A. pulcher, A. balios,

and A. elegans all have higher ratios, indicating

proportionately broader heads.

There are no species with differences between

the means for the sexes that are statistically significant, i.e., no sexual dimorphism exists in this

character.

TIBIOFIBULA/HEAD LENGTH.—This ratio, as

shown in Figures 9 and 10, demonstrates the dichotomy of the two groups, and the intermediacy of

A. mindoensis. Members of the ignescens group

generally have proportional values below 135, and

IGNESCENS

BOMOLOCHOS

ARTHURI

HALIHELOS

PACHYDERMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PALMATUS

LONGIROSTRIS

ELEGANS

100 120 14ifc0 160

FIGURE 9 Species comparison, males, for the ratio tibio

fibula/head length x 100.

BUFONIFORMIS

IGNESCENS

BOMOLOCHOS

ARTHURI

PACHYDERMUS

NEPIOZOMUS

BOULENGERI

MINDOENSIS

PULCHER

BALIOS

PLANISPINUS

PALMATUS

LONGIROSTRIS

ELEGANS

80 100 120 140 160 180

FIGURE 10.—Species comparison, females, for the ratio tibiofibula/head length x 100.

those of the longirostris group are generally above

135. A. bufoniformis is very distinct, overlapping

only one other species in its total range of variation.

In those species for which there are sufficient

individuals of both sexes for comparison, there is

no sexual dimorphism in the ignescens group, with

the exception of A. mindoensis. In this regard A.

mindoensis is more like the species of the longirostris group, where a significant difference between

the means for the two sexes occurs in all species

except A. elegans.

TIBIOFIBULA/KNEE TO KNEE.—This ratio (Figures 11 and 12) is an attempt to express quantitatively a relationship often used in the earlier literaELEGANS

LONGIROSTRIS

PALMATUS

BALIOS

PULCHER

MINDOENSIS

BOULENGERI

PACHYOERMUS

HALIHELOS

ARTHURI

BOMOLOCHOS

IGNESCENS

44 46 48 50 52 54 56

FIGURE 11 Species comparison, males, for the ratio tibiofibula/knee to knee x 100.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ELEGANS

LONGIROSTRIS

PALMATUS

PLANISPINUS

BALIOS

PULCHER

MINDOENSIS

BOULENGERI

NEPIOZOMUS

PACHYDERMUS

BOMOLOCHOS

ARTHURI

IGNESCENS

BUFONIFORMIS

42 4 4 4 6 48 50 52 54 56 58

FIGURE 12. Species comparison, females, for the ratio tibiofibula/knee to knee

X 100.

ture on anuran taxonomy. The lower limb is

placed parallel to the femur, which is extended at

right angles to the body, and the relationship of

the heels in this position is usually stated as overlapping, touching, or separated. Quantitatively, if

the heels overlap, the value of this ratio is usually

greater than 51, while if the heels are separated, it

is smaller than about 50. Most of the members of

the ignescens group give values below 50, while the

species in the longirostris group usually show ratios above 51. A. bufoniformis shows the lowest

values of the genus in Ecuador. A. mindoensis diverges sharply from the ignescens group in this

characteristic, with all the males having ratios

higher than 51, and only 2 of the 16 females with

ratios below 51. A. boulengeri is intermediate, with

values ranging from 48 to 52 in both sexes, although in every case in this species the heels do

not overlap, as a consequence of the method of

measurement.

Although the two principal groups are distinguishable, and the means for this character are

significantly different between species of the groups,

the ranges of variation often overlap extensively,

and this is not a good character for population and

species differentiation. It appears that the proportional relationships between these two parts of the

hind leg remain rather constant, even though in

the ignescens group the leg is short, stubby, and

muscular, while in the longirostris group the leg

is long, lean, and slender.

Sexual Dimorphism

In addition to the several characteristics discussed under the heading of comparative measurements, there are other cases of distinctive sexual

dimorphism in Ecuadorian Atelopus. The most

conspicuous sexual difference is associated with the

act of clasping, which in this genus is a matter

of some magnitude, in light of the length of time

that the clasp is maintained. Males of most species

have a cornified edge on the first digit of the forefoot, and in several species this cornified area is

expanded well onto the second digit and adjacent

areas of the palmar surface. This horny area permits external sex determination in almost every

case, since it is never present in females. The

amount of cornified tissue is considerably greater

than in most anuran species, and is presumably

very useful to the male in the maintenance of his

tenacious grip on the female while awaiting the

deposition of the eggs. Since amplexing pairs may

remain that way for as long as two weeks (see "Remarks" under A. ignescens), a good grip is rather

important.

Rivero (1963:109) has remarked, "Spiny or tubercular skin is apparently not a sexual character
NUMBER 145

in Atelopus, although in tubercular species, the

male is usually more distinctly so than the female." This has not been true in the species I

have examined, for all species with tubercles show

considerable sexual dimorphism in the number,

distribution, and prominence of warts, tubercles,

and particularly in spiny points, single or several,

on the tops of the warts or glandular areas. In

each case it is the female that is more heavily warty

or densely spinose, quite the reverse of the situation

in related amphibians, including other genera in

the Bufonidae. In some Ecuadorian species, the

males are almost smooth on the sides of the body

and the temporal region, while the females have

prominent spines in the same area.

Key to the Atelopus of Ecuador

1. Legs long, thin; body usually slender; heels overlap or touch when tibiofibulae are parallel

to femora at right angles to body; ratio tibiofibula/SD usually more than 45 2

Legs short, stout; body stout; heels do not touch when tibiofibulae are parallel to femora

at right angles to body; ratio tibiofibula/SD usually below 45 9

2. First finger almost entirely hidden in webbing; tip may be free 3

First finger distinct; webbing somewhat incised between first and second fingers 6

3. Belly unicolor, without spotting 4

Belly with clear-cut, marked, round black spots pulcher

4. Dorsum whitish with irregularly parallel, broad, black or brown stripes elegans

Dorsum not colored as above 5

5. Dorsum with broad band of light yellowish brown, filled with reddish brown marbling

and spotting ndndoensis

Dorsum chocolate brown, dorsolateral streak and dorsal spotting of light gray, bordered by

narrow brown or black edge palmatus

6. Limbs smooth 7

Limbs not smooth; may be shagreened, pustulate or warty 8

7. Ground color of dorsum of body light tan, with many small, rounded, regular darker

brown spots balios

Ground color of dorsum dark brown, with small, light yellowish spots often bilaterally

symmetrical longirostris

8. Entire body and limbs light brown, with small, scattered, dark spots above and below

halihelos

Dorsum light tan, vermiculated with light to dark brown, venter yellowish white, no

spotting ptanispimu

9. First finger distinct, free at least in part from webbing 10

First finger indistinct, entirely enclosed in webbing bufoniformis

10. Areas of spinules, warts, or pustules on some parts of head and/or body, usually on sides

and temporal region 11

Skin of head and body entirely smooth, no pustules, warts, or spinules boutengeri

11. Dorsum not unicolor, or, if so, not black 12

Dorsum unicolor, usually black, color extending well down sides and onto venter ..ignescens

12. Dorsum with strongly contrasting light (white or yellow) and dark (usually black) areas,

legs dark brown to black, usually with yellow spotting pachydermus

Dorsum not as described 13

13. Middorsum yellowish or greenish, with few scattered dark spots or uniform; no pustules

or warts on humerus, tibiofibula, or tarsometatarsus; dorsum of body usually smooth,

but few individuals have low warts posteriorly 14

Color not as described; all parts of limbs usually with warts, pustules, or spinules; dorsum

may or may not be warty 15

14. Humerus smooth, no pustules or glandular areas; ratio head length/SD x 100 greater

than 33 bomolochos

Humerus pustulate in females, with small rounded glands in males; ratio HL/SD X 10°

smaller than 33 ignescens

15. Middorsum of body with many low, rounded warts, which may have one to many whitish

spicules on them, venter usually dark brown with scattered darker spots nepioxomus

Middorsum of body smooth, no warts or pustules; venter yellow, without spots (bright

red in life) arthuri
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The males are smaller than the females in those

species where both sexes have been taken. The

forelimb of the male is stouter and more muscular

than in the female, presumably also associated with

the mating behavior, at least as observed in A.

ignescens (see elsewhere). The humeral area of

the forearm is usually noticeably thinner than the

radioulna area, which is quite thick. As a result

the males have arms reminiscent of the cartoon

character Popeye the Sailor.

Atelopus arthuri, new species

FIGURE 13

HOLOTYPE.—USNM 193470, female, from 15

kilometers north of Pallatanga, Chimborazo Province, Ecuador, at an altitude between 2800 and

2860 meters, collected by Arthur J. Peters, February 14, 1959.

DIAGNOSIS.—A member of the ignescens group

of Ecuadorian Atelopus, with a vermiculate dorsal

pattern of light brown and dark brown on both

body and limbs; belly clear yellow (tomato red in

life) with a brown patch on buttocks and brownish

spotting or vermiculation on chin and throat in

some individuals.

DESCRIPTION.—Snout from above with pointed

tip; distinct angle in canthus at level of nostril,

canthus forms straight line angled outward from

nostril to eye; snout, canthus, and upper eyelid

heavily fleshy and raised, with continuous fleshy

fold from posterior corner of eye to back of head.

Muzzle between canthi distinctly concave, rest of

dorsum of head flat. Snout from side distinctly

protruding beyond upper lip and lower jaw, nostril directed laterally and slightly posteriorly; loreal region vertical anteriorly, distinctly concave

immediately before eye; temporal and tympanic

areas glandular and pustulate. Dorsum of body

quite smooth or finely wrinkled, without pustules;

sides of body and most of upper surfaces of limbs

heavily pustulate; females with whitish spinules

singly or in clusters on top of pustules. Ventral

surfaces of body, chin, and limbs smooth or with

very fine wrinkles, no or very few pustules.

Forelimb very short; if pulled back along body,

tip of longest finger does not reach vent. Forearm

slightly larger in diameter than humeral region.

Forefoot fleshy, with fleshiness extending into web,

FICUKE 13.—Holotype, USNM 193470, of A. arthuri, new species.
NUMBER 145 11

which thus appears reduced, although actually including first phalanx of each digit; three phalanges

of third and fourth digits are free of web. No

tubercles on ventral surface of forefoot in most

individuals; some have series of fleshy swellings

both sub- and interarticular. Palmar tubercle large,

low, rounded; plantar tubercle very weakly defined.

Hind limb short, fleshy; when brought forward

along body heel may reach to points between posterior margin of head and axilla. Heels are well

separated when tibiofibulae are parallel to femora

at right angles to body. Foot with fleshy web, extending to tip of toe on all except fourth digit;

deeply incised between digits. Sole of foot smooth,

with poorly defined subarticular tubercles; outer

metatarsal tubercle smaller than inner, but raised

higher, making it more prominent; inner metatarsal tubercle flattened.

COLOR IN ALCOHOL.—Dorsum light grayish

brown with darker brown spotting, vermiculation,

or streaking; canthus, margin of eye, and parotid

swelling usually unicolor grayish brown (in some

individuals almost entire head is unicolor). Dorsum of limbs and feet as body, with same light

and dark vermiculations, except in some individuals, which show clear, almost colorless, slightly

yellow areas on first two digits and webbing of both

fore and hind feet. Sides of body as dorsum, with

tops of pustules appearing as white spots in some

individuals. Chest, belly, and ventral surfaces of

thighs and humerus light yellow, with viscera

and/or eggs showing through the skin in many individuals. A more or less enlarged brown patch

on the thighs below the anus, extending onto ventral surface for short distance. Chin and throat

yellow, almost always vermiculate or spotted with

dark brown, extending onto girdle region. Ventral surfaces of forearm, crus, and ankle areas of

limbs dark brown, occasionally punctate with yellow. Ventral surfaces of digits entirely yellow, with

palm and sole also yellow in some individuals.

When palm and sole are partly or wholly brown,

tubercles are yellow topped and contrast strongly

with brown areas. One individual (USNM 193466)

has a reddish brown venter rather than yellow.

COLOR IN LIFE.—Ground color of dorsum dark

brown, heavily reticulate with orange-yellow.

Ground color on sides dull reddish orange, with

dark brown reticulations. Large tomato-red spot

ventrally, on thighs, belly, and chest, entirely surrounded by a bright light green, which also covers

much of the ventral surfaces of arms and legs.

Dorsum of limbs usually as back, joints light

orange. Large metatarsal tubercle on outer edge

of foot bright yellow, lower surfaces of digits dull

orange, as is same surface of forefoot. Pupil black,

bright green ring around it, iris black with minute

green punctulations.

HOLOTYPE.—The holotype agrees with the description above in all characteristics for which no

variation is described. Where the species is variable, the holotype has the following characteristics:

skin on dorsum of body wrinkled in nuchal region, smooth on rest of back. Pustules on sides of

body and limbs with lighter areas, not clearly

spinulate. Ventral surfaces of body wrinkled, a

few pustules on throat. Scattered fleshy pustules

on ventral surface of forefoot. Plantar tubercle

not recognizable. Heel reaches to level of axilla.

Subarticular tubercles on hind foot somewhat more

prominent than in other individuals. Head with

brown vermiculations and spotting. No distinctive colorless areas on feet. Palm yellow, sole

brown, with lighter tubercles. Measurements (in

millimeters): snout to vent, 46.9; snout to end of

coccyx, 40.4; knee to knee, 32.3; tibiofibula, 15.5;

head length, 12.5; head width, 12.8; eye diameter,

3.0; snout to eye, 5.9; interorbital distance, 3.5;

width of eyelid, 3.5.

VARIATION.—Since the entire type series was collected by my son and me, we were able to gather

notes on the variation of the species in life. Individuals are extremely variable in the distribution

of the colors of the ventral surfaces, which are tomato red and bright green. One or two have the

dorsum almost totally green, with l^|le or no vermiculation. Others have more yellowish orange on

the back than dark brown. In addition, individuals

may have different colors than the majority, or different distribution of colors and pattern. One or

more individuals were taken with the following

variants: orange belly and flash areas; green spots

on chest, belly, and chin; entire venter green; dark

brown stripe or chevron across sternum; greenish

yellow reticulations on dorsum; bright green dorsolateral line from eye to groin or from tip of snout

to groin; and dorsum of feet green.

ECOLOGY.—The entire type series was collected

on February 14 and 15, 1959, at altitudes between

2800 and 2860 meters. The rainy season was late
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

during that year in that part of Ecuador, and the

frogs were actively breeding. Several mated pairs

were taken. The species is entirely diurnal, and

individuals were found wandering about over quite

dry terrain, using a slow walk and hopping only

when disturbed. The establishment of pairs appears to be entirely on a visual basis, and there

is no calling or chorus on the part of the exploring

males. The latter will raise themselves up high on

the forelimbs to look about for a female. Mated

pairs were found in pools formed in a fairly rapidly flowing mountain stream, but no egg deposition was observed. The voice in the species is characteristic of Ecuadorian Atelopus where known,

which is a low, chicklike peep, often not distinguishable at any distance. Occasionally males of

this species will trill the peep slightly.

ETYMOLOGY.—The species name is derived from

the first name of the collector of the holotype, who

provided me with pleasant company on the long

trip to Papallacta and back, and who collected

the first specimens of the new taxon while I was

busy fixing our supper.

SPECIMENS EXAMINED.—The following are paratypes: ECUADOR: Chimborazo Province, 15 km

N of Pallatanga, between 2800 and 2860 m altitude

(AMNH 79461, UKMNH 108938-39, UMMZ

131683, CAS 134199, MCZ 84199, USNM 193463-69,

193471-75).

Atelopus balios, new species

FIGURE 14

A.[telopus] longibrachius.—Rivero, 1968:22.

A.[telopus] spurrelli.—Rivero, 1968:22.

HOLOTYPE.—AMNH 17638, female, from the Rfo

Pescado, Guayas Province, Ecuador, collected by

G. H. H. Tate, May 28, 1922.

DIAGNOSIS.—A member of the longirostris group,

with long, thin legs and with heels overlapping

when tibiofibulae are parallel to femora at right

angles to body; first digit on forefoot distinct, not

hidden in webbing; limbs smooth. Similar to

longirostris, but differs in having a lighter ground

color with darker spotting, while longirostris has

a dark ground color with yellowish spots.

•".

FIGURE 14.—Holotype. AMNH 17638, of A. balios, new species.
NUMBER 145 13

DESCRIPTION.—Snout from above rounded or

with blunt point; outward curve of muzzle becoming inward curve from level of nostril to eye; snout,

canthus, upper eyelid, and temporal region somewhat fleshy. Dorsum of snout weakly concave, rest

of dorsum of head flat, lightly shagreened. Eye

slightly bulging dorsally. Snout from side protrusive over lower jaw and upper lip, forming blunt

point; nostril lateral, loreal region vertical or very

slightly concave. Eye slightly overhung by fleshy

margin of upper eyelid; temporal region more

heavily shagreened than dorsum, and also heavily

wrinkled. Dorsum almost smooth, very finely

wrinkled; sides with wrinkles forming deep creases;

chin and belly so heavily wrinkled that areas between wrinkles look like pustules.

Forelimb very slender, with forearm more fleshy

than humeral region, dorsum thickly shagreened,

ventral area wrinkled. When forearm is pulled

back along body, second and fourth digits reach

vent. Sole of forefoot fleshy, fleshy webbing encloses all but tip of first digit, extends as fringe to

ends of other digits, giving them a flattened appearance, webbing between toes incised to a point

about one quarter of distance to digit tips. All

subarticular tubercles almost totally indistinguishable, as are both plantar and palmar tubercles,

with latter occasionally forming low rounded

hump. Males with horny excrescence on first digit

of forefoot.

Hind limb slender, finely but densely shagreened

dorsally, less dense laterally, giving way to wrinkling on ventral surface. Tibiotarsal articulation

may reach to points between anterior edge of eye

and anterior margin of tympanic area when carried

forward along body. Heels touch or overlap when

tibiofibulae are parallel to femora at right angles

to body. Entire foot fleshy with thick webbing to

tips of toes, first three toes almost totally obscured

by webbing, which forms fringe on fourth toe from

level of tip of third and fifth toes. No subarticular

tubercles, very reduced outer metatarsal tubercle,

almost indistinguishable in some individuals, no

inner metatarsal tubercle.

COLOR IN ALCOHOL.—Dorsal ground color very

light tan, with numerous rounded spots, completely

separated from each other. Dorsum of legs, sides

of body as dorsum of body, side of head with dark

brown line from tip of snout through eye along

side of head to shoulder, line either continuous or

irregularly broken into spots along lower sides.

Upper lip light, rest of body below stripe usually

unicolor light yellowish white, a few individuals

with scattered spots on belly. Ventral surfaces of

limbs yellowish white, of feet dull yellow.

HOLOTYPE.—The holotype agrees with the description above in all characteristics for which no

variation is described. Where the species shows

variability, the holotype has the following characteristics: snout from above with blunt point, loreal

region vertical. Tibiotarsal articulation reaches

posterior corner of eye; heels overlap when tibiofibulae are parallel to femora at right angles to

body. Outer metatarsal tubercle present, distinguishable. Color pattern as shown in Figure 14,

lateral brown streak broken into series of spots, no

spots on belly. Measurements: (in millimeters)

snout to vent: 35.2; snout to end of coccyx: 33.5;

knee to knee: 28.4; tibiofibula: 15.7; head length:

9.2; head width: 8.6; eye diameter: 3.0; snout to

eye: 4.6; interorbital distance: 3.2; width of eyelid:

2.2.

REMARKS.—The AMNH catalog shows the type

locality as in Guayas Province, and notes made

by Tate would indicate it is near the Guayas-Azuay

border. Tate did not indicate what province he

was in when he collected the type series, but he

was on the mule trail from Naranjal to the highlands, and he estimated the altitude at the river

to be about 1400 feet above Naranjal, or approximately 500 meters. He noted that it was three hours

by trail from Naranjal to the river, which was at

the foot of the "real hills." He found the area

through which the river flows to be very wet, but

from there on the area became much drier. Since

the trail passes the 500-meter contour well before

the Guayas-Azuay border is reached, I agree the

specimens were probably collected in Guayas Province.

Rivero discussed the type series, and figured the

specimen here designated as the holotype (1968:22;

pi. 3: fig. 5). On the legend of the plate the AMNH

catalog number is given as 17738 rather than 17638,

which can be read on the photograph. Rivero considered the specimens possibly to be conspecific

with Atelopus longibrachius Rivero, a Colombian

species. When A. balios is compared with descriptions of the type material (Rivero, 1963:112; Cochran and Goin, 1970:135), the differences in color

pattern are distinctive, as pointed out by Rivero,
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and there are additional differences in the number

and distribution of pustules, the length of the forelimb, the presence and size of tubercles on the

palm and sole, and in other characters. Geographically the localities for the two taxa are widely

separated. Much collecting has been done by many

workers in the area between the two type localities,

but nothing resembling either taxon has turned up

in collections as yet.

In the same paper Rivero (1968:22) indicated

that some of the AMNH specimens from the Rio

Pescado belonged to Atelopus spurrelli Boulenger,

a Choc6an species. He has reference to AMNH

17548-49, which are marked differently, and the

pattern is certainly reminiscent of that of spurrelli.

I have noted, as well, that these two specimens have

a somewhat more prominent snout tip, which is

slightly hooked downward, as is seen in A. longirostris, and the third digit on the forefoot seems

to be markedly longer than in the type series. The

color pattern could be formed from fusion of the

discrete spots seen in the type series, however, and

this is what I think has happened, so I am regarding the specimens as members of the new taxon.

I do not regard them as typical, and they are not

designated as para types.

ETYMOLOGY.—The specific name is derived from

the Greek word balios, meaning "spotted" or

"dappled," and refers to the dorsal pattern of the

members of this taxon.

SPECIMENS EXAMINED.—ECUADOR: (Guayas

Province; Rio Pescado (AMNH 17619-30, 17636-

38, USNM 193957-58). All are paratopotypes. Assigned to this species, but not designated paratypes:

ECUADOR: Guayas Province; Rio Pescado

(AMNH 17548-49).

Atelopus bomolochos, new species

FIGURE 15

HOLOTYPE.—CAS 93910, female, from Ecuador,

Azuay Province, Sevilla de Oro, approximately

2800 meters. Collected January 9, 1958, by G. Frymire, A. Embree, and S. Strobel.

DIAGNOSIS.—A member of the ignescens group,

most similar to ignescens, from which it is distinguished by the light dorsal coloration often marked

with irregular dark spots; ratio head length/stand-

'

FIGURE 15.—Holotype, CAS 93910, of A. bomolochos, new species.
NUMBER 145 15

ard distance X 100 usually greater than 33; warts

and pustules absent on upper arm, crus, and ankle,

but present on sides from tympanic area to groin,

usually absent on dorsal and ventral surfaces of

body; first digit on forefoot distinct, almost totally

free of webbing.

DESCRIPTION.—Snout from above gently rounded

or with blunt point; distinct angle above nostril;

snout, canthus, and upper eyelid fleshy and raised,

with flattened, glandular, fleshy ridge from posterior corner of eye to end of head. Dorsum of

head between fleshy areas slightly concave or flat,

smooth. Eyes protrude laterally, dorsal area of eye

in same plane as rest of head. Snout from side

extends short distance beyond level of lower jaw

and upper lip, nostril lateral, occasionally directed

slightly backward, on raised, rounded area. Loreal

region distinctly concave; fleshy area over eye fused

with upper eyelid; upper lip slightly fleshy. Temporal area studded with glandular prominences,

rounded in males, covered with spiny points in

females. Middorsum of body completely free of

warts, glands, or pustules; usually quite smooth.

Dorsolateral area of neck with heavy, glandular

fold, shelflike in appearance, extending back from

dorsolateral temporal fold, sides wrinkled and

folded, covered by small white or yellow pustules

from tympanic area to groin, and on upper arms

and thighs; pustules may be single and evenly scattered or bunched together. Chin, chest, and belly

with wrinkles and folds, most prominent on chest.

No pustules or spicules on ventral surfaces, although occasional individuals show very tiny spiculations on chest.

Forelimb short and heavy; upper humeral region

with spicules, but lower humeral region and forearm free of spicules, smooth or coarsely wrinkled.

When forelimb is stretched along body, fingertips

do not extend beyond posterior margin of thigh.

Flesh of forefoot swollen; all digits free, with fleshy

webbing only at base. Sole of forefoot covered with

raised, rounded, glandular areas, subarticular tubercles not differentiated from these pustular areas;

palmar tubercle larger and well defined. Males

with horny excrescence on first and second digits,

first usually strongly flexed, with prominent, thick,

rounded fleshy hump dorsally, keratinized area

wide and extending from tip of digit to its base;

second digit with cornified area narrow, extending

from base of digit to terminal articulation.

Hind limb short, stout; femoral region spinose

on upper thigh only, close to body, rest of limb

smooth or slightly wrinkled, occasionally with

glandular areas on thigh and crus. Tibiotarsal

articulation reaches to points at arm insertion when

carried forward along body; heels are separated

when tibiofibulae are parallel to femora at right

angles to body. Foot fleshy, with very indistinct

scattered pustules on sole; webbing fleshy, extends

to digit tips as fringe except on first toe, which is

almost completely hidden in webbing; webbing

between all digits except first and second deeply

incised. Outer metatarsal tubercle raised and prominent, inner metatarsal tubercle indistinct.

COLOR IN ALCOHOL.—Dorsum of head, body, and

limbs light, may be yellowish, yellow-green, or light

brown; occasionally uniform, but usually with few

to many irregular, scattered, dark brown to black

spots. Dark dorsolateral line begins on side of

snout just below fleshy fold, runs through eye and

across temporal region onto sides, where it may be

continuous and broad, narrow and jagged, or almost totally absent. Ventral surfaces unicolor

white, yellowish white or yellow; occasional individual with one or two small brown spots on

throat or below anus. Spots often present at limb

articulations, may extend around limb, but in no

instance continuous ventrally. Sole of forefoot

always unicolor, sole of hind foot usually unicolor,

may be spotted or streaked.

HOLOTYPE.—This specimen conforms with the

description given above for the species, where no

alternatives are given. In the case of alternatives,

it has the following characteristics: snout rounded;

dorsum of head slightly concave, temporal region

lightly pustulate. Skin on chest and throat lightly

spiculate. Dorsum yellowish, with many black

spots, dorsum of limbs with black reticulation; two

black spots on throat at chest. Measurements of

type (in millimeters): snout to vent: 50.4; snout

to end of coccyx: 45.0; knee to knee: 34.6; tibiofibula: 17.0; head length: 12.7; head width: 12.6;

eye diameter: 4.0; snout to eye: 5.7; interorbital

distance: 3.8; width of eyelid: 3.6.

REMARKS.—The tendency seen in these individuals to have spots of color at the articulations of

the limbs is reminiscent of the stronger development of rings around the joints in A. botdengeri.

This species is comparatively large, and could cer-
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tainly be considered as a close relative of boulengeri.

Of the seven species described as new in this

paper, bomolochos is perhaps of most questionable

validity. It is clearly closely related to ignescens,

and, as is pointed out in the discussion of that

species, the variability of ignescens is considerable.

It could easily encompass the differences on which

bomolochos is based, although I have as yet seen

specimens with the particular constellation of characters used to define bomolochos only from a

comparatively limited area. There is a distinct possibility that the two taxa are sympatric, although

there seem to be no existent collections where a

single series contains both taxa. Both the California Academy and the University of Kansas have

had active collectors working along the PanAmerican Highway in Canar, Azuay, and Loja

Provinces in the last few years, and a fairly continuous series of ignescens can be put together from

the available material. While these southern specimens are not identical with northern populations,

they are not so distinct that I think any purpose

would be served by describing them as different.

A single individual tentatively called bomolochos,

CAS 93906, is from a locality well to the north of

the remaining sample, and was also collected along

the Pan-American Highway. There is a small series

from Cuenca which I feel is legitimately associated

with the new taxon. This locality, too, is on the

highway. Specimens called ignescens in this paper

come from both north and south of these two localities on the highway, and one gains a distinct

impression of either interdigitation or sympatry.

The two taxa, bomolochos and ignescens, are not

distinguishable on the basis of any of the measurements shown in the figures, with the possible exception of head width/head length. There are insufficient males of bomolochos available for statistical analysis, but the females of bomolochos are

distinct from those of ignescens with regard to this

character. Otherwise, the variation in bomolochos

is almost completely within that expected or observed in ignescens.

It is worth noting that both workers from the

University of Kasnas and several of my field parties have collected in the area from which bomolochos is recorded without obtaining specimens.

My party, including two students, an experienced

Ecuadorian collector, and Dr. C. F. Walker, was

in Sevilla de Oro in July 1962, but we took no

Atelopus at all. The ignescens material at the

University of Kansas was collected in the months

of May, June, and July. The California Academy

collectors of the type material were in the area in

the months of January and February. This seems

to indicate a seasonal occurrence and disappearance for bomolochos, but may simply reflect the

breaks of the game.

ETYMOLOGY.—The specific name is directly from

the Greek word bomolochos, meaning "toady," or

"lickspittle." While perhaps a stretch of the imagination for those who believe in strict interpretation of the literal meanings of words, it delights

me to think of the species as a "toady type frog,"

or, for that matter, since it is a bufonid, a "toady

type toad."

SPECIMENS EXAMINED.—The following are designated as paratypes: Azuay: Sevilla de Oro, 2800 m

(CAS 85341-42, 93910; holotype: 93912, USNM

193956). The following are tentatively identified

as bomolochos, but are not designated as paratypes: Azuay: Cuenca, 2535 m (CAS 85139-41,

85161). Canar: between Juncal and General Morales, approximately 2500 m (CAS 93906).

Atelopus boulengeri Peracca

FIGURE 16

Atelopus Boulengeri Peracca, 1904:20 [type locality: "di

Gualaquiza e parecchi di S. Jose, Ecuador orientale"].

Atelopus bicolor Noble, 1921:3, fig. 4 [type locality: Cordillera

Kutuku, east of Macas, Ecuador, 1800-2000 m].—Rivero,

1963:107.

A.[telopus] boulengeri.—Nieden, 1926:82.—Rivero, 1963:107.

SYNTYPES.—Six specimens, UT An. 560, from

San Jose, and eleven specimens, UT An. 559, from

Gualaquiza, in the Zoological Museum of the University of Torino, Italy.

DESCRIPTION.—Snout from above with rounded

tip, outward curve of muzzle becoming inward

curve from nostril to eye; snout, canthus, and upper

eyelid fleshy and raised, with fleshy ridge from posterior corner of eye to edge of head. Dorsum of

head between these raised, fleshy areas flat, broad

and smooth, lying in same plane as broad, flat back.

Snout from side slightly protrusive beyond upper

lip and lower jaw, nostril lateral, below fleshy area

of canthus, and directed slightly posteriorly. Loreal

region strongly concave behind nostril; eye over-
NUMBER 145 17

FIGURE 16.—A representative individual, USNM 193491, of A. boulengeri Peracca.

hung by fleshy margin of eyelid, upper lip somewhat fleshy. No tubercles or pustules anywhere on

head or body. Dorsum and sides of body smooth or

finely wrinkled, belly with more prominent and

distinct wrinkles.

Forelimb slender, forearm usually somewhat

more fleshy than humeral region, dorsum weakly

pustulate, rest of arm smooth. When forelimb is

pulled back along body, palm of hand reaches vent.

Entire hand fleshy, quite heavy around first digit,

which is clearly distinct and prominent, second

digit shorter than fourth, subarticular tubercles

at base of digits only. Tips of digits prominent,

slightly swollen ventrally. Palmar tubercle distinct,

plantar tubercle present. Edge of prepollex with

horny excrescence in males.

Hind limb slender, smooth except for few low,

very indistinct pustules on dorsum. Tibiotarsal

articulation reaches to points between posterior

margin of eye and arm insertion when carried

forward along body; heels do not overlap or touch

when tibiofibulae are parallel to femora at right

angles to body. Entire foot with fleshy webbing to

tips of toes; webbing somewhat excised between

last three toes and reaching tip of fourth toe as

fringe only; first and second toes almost totally

hidden in webbing, tips of all toes free and somewhat prominent, with slight swelling. Subarticular

tubercles weak, outer metatarsal tubercle raised

and prominent but smaller than the flattened,

rounded inner metatarsal tubercle.

COLOR IN ALCOHOL.—Dorsum usually unicolor,

chocolate brown, dark brown, or blackish; extending slightly onto sides of body, where it is sharply

set off by yellow of sides and belly, both of which

are also unicolor. Dorsum of head with dorsal color

extending to varying degrees anteriorly; in some

it.ends at eye level, in others it covers snout. Side

of head, lips, chin, and throat all monochrome

yellowish. Limbs usually slightly darker yellow

than venter, with rings of brown around all joints.

Dorsum of first finger and first two toes often

lighter than rest of foot. Ventral surfaces of feet

usually dark, with lighter tubercles. Brown of

dorsum usually extends over posterior margin of

legs and forms brown "seat" or "diaper" on rear

of venter, with anus usually bright yellow in center

of this brown area.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

COLOR IN LIFE.—Rich chocolate brown dorsally,

with bright yellow laterally and ventrally.

ECOLOGY.—The two specimens collected in 1962

were found along a trail in fairly heavy subtropical

forest, in full daylight, about 11:00 A. M. Both

were walking, not hopping, along the trail, fully

exposed, and made no effort to avoid being captured.

SYNTYPES.—The 17 syntypes agree completely

with the description above, with the series exhibiting all the variations mentioned in the description.

Some of the specimens from Gualaquiza are somewhat dried, and the digits do not have either as

thick or as full a webbing as in fresh specimens,

but the San Jose series is in good condition, and

the digits are as described. Only one specimen in

the series shows a yellow area at the anus; the

rest have the anal region the same color as the

dorsum.

COMPARISONS.—The differences between boulengeri and bicolor appear to be the consequence

of differences between investigators, rather than

between the types. The primary distinguishing

characters pointed out by Noble in the description

of bicolor were the development of the webbing

and several proportions. Noble said the digits of the

forefoot were webless, while Peracca described the

webbing between the second, third, and fourth

fingers as rudimentary, and as relatively well developed between the first and second fingers. The

area between these digits is actually quite fleshy,

and it appears that Peracca considered this fleshiness equivalent to webbing, while Noble, who

illustrated this area clearly in his figure, thought it

should not be called webbed at all. The description

of the webbing of the hind foot by Peracca is

quite consistent with Noble's figure, although Peracca described the foot as strongly webbed, while

Noble said, "toes webbed to the tips . . . but the

web deeply indented, making the toes appear half

webbed." The differences in proportions are quite

minor, and could easily be the result of different

techniques in measuring. Thus, in boulengeri the

head length is contained in the body length 2i/2

to 3 times, according to Peracca, while in bicolor,

according to Noble, the body is 3i/2 times as long

as the head. But Noble clearly indicated that the

head length meant from snout to angle of jaw, and

no one knows for sure how Peracca made his head

length measurement. There are several other standard head length measurements that might give

a larger value, and thus a smaller proportion, for

Noble's specimens.

Although there may have been some geographical

validity for retaining the two species prior to our

field work in 1962, the two specimens we obtained

weaken this validity considerably. The type series

for boulengeri came from Gualaquiza, a town in

the valley of the Rio Zamora, at 950 meters. Noble's

specimens came from quite a bit higher in an

isolated Cordillera to the east of the valley of the

Rio Upano, a tributary of the Namangoza, which

joins the Zamora to form the Rio Santiago. The

two new specimens come from the eastern slopes

of the main Andean chain at an altitude of 1560

meters, and some distance to the west of the valley

of the Namangoza. This indicates an altitudinal

range of about 600 meters, with the species occupying the foothills and mountain slopes of the headwater streams of the Santiago up to an altitude

of 1560 meters.

Rivero (1963:110) regarded A. bicolor Noble

as probably not referable to any other Ecuadorian

species, but later (1968:23) indicated that it is

"certainly a synonym of A. boulengeri Peracca."

He records four specimens from Turula, which he

indicates as being east of Macus, but there is only

one specimen in the AMNH from Turula, which

is a "tambo" on one of the mule trails from Riobamba to the Amazonian region, and lies northwest

of Macas. The bicolor paratypes, AMNH 13133-35,

are not from Turula (see below).

SPECIMENS EXAMINED.—ECUADOR: no additional data (AMNH 17427-28, 17430-31). MoronaSantiago Province; Cordillera Cutucii (AMNH

13132—holotype of bicolor, AMNH 13133-35—

paratypes of bicolor, UKMNH 108940, USNM

193493), Gualaquiza (UT An. 559—syntypes of

boulengeri), Rio Piuntza, 1830 m (UKMNH

147071-81), San Jose (UT An. 560—syntypes of

boulengeri), Turula (AMNH 33893), between

Mirador and Copal, on mule trail from Sevilla de

Oro to Mendez, 1600 m (USNM 193491-92). Loja

Province; San Ramon (JAP 8934).

Atelopus bufoniformis Peracca

FIGURE 17

Atelopus bufoniformis Peracca, 1904:20 [type locality: El Pun,

Ecuador].
NUMBER 145 19

FIGURE 17.—A representative individual, USXM 193537, of A. bufoniformis Peracca.

SYNTYPES.—Two specimens, UT An. 561, in the

Zoological Museum of the University of Torino,

Italy.

DESCRIPTION.—Snout from above angulate and

pointed, with a sharp angle at level of nostril;

canthi from nostril to eye almost parallel, without

a curve; eye strongly protruding and directed

slightly forward; canthus fleshy and raised above

median area of snout; dorsum of eye fleshy with

many wrinkles and grooves. Dorsolateral area of

head behind eye with a prominent ridge from eye

level extending posteriorly along back, converging

on ridge from opposite side to a point above shoulder, at which point they diverge until they disappear. Second, more glandular ridge arises at corner

of eye and runs over tympanic region, either ending

in small boss at eiul of head, or continuous with a

glandular dorsolateral fold. Rest of head dorsally

covered with wrinkles, grooves, and pustules.

Snout from side extends very slightly beyond end

of lower jaw, and is either bluntly rounded or

weakly pointed. Loreal region vertical, not concave; entire side of head with rather thick skin,

wrinkled, grooved, and with small, fleshy, flattened

warts, increasing in prominence over tympanic

area. Entire body, limbs, chin, throat, and belly

heavily pustulate or warty, with pustules low and

flattened.

Forelimb very short, slender. When limb is carried along body, digits barely reach insertion of

hind leg. Hand with very fleshy webbing enclosing

first digit completely, second and fourth almost

as completely, and on third digit to tip as fringe,

with whole foot giving appearance of toad wearing a mitten. Entire lower surface of hand as

heavily pustulate as rest of body, obscuring subarticular tubercles completely; plantar tubercle

very vaguely defined. Hind limb short, slender;

tibiotarsal articulation reaches to points on side

well behind axilla when leg is carried forward

along body; heels widely separated when tibiofibulae are parallel to femora at right angles to

body. Entire foot with very thick, fleshy webbing,

enclosing fully all digits except fourth, which is

fringed to tip by web. No distinguishable tubercles

on sole, which is fleshy and wrinkled; outer metatarsal tubercle vaguely defined, inner metatarsal tubercle even more vaguely defined.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

COLOR IN ALCOHOL.—Dorsum of body, head, and

limbs entirely very dark brown to black; only occasional individuals with somewhat lighter brown

vermiculations on back. Venter of body brown,

with more or less vague dull yellowish mottling.

Sole of feet lighter than rest of ventral surfaces.

COLOR IN LIFE.—USNM 193537 was very reddish

in life, and the warty pustules on the venter were

a very bright red. USNM 193538 had yellow pustules on the venter.

SYNTYPES.—The two syntypes are badly dried,

but it is still possible to identify them as bufoniformis in the sense of this paper. The larger of the

two syntypes is the individual for which measurements are presented in the original description.

The second specimen is about 20 millimeters long.

The larger specimen fits the description above

except as follows: the glandular ridges arising at

the corners of the eye are continuous with the

glandular dorsolateral folds. The snout from the

side is bluntly rounded. The specimen is too badly

dried to carry foot forward, but Peracca said that

the tibiotarsal articulation "raggiunge scarsamente

la spalla."

REMARKS.—This is a very small species. The

largest individual seen measured 33 millimeters

from snout to end of coccyx, and most specimens

were less than 30 millimeters long. The shortened

limbs and increased padding or fleshiness of the

feet seem to be highly adaptive, providing reduced

surfaces for heat loss. The species seem well adapted

for high altitude life, and has been taken as high

as 3700 meters.

Cochran and Goin (1970:120) described and

figured a single specimen of this species from the

Paramo de las Papas, Cauca, Colombia. It is correctly identified, and thus A. bufoniformis shows

a wider distribution than any other highland

species except A. ignescens. As I have pointed out

elsewhere in this paper, these are the only two

species that have been able to break out of the

isolating mechanisms faced by all high altitude

species of Atelopus, and extend their distribution

into higher areas of both Ecuador and Colombia.

SPECIMENS EXAMINED.—ECUADOR: Napo Province; El Pun (UT An. 561-syntypes), Environs of

Santa Barbara (USNM 193537-40). Carchi Province; Paramo del Angel, 23 km SW of Tulcan,

3700 m (UKMNH 117880). Imbabura Province:

Cordillera de Intag, La Delicia, 2700 m (UKMNH

132126), N slope of Nudo de Mojanda, 3400 m

(UKMNH 131797-98).

Atelopus elegans (Boulenger)

FIGURE 18

Phryniscus elegans Boulenger, 1882a: 155 [type locality: Tanti,

Ecuador, 2000 feet]; 1882b:464, fig. 1.

Atelopus elegans.—Boulenger, 1894:375; 1898:119.—Peracca,

1904:22.

A.[telopus] elegans.—Nieden, 1926:81.

Atelopus [elegans] elegans.—Rivero, 1963:109.

HOLOTYPE.—British Museum (Natural History),

one female.

DESCRIPTION.—Snout from above rounded, not

ending in point; distinct angle in canthus at nostril level; canthus and upper eyelid not fleshy,

dorsum of head smooth and flat. Snout from side

with distinct projection, ending in blunt point,

extending beyond symphysis of lower jaw; loreal

region vertical, not concave. Head dorsally and

laterally smooth, without spines, warts, or pustules.

Body slender, skin completely smooth.

Forelimb long, slender; when laid back along

body, palm reaches anus. Webbing fleshy, enclosing

and almost completely obscuring first digit; full

webbing between first and second digits; web extends to tips of other digits but is deeply incised

between them. Fourth digit extending beyond level

of penultimate joint of third; considerably longer

than second digit. Palm of hand fleshy, with deep

folds; palmar tubercle low, rounded; plantar tubercle weak and indistinct.

Hind limb long, slender; when brought forward

along body, tibiotarsal articulation reaches points

between loreal region and posterior margin of

eye; heels overlap when tibiofibulae are parallel

to femora at right angles to body. Hind foot with

thick, fleshy webbing; first and second digits almost totally enclosed, with only their tips distinguishable; other digits with full webbing that

is shallowly incised, and extends to tip of fourth

digit as fringe only. Fifth toe longer than third.

Sole of foot quite fleshy with deep grooves or folds,

no clearly defined subarticular tubercles; outer

metatarsal tubercle poorly defined, inner metatarsal

tubercle almost indistinguishable.

COLOR IN ALCOHOL.—Dorsal ground color white,

with more or less regular pattern of brownish or
NUMBER 145 21

FIGURE 18 A representative individual, UMMZ 51270, of A. elegans Boulenger.

black consisting of parallel stripes beginning on

eyelid, where they may be joined by bar between

eyes, and extending to just short of anus, which is

usually in isolated spot of brown narrowly separated from stripes. Dorsal stripes irregularly connected by vermiculations and lateral bars, or separated by additional stripes running lengthwise,

by light spots surrounded by brown borders, or

by brown spots surrounded by white. Solid white

line from eye to groin along dorsolateral line in

most individuals, bordered below by dark line

beginning at tip of snout, passing along lateral

area of canthus, and continuous from eye to groin.

No pigment on sides or belly below this lateral

dark streak. Dorsal areas of limbs and feet mottled,

streaked, and striped with dark brown to black, as

body; ventral areas of appendages unicolor yellowish white.

COLOR IN LIFE.—Unknown, but several specimens are very yellowish in ground color, indicating that this might be the living color.

REMARKS.—Rivero (1963:109) regarded Atelopus

gracilis Barbour, from Gorgona Island, Colombia,

as a subspecies of Atelopus elegans. Cochran and

Goin (1970:133) made elegans a subspecies of A.

varius, with gracilis a strict synonym. They did not

refer to Rivero's action in their synonym. Neither

Rivero nor Cochran and Goin presented any documentation for the changes they made. Rivero

stated that specimens from Ecuador "seem sufficiently distinct from Gorgonan material to make

the island form deserve the racial name A. elegans

gracilis." He does not describe the differences or

define the race. Cochran and Goin reduced several

species to the subspecific level within A. varius,

but again the reasons for doing so were not clearly

stated. Even though it is entirely possible that

elegans represents the southern end of a genetic

continuum which embraces both high and low

altitude populations, I have seen no evidence to

support the concept. A. elegans is a distinct, easily

recognized, well marked taxon, and I prefer to
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

recognize it as a full species until intergradation

with the subspecies of varius can be demonstrated.

The relationship between gracilis and elegans may

be subspecifk, but I have not examined the gracilis

material, and leave that question in abeyance for

other investigators.

In 1955, I referred to the locality called "Buliin"

in the American Museum catalogs as unlocated.

Levi-Castillo, in an editorial footnote to my remarks, suggested that it was in reference to "BuluBulu," a river in Guayas and Caiiar Provinces. The

United States Board on Geographical Names (1957,

p. 138) lists a place called "Pulun" at the coordinates l°05'N, 78°40'W, and it is shown at those

coordinates on the 1:1000000 map of the Instituto

Geografico Militar del Ecuador (1950). The American Museum specimens were collected at about the

same time as material from Salidero and Rio

Durango, both of which are shown on a map of

Esmeraldas Province in Acosta-Solis (1944, opp. p.

574). Both localities are very close to Pulun, which

also appears on the map in the work by AcostaSolis. The coordinates for Salidero would be approximately l°04'N, 78°38'W, and the town lies

on the Rio Bogota. The Rio Durango enters the

Bogota from the east about halfway between Salidero and Pulun. I am, therefore, using the name

"Pulun" for the place where the AMNH specimens

were collected.

SPECIMENS EXAMINED.—ECUADOR: Esmeraldas

Province; Pulun (AMNH 10581-84, UMMZ

51270), Quininde (USNM 193534), Rio Cupa

(UKMNH 108911, USNM 193535), Rio Durango

(AMNH 10585-88), Salidero (AMNH 10577,

10579-80, MCZ 2615, 7605). Imbabura Province;

Lita, Rio Mira (USNM 193536).

Atelopus halihelos, new species

FIGURE 19

Atelopus [species].—Rivero, 1968:21.

HOLOTYPE.—AMNH 16716, male, from Ecuador,

Morona-Santiago Province, Cordillera Cutucii,

collected by E. Feyer, March 1921.

DIAGNOSIS.—A member of the ignescens group,

although limbs are more slender than other species;

most similar to nepiozomus, from which it differs

in having an almost indistinct first digit on the

forefoot and in coloration, particularly in the

5 I t

<

FIGURE 19_Holotype, AMNH 16716, of A. halihelos, new species.
NUMBER 145 23

presence of a light whitish patch below the anus,

in contrast to the dark "diaper" of nepiozomus.

DESCRIPTION.—Snout from above with rounded

point, distinct angle in canthus rostralis at nostril;

snout, canthus, and eyelid fleshy and raised, with

either fleshy ridge or row of raised pustules from

corner of eye along dorsolateral margin of head.

Dorsum of head flat, with scattered, low, glandular

pustules. Snout from side protrusive, pointed, extending well beyond upper lip and lower jaw;

nostril lateral, below fleshy area of canthus, at top

of rounded prominence. Loreal region flat or

slightly concave; eye overhung by flap formed of

fleshy area, flap has free edge and can be lifted away

from the eyelid; upper lip swollen. Temporal

region with low, glandular pustules, which may

bear pointed spicules in either sex. Dorsum and

sides of body smooth, with many low, rounded,

scattered warts. Ventral surfaces may be either

comparatively smooth or with flattened, indistinct

glandular patches separated by wrinkles and folds.

Forelimb slender in females, shorter and more

muscular in males, particularly in the forearm.

When limb is carried along body, digit tips reach

beyond posterior margin of thigh. Dorsal and lateral surfaces of limb covered with warts, more

prominent in males than in females. Hand with

fleshy webbing, first digit almost entirely enclosed,

other digits free. Males with keratinized pad on

dorsum of first digit, no horny excrescences on second digit. Sole of forefoot smooth or with few low

pustules; plantar tubercle weak, defined by absence

of pigment in area.

Hind limb long, slender, covered dorsally and

laterally with low warts; tibiotarsal articulation

reaches to area between eye and end of head when

leg is carried forward along body; heels touch or are

barely separated when tibiofibulae are parallel to

femora at right angles to body. Webbing thick but

not heavily fleshy, extending to tips of all but

fourth digit, incised between all digits except first

and second; only tip of first digit free of web. Sole

almost entirely smooth, subarticular tubercles usually present, but low and often indistinct; outer

metatarsal tubercle fairly prominent, distinct; inner metatarsal tubercle larger than outer, but low

and indistinct.

COLOR IN ALCOHOL.—Entire body and limbs light

brown, with small, scattered, irregular dark brown

or blackish spots on all surfaces. Ventral area of

thighs and belly immediately below anus very light

colored, often whitish, without spots.

HOLOTYPE.—This specimen conforms with the

description given above for the species in those

characteristics for which no variation is mentioned.

In the case of variable characters, it has them as

follows: a row of raised pustules along dorsolateral

margin of head behind eye; loreal region slightly

concave. Warts in temporal region do not bear

spicules. Chest and throat with low, indistinct

pustules. Heel carried forward along body reaches

temporal region; heels do not touch when tibiofibulae are parallel to femora. Soles of both fore

and hind foot with few indistinct tubercles, subarticular tubercles on hind foot slightly more

prominent. Measurements of type (in millimeters):

snout to vent: 25.3; snout to end of coccyx: 23.4;

knee to knee: 20.3; tibiofibula: 9.8; head length:

7.8; head width: 7.2; eye diameter: 2.6; snout to

eye: 4.0; interorbital distance: 2.6; width of eyelid:

2.2.

REMARKS.—This series of specimens was discussed

by Rivero (1968:21), and he felt their affinity was

closest with A. bufoniformis. Since there are neither

enough females of A. halihelos nor males of A.

bufonifmmis available for statistical analysis, the

two species cannot be directly compared on the

graphs provided earlier in this paper. Relative position with regard to other species, however, shows

that halihelos is more slender, with longer limbs,

and it appears more closely related to A. nepiozomus and other species on the lower Amazonian

slope than to the highly specialized, high altitude

A. bufoniformis. As in the latter species, it appears

that halihelos is a diminutive species, with the

largest specimen in the type series less than 32 millimeters long.

There is no way to be sure about the actual distribution or altitudinal relationships of this new

species. The Cordillera de Cutucii is a fairly isolated

mountain mass to the east of the valley of the Rfos

Upano and Namangoza, with the foot of the Cordilleras at approximately 1000 meters, and the highest point somewhat more than 2000 meters. I would

predict that the species will be found at altitudes

above the lower edge of the Cordillera, and possibly to the highest points. It seems to be a situation

similar to that of A. planispinus, which inhabits

the slopes of a similar isolated mountain mass to

the north.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ETYMOLOGY.—The specific name is derived from

the Greek hales or halis, meaning "in abundance,

in heaps, crowds or swarms," and helos, a "wart"

or "callus." The reference is to the large number

of glandular swellings on the body and limbs of

individuals of this taxon.

SPECIMENS EXAMINED (in addition to holotype).—

The following are paratopotypes: ECUADOR: Morona-Santiago Province; Cordillera Cutucii (AMNH

16717-24, USNM 193959).

Atelopus ignescens (Cornalia)

FIGURE 20

Phryniscus ignescens Cornalia, 1849:316 [type locality: "in

locis humidus circa Latacunga prope Quito," Ecuador].

Phryniscus laevis Giinther, 1858:43, pi. 3: fig. A [type locality:

"Panama. Quito. Chili"]; 1859:417.—Boulenger, 1882:151.

Phrynidium laeve.—Cope, 1867:196.

Atelopus laevis.—Cope, 1868:117.—Orton, 1871:693.

Atelopus ignescens Espada, 1875:139, pi. 1: figs. 7-9.—Boulenger, 1880:46; 1898:119 Werner, 1901:600.—Peracca,

1904:19.—Parker, 1934:268; 1938:440.—Rivero, 1963:109;

1969:142.

Phryniscus laevis exigua Boettger, 1892:22 [type locality:

"Zurucuchu, W-Andes of Cuenca, 3250 m, Ecuador"].

A.[telopus] ignescens.—Nieden, 1926:83.

Atelopus carinatus Andersson, 1945:15, fig. 5 [type locality:

"Eastern Ecuador"].—Rivero, 1969:142, fig. 2.

Atelopus ignescens ignescens.—Rivero, 1965:137.—Duellman

and Lynch, 1969:236.

HOLOTYPE.—In the "Museo Mediolanense,"

present state unknown.

DESCRIPTION.—Snout from above not prominent

or projecting, gently rounded, with little or no

difference in angle between tip of snout and eye;

snout, canthus, and upper eyelid heavily fleshy and

raised, with a glandular, fleshy ridge from posterior

corner of eye to corner of head. Dorsum of head

between fleshy areas flat, smooth, or very finely

wrinkled. Snout from side only slightly protrusive

beyond lip and lower jaw, nostril lateral and below

fleshy area of canthus. Loreal region vertical or

slightly concave; eye overhung by heavily fleshy

upper eyelid, upper lip not fleshy. Temporal area

studded with pointed tubercles in females, or with

fleshy, glandular bumps in males. Middorsum of

body completely free of warts, glands, pustules;

either smooth or finely wrinkled. Dorsolateral area

and sides of body in females heavily studded with

rounded glands that have one to several raised,

pointed spicules on them, giving spiny appearance;

same area in males covered with low, rounded

FIGURE 20.—A representative individual, JAP 4176, of A. ignescens Cornalia
NUMBER 145 25

glandular areas without spicules. Chin and belly

finely wrinkled and occasionally lightly shagreened

on chest in males, same in females, but with tendency in some individuals to have darkened spiny

area across chest.

Forelimb short, stubby, and fleshy; humeral region heavily studded with spicules in females, covered by small rounded glands in males; forearm in

both sexes heavily fleshy but quite smooth, contrasting strongly with upper arm. When forelimb

is pulled back along body, tips of digits do not

reach past hind leg in males or past groin in females. Entire forefoot fleshy, all digits free and

prominent, with fleshy webbing only at base. Sole

of foot covered with pustules, subarticular tubercles not differentiated from these pustules, palmar

tubercle larger and clearly defined. Males have

horny excrescences on both first and second digits;

first digit usually strongly bent under and flexed,

with prominent thick, rounded, fleshy hump dorsally, keratinized area extends from tip of digit to

base; second digit with cornified area only at base.

Hind limb short, stout, fleshy; femoral region

studded with spines in females, heavily pustulate

in males, crural area smooth and fleshy. Tibiotarsal

articulation reaches to level of arm insertion when

carried forward along body; heels are well separated

when tibiofibulae are parallel to femora at right

angles to body. Foot fleshy with many pustules on

sole, webbing fleshy, extends to tips of digits as

fringe except on first toe, which is almost completely hidden within webbing; webbing between

all digits, including first and second, deeply incised.

Subarticular tubercles not clearly distinguishable

from pustules, outer metatarsal tubercle small but

raised and prominent, inner metatarsal tubercle

larger but not as raised.

COLOR IN ALCOHOL.—Dorsum of head, body, and

limbs uniform black, extending down sides and

ending abruptly at edge of belly, which is yellow

to yellowish white. Some individuals have spotting

on chest. Ventral surfaces below anus and on limbs

often light brown to black; pigment distribution

ventrally differs considerably from one individual

to the next. Lower lip usually dark, same color as

upper lip, and sharply distinct from rest of chin.

COLOR IN LIFE (from a series of 25 individuals

collected near Quito June 6, 1959).—Dorsum dark

brown to black, lightening slightly on dorsum of

limbs. Dorsal color sharply set off from ventral

color along a line running from corner of mouth

along the lateroventral line. Ventral surfaces highly

variable, from a light cream unicolor to light cream

with yellowish or orange border to unicolor yellowish to unicolor orange. The eye is a deep black,

no pupil color can be distinguished. Often with

a bright orange or yellow spot on ventral side of

crus, and some have same on dorsum of foot. In

this same series of 25, the following variation in

color of the ventral surfaces was noted (individuals

can occur in more than one tabulation): venter

mostly dull orange-5; cream with orange in axilla

and inguinal region-4; yellowish, with brightest

yellow in limb insertions-2; yellow spot on dorsum

of foot-6; yellow spot on ventral surface of crus-8;

yellow venter with reddish orange in limb insertions-2; unicolor yellow-2; venter of limbs maculated with bright orange-1. A series from 5

kilometers west of Papallacta all had bright, tomato-red bellies in life but of course, now are

identical with other alcoholics in belly color.

REMARKS.—Tadpoles presumably of this species

have been described by Duellman and Lynch

(1969:236). The material was taken at 3620 meters

on the slopes of Cotopaxi Volcano. The tadpoles

have a large suctorial disc behind the mouth which

is used to anchor the individual to the underside

of stones in swiftly flowing water.

It will be noted that I have continued the synonym of A. carinatus Andersson with A. ignescens.

Rivero (1969:142) has recently reexamined the

holotype of Andersson's species, the only known

specimen, and thought that the species should be

considered valid on the basis of several characters,

including the "nuchal tank" formed by the protruding posterior margin of the frontoparietal and

the raised edges of the suprascapulae, the shape of

the snout, and the size of the eye. The first of these,

I am convinced, is the result of the way the specimen was preserved. The others are extremely

difficult to evaluate or quantify, seem of minimal

value taxonomically, and are, to my eye, at least,

occasionally duplicated in large series of ignescens

from a single locality. It should also be pointed out

that the type locality, vague though it is, is an area

of intensive activity by herpetological collectors,

and no additional specimens resembling A. carinatus has ever been obtained in eastern Ecuador,

while hundreds of animals looking very much like

the type continue to be collected in the inter-
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Andean valleys. Until added material from the

supposed type locality is available, I shall continue

to regard this weakly denned form as a synonym of

A. ignescens.

ECOLOGY.—A. ignescens is adapted to the most

rigorous climates found in Ecuador where amphibians occur. It inhabits the high inter-Andean

valleys, and has sufficient eurytopic abilities that it

has crossed the inter-Andean nudos to invade

neighboring valleys. It has also moved up the

slopes of the highest mountains above tree line,

where it is found in the cold, windy, and dry

paramos. It is tolerant of temperatures well below

levels at which other species of Atelopus become

inactive, except perhaps A. bufoniformis.

The species engages in mass migrations, apparently at the beginning of the breeding season, and

can be seen in the hundreds at such a time. I have

see two such migrations, the first of which was

moving across the Pan-American Highway north

of Latacunga on December 31, 1958. Large numbers of individuals were smashed on the road over

a distance of four or five miles. All living individuals were crossing the highway in the same direction. Some mated pairs were seen.

The second such migration was seen in the

paramos on the road from Quevedo to Latacunga,

on January 22, 1959, at an altitude of 3600 meters.

In this case, the movement was again in a single

direction, and definitely downhill. A light drizzly

rain was falling, and sufficient rain had already

fallen to make the road somewhat treacherous,

with standing puddles. In both cases, the migration seems to be toward standing water, presumably

the place of egg deposition, but this was not verified by direct observation.

COURTSHIP AND MATING BEHAVIOR.—The migration seen in the paramos included many clasped

pairs as well as males ready to clasp. The males,

always smaller than the females, clasp with an

axillary grip. I followed one such pair for almost

two hours, hoping to see egg deposition, but the

female simply wandered about in what appeared

to be a random way, never pausing anywhere long

enough for any action to take place. The rate of

progress is a slow, deliberate walk, never a hop or

a jump. At one time the female entered a small

puddle about five feet across and at most a foot

deep, but she never broke either her stride or her

pace, continuing to walk very deliberately across

the bottom of the puddle and out the other side.

Experiments performed later with mated pairs in

the compound of Robin Copping's house in Quito

demonstrated that once the male has mounted the

female he will remain in place no matter what the

female does until the eggs are deposited. This

may be a matter of several weeks, during which

time the female feeds and moves about normally,

but the male does not feed while he waits. The

period of waiting for the male may not be so long

under normal conditions, of course, but I suspect

it is usually much longer than in other frogs.

The activity period is diurnal, and I have never

seen members of this species active at night. The

males seek out the females entirely visually. While

they do make a slight sound, it is a very low, chicklike peep, difficult to hear even at short distances,

and made while walking about looking for a female, rather than made in a stationary position to

which a female could be attracted by the "call."

As they walk about they will occasionally pause,

raise themselves high on the forelimbs, and look

about, scanning the horizon for other members of

their species. There is no sexual discrimination in

this scanning, because the individual moves off in

the direction of another frog as soon as the second

is seen, and attempts to mount. If the second is a

male, the first is repelled by slow kicking and soft

peeping. If the second is a female, it can usually

be successfully mounted. If the second is a female

with another male already in mounted position,

the approaching animal is not put off for a second

by the fact of prior possession. He will, if permitted to do so, mount the first male, and clasp

him. Almost invariably, however, he is repelled by

the first male. Since all the movements by this

species are slow and deliberate, the repulsion almost appears to be in slow motion, but it still is

effective. The mounted male raises his hind foot

on the side of the approaching male, puts it very

deliberately in the face of the intruder, and softly

but forcefully kicks him away. The intruder usually describes a gentle arc, landing on his back,

and the mated pair moves away as he slowly struggles to turn over. If they have not gone far enough

by the time he recovers his equilibrium, a second

attempt will be made and the same sequence enacted.

SPECIMENS EXAMINED.—I have seen over 500 individuals of this species, from all of the inter-
NUMBER 145 27

Andean valleys of Ecuador, from Tulcan south to

the region of southern Cuenca Province. It seems

of little value to list them in detail until someone

has examined them more carefully for infraspecific

variation.

Atelopus longirostris Cope

FIGURE 21

Atelopus longirostris Cope, 1868:116 [type locality: "Valley of

Quito," Ecuador].—Orton, 1871:693.—Espada, 1875:155.—

Nieden, 1926:81.—Guibe, 1948:32.

Atelopus varius [not Stannius] Boulenger, 1880:46.

Phryniscus longirostris.—Boulenger, 1882:153.

Phryniscus boussingaulti Thominot, 1889:28 [type locality:

"Entre Latacunga et Guayaquil, au sud de Quito," Ecuador].

HOLOTVPE.—Apparently lost. Not located in

either the Academy of Natural Sciences of Philadelphia or the National Museum of Natural History.

DESCRIPTION.—Snout from above strongly projecting, with comparatively sharp point, distinct

inward curve from snout tip to nostril, canthi

almost parallel from nostril to point just before

corner of eye where each canthus flares outward;

area from snout tip along canthus, outer eyelid

border, and to corner of head raised and fleshy.

Dorsum of head flat, shagreened; eyes prominent,

protrusive. Snout from side strongly protrusive

beyond lower jaw, with upper lip distinct from

protrusion as well; nostril lateral, posteriorly directed, slightly below fleshy canthus, and almost

directly above end of lower lip. Loreal region

sunken, eye overhung by fleshy eyelid margin; vertical ridge behind eye and horizontal ridge from

eye to corner of head; side of head and tympanic

region shagreened, with few larger, white-topped

pustules. Dorsum and sides of body heavily shagreened, with scattered white-topped pustules on

sides only, belly almost smooth or with finely wrinkled surface. Vertebral column forms distinct,

raised middorsal ridge from shoulder level to coccyx.

Forelimb very slender, forearm more fleshy than

humeral region, pustulate on shoulder and dorsum of humeral region, shagreened elsewhere. Entire hand somewhat fleshy, heaviest around first

FIGURE 21. A representative individual, USNM 193834, of A. longirostris Cope.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

digit, which is distinct although almost totally enmeshed in fleshy webbing. Tips of digits slightly

swollen; second digit shorter than fourth; second,

third, and fourth digits almost free of webbing.

Subarticular tubercles present only at digit base,

weakly developed; large palmar, small plantar tubercle.

Hind limb long, slender, shagreened but without

pustules; tibiotarsal articulation reaches points between nostril and posterior corner of eye when

carried forward along body; heels overlap or touch

when tibiofibulae are parallel to femora held at

right angles to body. Fleshy webbing of foot extends to swollen tips of digits, but is incised between digits; first digit completely hidden in

webbing, tip of second distinct. Subarticular tubercles present but weak; outer metatarsal tubercle

raised, prominent; inner metatarsal tubercle

elongate, flattened, kidney shaped.

COLOR IN ALCOHOL.—Dorsum brown, from reddish brown to very dark brown, snout tip light

brown, sharply distinct from top of head, which

usually has few scattered white spots, characteristically one on back of each eyelid. Dorsum with

scattered, often bilaterally paired light spots; dorsolateral area over shoulder usually with elongate

white spot, almost forming stripe in some individuals, usually followed dorsolaterally by one or

two additional elongated spots or streaks. Sides

usually unicolor, occasionally lightly vermiculated,

abruptly changing into very light, immaculate yellow of belly, from chin to anus, with occasional

individual showing brown vermiculations ventrally. Legs as body, usually unicolor brown above,

spots laterally and dorsolaterally, light yellow below, posterior surface of thigh usually vermiculated. Tips of digits yellow, soles of feet usually

dark, but some with yellow areas.

COLOR IN LIFE.—Dorsum blackish brown, shading into a light chocolate brown low on sides.

Spots on head, back, sides, and dorsum of legs dull

yellow. Those on back of femur white. Tip of

snout yellowish brown, brighter than rest of spots.

Series of tiny white stipples along sides and between eye and arm, behind temple. Eye black all

over. Belly white, not flesh colored. Tips of digits

very light orange. Stripe along upper lip yellow

as dorsal spots, but grades into white of belly near

corner of mouth. In a female heavy with eggs, the

eggs show brightly through the belly flesh, giving

a strong orange tint to ventral surfaces.

ECOLOGY.—I have collected this species in several

places on the western slope, and always found the

specimens in rocky areas on the edges of streams.

I have found individuals under rocks and also out

walking about in and over the rocks in full sunlight. The habitat is not usually a shaded one, but

is very exposed. A pair in amplexus was taken at

Apuela on April 17, 1959, which was at the end

of the rainy season. The female was heavy with

eggs.

COMPARISONS.—An obviously close relative of

longirostris is A. varius glyphus, described by Dunn

(1931). The type and paratype series are in the

National Museum of Natural History, and I have

compared them directly with the series of longirostris available from Ecuador. In addition to the

general similarity in color pattern, both species

have rather slender bodies, elongate, thin limbs,

and a comparatively prominent, overhanging

snout. It is possible that the relationships of

glyphus are not with varius at all, and it should be

regarded as a distinct species. The two paratypes

from Porto Obaldia, noted as quite different by

Dunn (1931:396), are very reminiscent of A. varius

loomisi Taylor (1952).

I have examined the holotype of Phryniscus

boussingaulti Thominot, and see no reason not to

continue its synonymy with longirostris. The specimen is badly faded dorsally, retaining no trace of

pattern. There is a series of clear, rounded, whitish spots laterally, which, although not typical of

longirostris, is almost duplicated in some of my

specimens (e.g., USNM 193833). Photographs of

the holotype were taken and are on deposit at the

National Museum of Natural History.

REMARKS.—The type locality of this species,

given as the "Valley of Quito" by Cope, is open

to the same question as for the other four species

described from the same place and collected by

the Orton Expedition. Although one of these species (Leimadophis alticolus) has been found since

in the immediate environs of Quito, the other three

(Bufo andianus, Dryadophis pulchriceps, and

Dipsas oreas) are clearly western slope species, usually at altitudes well below that of Quito. The

road to Nanegal, on the western slope, is an easy

one to travel, and a three-hour drive today puts

one into the range of all of the species. I believe
NUMBER 145 29

that a local collector brought into Quito the material that eventually served as the types of these

species for Cope, just as today several local collectors still find valuable and interesting material

around Mindo, Tandayapa, and Nanegal.

This problem is complicated by the fact that Orton (1871) indicated that the type came from Hacienda Antisana, which he said was at an altitude

of 13,300 feet. This locality, according to Cope

(1868:96) is on the western slope of Volcan Antisana. Cope indicated that the total collection at

this locality was "a small black frog," a description more appropriate to A. ignescens than longirostris. I think we can safely assume that Orton

was depending upon his memory when he indicated the type locality, and in this case it was

faulty. It is of additional interest to note that the

USNM catalogs do not include any records of

specimens from Antisana collected by Orton, so it

appears that the "small black frog" from there was

deemed too insignificant to catalog.

A series of specimens collected by P. Hershkovitz in 1935 at Carolina, Imbabura Province, at

900 meters, is tentatively assigned to this taxon.

In general appearance and morphology they closely

resemble longirostris, but they are all quite small.

The difficulty in calling them longirostris is caused

by their color pattern. They are quite similar to

each other, in the occurrence of many more

rounded and more closely approximated yellow

spots dorsally than is typical of longirostris, and

the large, elongate spot immediately behind the

head dorsolaterally, so characteristic of longirostris,

is absent in all four specimens, as is the lighter

coloration of the projecting snout. There is a

more or less distinct, continuous yellow dorsolateral line on the specimens, while in longirostris

the same area is occupied by a discontinuous series

of spots. The specimens are the northernmost record for the species as I define it, and may show

incipient divergence from the parent taxon. The

possibility that the specimens represented the taxon

called Atelopus longirostris marmorata by Werner

occurred to me, but the dorsal pattern cannot be

called "marmorate," in my opinion, although it is

closer to that than are typical longirostris, and I

have considered Werner's taxon unidentifiable (see

elsewhere, under incertae sedis).

The large series of specimens available from

Ecuador provides a sufficient basis to demonstrate

that Rivero (1963:112) was correct when he indicated that the Colombian specimens he tentatively

assigned to longirostris (CNHM 43850-51) probably would not prove to be conspecific with that

species. Part of his doubt came from the difference

in known altitudes for Quito and Cisneros, in

Colombia, but I have shown above that Quito is

not a valid locality for longirostris. The series of

specimens from Santander called longirostris by

Cochran and Goin (1970:143) unequivocally do

not belong to that taxon. Therefore, longirostris

is not known from Colombia at the present time,

but is confined to the lower western slopes of the

Andes in Ecuador.

A series of specimens in the American Museum

(AMNH 49943-46, 49948-50) are recorded as having been collected at "Mera, Rio Pastaza," which

is a town on the principal road descending from

the high Andes to lowland Amazonia. This is

such a zoogeographic unlikelihood that I challenge

the validity of the original data rather than record

the occurrence of the species on the Amazonian

slope. The records show that the specimens were

purchased from a professional collector, and the

history of Ecuadorian herpetology is replete with

erroneous records based on such purchased material. The specimens are in my opinion clearly

members of the species longirostris, and I am sure

that they came from the lower western slopes of

the Andes in Ecuador. In addition to the fact that

the locality does not make zoogeographic sense,

it should be pointed out that collectors have been

delivering specimens to European, United States,

and Latin American collections from the Mera

region for over 200 years, since it lies on one of

the main trails to the east, but there are no records

of other specimens of this species from the area.

I do not include this locality as part of the known

range of the species until verification of its occurrence there by further collections has been made.

SPECIMENS EXAMINED.—ECUADOR (PM 207,

holotype of boussingaulti): Esmeraldas Province;

Alto Tambo (USNM 193853), Carondolet (USNM

193847-48). Imbabura Province; Apuela (USNM

193819-21, 193823); near Penaherrera (UMMZ

83662-66). Pichincha Province; Chontapamba

(MCZ 84203-04, USNM 193834, 193854-64),

Llambo, on road to Gualea (BMNH 1972.737,

USNM 193824-31), Mindo (UMMZ 55516, USNM

193832, 193841, 193866), 10 km northwest of Nane-
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

gal Chico (USNM 193817), Nanegal Chico

(UKMNH 108942-13, UPR-M 5090-91, USNM

193833, 193837-10), Rio Blanco (USNM 193846,

193852), Rio Mindo (USNM 193842-43), Rio

Toachi (USNM 193849), San Tadeo, near Mindo

(USNM 193865), Tandayapa (BMNH 1972.736,

CAS-SU 11388-89, USNM 193822, 193835-36,

193851-52). Province unknown: Miligali (USNM

193844-45).

Tentatively allocated to taxon: Ecuador: Imbabura Province, Carolina, .900 m (AMNH 50969,

UMMZ 83656, 83658-60).

Atelopus mindoensis, new species

FIGURE 22

HOLOTYPE.—USNM 193554, from Mindo, Pichincha Province, Ecuador, 1200 meters, collected by

Manuel Olalla, May 1959.

DIAGNOSIS.—A diminutive member of the longirostris group, most similar to A. palmatus, with

adult females usually shorter than 30 millimeters

in standard length. Males usually shorter than 20

millimeters. First digit on forefoot almost entirely

hidden in webbing; ventral surfaces unicolor, no

spotting, dorsum with a broad yellowish brown

band and reddish brown spotting and marbling.

DESCRIPTION.—Snout from above bluntly

pointed, with canthus forming straight line from

tip of snout to nasal eminence, from that point

to corner of eye it forms a gentle inward curve.

Snout and edge of canthus fleshy anterior to nasal

eminence, but no fleshiness on canthus from nostril

to eye, on eyelid, or on temporal region. Slight

depression in center of snout, as consequence of

fleshy margin; rest of dorsum of head flat. Snout

from side prominent, projecting over lower jaw;

nostril closer to tip of snout than eye, on raised

area and directed laterally; loreal region vertical,

gently rounded, no loreal depression anterior to

eye. Temporal region with scattered pustules and

spines, differing in number and prominence between individuals. Body liberally studded dorsally

FIGURE 22— Holotype, USNM 193554, of A. mindoensis, new species.
NUMBER 145 31

and laterally with tiny pustules widely scattered

but at least in part organized into tracts; scattered

larger, pointed, usually pigmentless, rounded pustules along dorsolateral line and laterally; limbs

also studded with tiny spicules, concentrated in

raised, fleshy areas along length of limbs.

Forelimb slender in females, shorter and heavier

in males, forefoot has somewhat fleshy webbing,

with first digit completely covered by it, second

digit included by a border of web; webbing deeply

incised between second and third and between

third and fourth digits, with fleshy nature of webbing making it appear absent between these three

digits. No subarticular tubercles on slightly fleshy

ventral surfaces of digits; plantar tubercle low,

rounded, almost indistinct.

Hind limb slender, heel reaches to points between posterior margin of eye and axilla when

hind limb is brought forward along body; heels

touch or slightly overlap when tibiofibulae are

parallel to femora at right angles to body. Webbing on foot extensive and somewhat fleshy, with

first toe almost completely indistinguishable within

it, second and third digits completely and continuously webbed; fourth toe webbed to tip, but webbing is hardly more than a border on ultimate

phalanx; full webbing on fifth digit. Ventral surface of foot is devoid of tubercles; fairly large,

raised, prominent, pigmentless outer metatarsal

tubercle present.

COLOR IN ALCOHOL.—Dorsal ground color rather

light yellowish brown, forming broad band length

of back, filled with reddish brown marbling, spotting, and vermiculation in younger individuals,

practically totally obscured in older individuals,

which tend to have back almost unicolor reddish

brown. Some individuals have irregular stripe from

eye to groin, bordered below by lighter stripe beginning at posterior margin of orbit and disappearing on sides. Sides are usually reddish brown,

sharply distinct ventrolaterally from belly color;

some specimens have series of clear, yellowish spots

on lower sides. Venter is clear light yellow, usually

quite transparent, with eggs visible in body cavity

of ripe females. Ventral surfaces of limbs often

completely invaded by dorsal color, but in some

individuals yellow of belly is continuous with yellow color of foot. No spotting of any sort on throat,

belly, or soles, except in one female with few light

brown spots on chin. Side of head dark brown, including upper lip; area below eye often with light

yellow crescent. Lower jaw entirely yellow.

HOLOTYPE.—The holotype conforms with the

description given above for the species, where no

variation is mentioned. Where alternatives appear

in the species description, the holotype has the

following characteristics: temporal region with

about six large, white-tipped pustules and many

scattered spicules. Heel reaches to the posterior

margin of the head when hind limb is brought forward; heels overlap slightly when tibiofibulae are

parallel to femora at right angles to body. Dorsum

almost unicolor, with irregular, somewhat lighter

area dorsolaterally that is vermiculated on its inner

margins. Clear yellowish spots present on lower

sides, some fused with light yellow ventral color.

Eggs visible in body cavity. Ventral surfaces of

limbs dark brown, with few scattered lighter spots.

Light crescentic area present below eye, lower lip

with dark brown margin, rest of chin and throat

clear. Measurements (in millimeters): snout to

vent: 29.0; snout to end of coccyx: 27.2; knee to

knee: 22.0; tibiofibula: 11.3; head length: 8.6; head

width: 7.7; eye diameter: 4.0; snout to eye: 5.2;

interorbital distance: 3.8; width of eyelid: 3.9.

ETYMOLOGY.—The species name is derived from

the name of the type locality, a most pleasant little

town where Beatriz Moisset and I spent several

delightful days in the field in 1965.

SPECIMENS EXAMINED.—The following are paratypes: ECUADOR: Pichincha Province; Milpe, 900

m (USNM 193562-63), Mindo (BMNH 1968.115,

UKMNH 108944-45, UMMZ 131684, USNM

193556-59), Rio Blanco, near mouth of Rio Yambi,

about 700 m (CAS 134204, USNM 193561), Rio

Lelia (AMNH 79462), region below Sigchos (MCZ

84200, USNM 193564), Rio Toachi(USNM 193555,

193560). The following are not designated as paratypes: ECUADOR: Esmeraldas Province; Carondolet (USNM 193570), Alto Tambo (BMNH

1968.116, USNM 193571-72, UPR-M 5092). Pichincha Province: Mindo (USNM 193565), region below Sigchos (USNM 193569), region of Gualea

(USNM 193566-68), Tandayapa (GOV 8696,

USNM 193573).
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Atelopus nepiozomus, new species

FIGURE 23

HOLOTYPE.—USNM 193543, female, from

slightly above Suro Rancho (also called Gualecenita), a single house on the trail between Limon

and Gualeceo, approximately 16 kilometers airline

west of Limon (which is now known as General

Plaza), Morona-Santiago Province, Ecuador, approximately 3000 meters, collected by James A.

Peters, August 23, 1962. See Figure 24 for map

showing locality.

DIAGNOSIS.—A member of the ignescens group,

very similar to ignescens, bomolochos, and arthuri,

differs from all three in presence of scattered dark

spots ventrally; dorsum with many low, rounded

warts and pustules which may be spiculate; dorsum

usually spotted and variegated with darker brown

on lighter ground color; also very similar to halihelos (for contrasting characteristics, see diagnosis

of halihelos).

DESCRIPTION.—Snout viewed from above roundly

pointed, with canthus forming slight inward curve

from tip of snout to nostril and then much deeper

inward curve from nostril to eye; canthus fleshy

and raised, with nostril below it on a lateral prominence; snout area between canthi slightly depressed. Upper eyelid fleshy, followed by fleshy

ridge from corner of eye to back of head. Frontal

and occipital areas flat, markedly lower than fleshy

margins of head. Snout viewed from side blunt,

slightly projecting beyond lower jaw; nostril closer

to tip of snout than eye, located on lateral eminence, directed laterally and slightly posteriorly,

loreal region distinctly concave. Area behind eye

to shoulder with raised, rounded warts, each of

which may have several white-topped spinules on

it. Dorsum and sides of body and legs with similar

low, rounded warts (more prominent in females

than in males), which may bear one to many whitish spinules or lack them entirely; generally more

and larger warts laterally than dorsally. Chin,

throat, belly, and lower surfaces of limbs usually

finely wrinkled, but devoid of warts and spinules.

Forelimb slender in females, heavier in males;

fingertips do not reach beyond body when leg is

carried back along body. Webbing very reduced

on forefoot, w

T

ith first digit clearly distinguishable;

webbing appears completely absent, but is actually

very fleshy between digits at base; greatest extent of

FIGURE 23—Holotype, USNM 193543, of A. nepiozomus, new species.
NUMBER 145

webbing between first and second digits, where it

includes about half of first digit. First digit quite

short, fourth longer than second, third much longer

than fourth. Many low, rounded, fleshy tubercles

on sole of forefoot and at digit articulations; outer

metatarsal tubercle larger than others, but not pronounced or raised.

Hind limbs short; tibiotarsal articulation reaches

to arm insertion or slightly farther when carried

forward along body; heels do not overlap when

tibiofibulae are parallel to femora at right angles

to body. Webbing extends to tips of all toes, but

is deeply incised between all except first and second, where it is complete and includes all of first

toe; webbing forms distinct border on all toes, narrowing toward tips, giving tapered appearance to

individual toe. Low, rounded tubercles at articulations and on sole; inner metatarsal tubercle low

and rounded as others, outer metatarsal tubercle

smaller but distinctly raised, pointed, and hard.

COLOR IN ALCOHOL.—Ground color of dorsum of

legs and body pinkish gray, with marbling, spotting, and variegations of reddish brown. Ground

color usually predominates, and in one individual

darker area is confined to thin pencil stripe of

color on vertebral column. Darker areas associated

with warty swellings, so that where wartiness is

heavier there is greater amount of darker marbling.

Sides of head dark brown, with distinct and sharp

color change at canthus and along dorsolateral

ridge of head. In few individuals a poorly defined

stripe of ground color from eye along dorsolateral

margin to groin, usually not present or indistinct.

Ventral surfaces of body yellowish to yellow-green,

with some spots or streaks of dark brown usually

present, most commonly on chin, throat, and chest;

where thin, midline streak may broaden into one

or several spots in tandem. Triangular, diaper-like

patch extending from anus onto ventral surface of

thighs. Dorsal surfaces of feet similar to body and

legs, but about half of specimens have first three

toes and first two fingers distinctly lighter than

rest of foot, in some becoming quite yellow. Ventral surfaces of feet usually unicolor light yellow,

but in some individuals may be dark spotted.

COLOR IN LIFE.—Dark greenish above with dark

brown spots. Ventrally orange on thighs, arm insertions, and chin. Belly yellowish. Spots on throat

and below anus very dark brown.

HOLOTYPE (Figure 23).—The holotype agrees

with the species description above in all characters

for which no variation is mentioned. For those

characters which vary, the holotype is as follows:

temporal and tympanic areas and sides of body

with white-tipped spinules on warts, those on body

not multiple on a single wart. Warts on legs lack

white tips, except very near arm and leg insertions. Tibiotarsal articulation reaches point of

insertion of forelimb; heels separated when tibiofibulae are parallel to femora at right angles to

body. No definable dark stripe from eye to groin,

whole side unicolor dark brown. Ventral surfaces

of feet with dark areas. Measurements (in millimeters): snout to vent: 32.4; snout to end of coccyx:

29.3; knee to knee: 24.9; tibiofibula: 11.5; head

length: 11.0; head width: 9.5; eye diameter: 4.8;

snout to eye: 6.9; interorbital distance: 5.2; width

of eyelid: 4.0.

REMARKS.—All of the specimens in die type series

were collected during our 1962 field work, which

was made up of a series of transects on the eastern

slopes of the Andes. Since the routes followed were

often only mule trails, it is not easy to identify

stopping places and collecting localities where we

worked during the week or so that a descent or

ascent might take. Figure 24 shows all of our collecting spots during the final transect, which took

us down one mule trail and back up a second a

litle farther south. Most of the names on the map

are "tambos," or single houses which provide overnight facilities for mule drivers, and cannot be

found on any other maps or in other gazetteers of

Ecuador. They are, however, comparatively permanent, because they occur at approximately one

"mule-day" intervals on the trail, and require a

good-sized flat area for a house and a corral. Only

when highways are open to towns such as General

Plaza, Macas, and others in the lowlands, will the

mule traffic stop, and these houses will revert to

the forest.

ETYMOLOGY.—The species name is derived from

the Greek word nepios, meaning "infant," and

zoma, referring to a girded garment or the position

of the girdle, i.e., the girded garment of an infant,

as close as one can come to "diaper" in Greek. It

refers to the patch of dark pigment that encloses

the anus, upper thighs, and the posteriormost part

of the venter of these frogs, which gives them the

look of a diapered baby.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

• •?< J I

SAN VICENTE^

! SAN, JUAN IBOSCOJ PLAN DE MILAGRO

—

FIGURE 24.—Route of collecting transect made in August 1962, by J. A. Peters and party,

showing localities along mule trails from whence material was obtained. Contour lines are

in meters. (Based on the Mapa Geografico de Ecuador, published by the Instituto Geografko

Militar, Quito, 1950.)

SPECIMENS EXAMINED.—The following are designated as paratypes: ECUADOR: Morona-Santiago

Province; Cerro Negro, 3450 m (USNM 193544),

near Pailas, circa 2150 m (USNM 193545), San

Juan Bosco, 2000 m (BMNH 1972.735, AMNH

79463, USNM 193546-47), San Vicente, circa 3450

m (UKMNH 108946, USNM 193549-51), Suro

Rancho, 3400 m (USNM 193548). The following

were examined but are not considered paratypes:

ECUADOR: Morona-Santiago Province; San Vicente, circa 3450 m (USNM 193552-53, JAP

7972-73).

Atelopus pachydermus (Schmidt)

FIGURE 25

Phirix pachydermus O. Schmidt, 1857:1") [type locality: "NeuGranada"]; 1858:257, pi. 3: fig. 26.

A.[telopus] pachydermus.—Rivero, 1963:107; 1968:23.

Atelopus pachydermus Rivero, 1965:138.

HOLOTYPE.—In the Zoological Museum of Cracow, Poland. Present condition not known.

Schmidt (1858) gave additional information concerning the type locality, stating it to be "Westen

von Neu-Granada, bei Bonaventura, in einer Hohe

von 5000'."

DESCRIPTION.—Snout from above blunt, rounded,

not projecting, distinct angle in canthus at nostril,

canthi from nostril to corner of eye form straight

line, angled slightly outward. Canthus and outer

eye margin slightly fleshy, with distinct ridge from

posterior corner of eye to end of head. Dorsum

of head flat, without ridges, tubercles, or fleshy

prominences; eye does not protrude dorsally. Snout

from side slightly protrusive beyond end of lower

jaw; nostril directed laterally, slightly below canthus; loreal region weakly concave; eye slightly

overhung by fleshy edge of eyelid; temporal region

without tubercles or spines, but in some individuals

with glandular swellings. Dorsum of body broad,

flat, with minute wrinkling and folding; in some

individuals scattered, rounded, glandular prominences. Sides of body studded with rounded,

glandular swellings, which are usually covered with
NUMBER 145 35

FIGURE 25.—A representative individual, USNM 193495, of A. pachydermus Schmidt.

several to many whitish spinules. Chin, chest, and

belly wrinkled or lined, but without pustules.

Forelimb short, stout in males, more slender in

females, forearm more fleshy than upper arm in

both sexes; dorsum of arm with glandular swellings,

rest smooth with wrinkles or folds. When arm is

pulled back along body, digits reach level of hind

leg perpendicular to body in females, not so far

in males. Forefoot slightly fleshy, webbing very

reduced, with all digits distinct and free; first digit

more swollen than others, wth horny excrescence

from tip of digit to base in males, covering a raised

hump of flesh, second digit with cornified area near

base of digit. Subarticular tubercles absent, but

palm with many raised glandular areas, often with

pair at base of each of second and third digits;

large palmar tubercle, less prominent plantar

tubercle. Tips of digits slightly swollen.

Hind limb rather stout, occasionally glandular

along femora, more often heavily wrinkled or

folded. Tibiotarsal articulation reaches to points

between end of head and arm insertion when carried forward along body; heels do not overlap when

tibiofibulae are parallel to femora at right angles

to body. Entire foot with somewhat fleshy web,

covering all but tips of first and second toes; extends to tips of other digits, but is deeply incised

between them. No subarticular tubercles, no tubercles on sole, outer metatarsal tubercle low, rounded,

somewhat indistinct, inner metatarsal tubercle

ovate and low.

COLOR IN ALCOHOL.—The typical pattern for this

species is shown in Figure 25. It may be described

in general as: ground color or entire body creamyellow, with irregular black vermiculations and

spots dorsally, which may be so heavy as to cover

head and back completely. Sides, venter, and legs

also spotted, stippled, and vermiculated with black,

with belly and undersides of limbs less spotted

than elsewhere. Color patterns extremely variable.

COLOR IN LIFE.—The lighter dorsal areas are

bright yellow. Many of the warts and pustules have

white tops. The ventral surfaces are white in almost all individuals, but some may have orange
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

areas or spots, and some have red bellies, similar

to those seen in specimens of ignescens from nearby

localities.

HABITAT.—This high altitude species is extremely common in the vicinity of Papallacta,

where they were found along the borders of the Rio

Papallacta, under rocks and stones in moist, spongy,

runoff areas.

VARIATION.—All species of Atelopus tend to be

variable in coloration, but the available material

of A. pachydermus shows a striking amount of

variability. Figures 26 and 27 show a series of adults

from the immediate environs of Papallacta, arranged to show the decrease in amount of light

pigment in the series. While no individuals were

taken in the region of Papallacta which lack the

dark pigment entirely, some of the specimens from

the region of Oyacachi are apparently entirely

free of any dark spotting and mottling, and in life

must present a striking golden appearance. In

addition to the variation in the amount of pigment

present in adults, there is a distinct ontogenetic

color change. Individuals 27 millimeters or less in

body length lack the strong contrast between the

dark and light areas seen in adults. The dorsal areas

that presumably will be yellow in adults are dark

gray or light brown in juveniles, and the darker

areas are not strongly differentiated from the

lighter. Some juveniles appear almost totally black

dorsally. Ventral color in the juveniles is the same

in adults.

REMARKS.—Rivero (1965:138; 1968:23) has discussed the use of the name Atelopus pachydermus

(Schmidt) for the population described here. He

examined the AMNH material from Oyacachi, and

tentatively assigned it to this taxon, and I have

followed him in this action. As Rivero pointed

out, however, there are problems involving the type

locality of Schmidt's taxon. The holotype of pachydermus was said to have come from above Buenaventura, at 5000 feet, presumably in the province

of Valle, on the western slope in Colombia. All

FIGURE 26.—A scries of specimens from the environs of Papallacta selected to show the variability

in dorsal pattern and distribution of colors in A. pachydermus. Numbers refer to field series, and

all have been recataloged.
NUMBER 145 37

of my material is from the eastern slope in Ecuador,

almost entirely at altitudes considerably above that

of the type locality. I consider it most unlikely that

any Atelopus species is capable of maintaining a

genetic continuum between these two areas, and if

the existence of a species answering the description

of pachydermus in the region of the type locality

is verified, my material will probably need a new"

name. I continue to use the name pachydermus

here for two reasons. First, the species has never

again been taken at or near the type locality, nor

has anything resembling it shown up there. It is

entirely possible that the type may have come

from the eastern slope, not the western, of Colombia, with either an erroneous locality given in the

description by Schmidt, or, as Rivero (1968:23)

suggested, there is a village called Buenaventura

on the eastern slopes of Colombia. Second, my specimens do fit the original description fairly well,

and the series includes specimens very similar to

the holotype in coloration. They are found at

comparatively high altitudes in northern Ecuador,

and there seems to be a good likelihood that the

same species could range into Colombia. The name

is certainly appropriate to my specimens, and I use

it for them temporarily, until further evidence

accumulates on the taxon.

A series of specimens from the collections of

Gustavo Orces-V. (GOV 8698-700) are recorded

as collected at Loja, Ecuador, but I am sure this

is an error in the transcription of the data, perhaps made by me when transferring that collection

to the National Museum. The series is unicolor

light golden, and is identical with material from

Oyacachi. I do not believe the same taxon, and

the same striking variation in pattern, also occurs

in Loja, and I have not included that locality in

the following list.

SPECIMENS EXAMINED.—ECUADOR: Napo Province; 2 km E of Chalpi, 2755 m (USNM 193520-

21), Oyacachi, 3300 m (AMNH 20505-21, BMNH

1972.732-733, CAS-SU 10348-50, 134200, JAP 3791,

MCZ 84201-02, USNM 193494, 193522-33), Pueblo

Viejo, Oyacachi (AMNH 20492-93), Papallacta

and environs, 3150 m (KU 117876-79, 142950-56,

UMMZ 131020-21, UPR-M 5088-89, USNM

193495-518).

FIGURE 27.—Ventral view of same series as in Figure 26.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Atelopus palmatus Andersson

FIGURE 28

Atelopus palmatus Andersson, 1945:12, fig. 4 [type locality:

"Rio Pastaza, Ecuador. 1000 to"].—Rivera, 1963:110; 1968:

21; 1969:142.

SYNTYPES.—Stockholm Royal Museum number

1908, 5 individuals.

DESCRIPTION.—Snout from above with pointed,

projecting tip; canthus forms straight line from

snout tip to level of nostril and then forms inward

curve from nostril to corner of eye. Snout, canthus, and eyelid raised, but weakly fleshy; bony

ridge from posterior corner of eye to corner of head,

ending in raised boss. Dorsum of head finely

shagreened, forming a plane surface, eyes slightly

raised above plane. Snout from side protrudes well

beyond both upper lip and lower jaw, may be

slightly concave laterally, and may curve slightly

downward at tip, giving "hooked" appearance.

Nostril lateral, slightly below canthus, loreal region slightly concave. Vertical ridge joins dorsolateral ridge behind eye. Side of head shagreened,

without raised tubercles or glandular areas. Dorsum

and sides of body lightly shagreened; vertebral

column forms indistinct ridge. Chin and belly

finely shagreened or lightly wrinkled and folded.

Forelimb long, slender; forearm heavier than

humeral region, dorsum lightly studded with small

pustules or finely shagreened, ventral surfaces

smooth or finely wrinkled. Forefoot with webbing

enclosing all but tip of first toe, weakly developed

on other digits. Subarticular tubercles absent or

very weakly developed, palmar tubercle large,

prominent, no plantar tubercle. Edge of prepollex

with horny excrescence in males.

Hind limb slender, lightly pustulate along dorsum of upper leg, smooth elsewhere. Tibiotarsal

articulation reaches to points between nostril and

anterior border of eye when carried forward along

body; heels touch or overlap when tibiofibulae are

parallel to fermora held at right angles to body.

Webbing extends to tips of toes, with first and

second digits almost totally obscured; webbing

rather deeply excised between other toes, reaches

FIGUXE 28—A representative individual, FNHM 36657, of A. palmatus Andersson.
NUMBER 145 39

tip of fourth toe as a fringe. Subarticular tubercles

weak and occasionally indistinct, outer metatarsal

tubercle prominent, inner metatarsal tubercle

absent.

COLOR IN ALCOHOL. — Dorsal ground color

chocolate-brown, dorsum of head spotted and variegated in light gray, usually with dorsolateral streak

or series of spots of same color from eye to groin,

scattered spots with irregular margins on back.

All light gray areas bordered with narrow dark

brown to black margin. Ground color extends down

side of head and body about halfway, with lips,

shoulder, and lower sides as well as all ventral

surfaces unicolor light brown to yellow. Limbs as

body dorsally and ventrally; dorsum of first digits

on both fore and hind foot usually yellowish, with

no brown pigment dorsally.

COLOR IN LIFE.—"Dorsal surfaces dark brown

with bright green spots. Flanks orange-yellow.

Ventral surfaces, including inner 2 digits of foreand hindlimbs, deep yellow. Groin bright orangeyellow. Iris black with minute greenish yellow

flecks and ring around pupil." (Taken from field

notes on KU 122544, made by W. E. Duellman.)

REMARKS.—Rivero (1969:142) has examined the

syntypes of A. palmatus, and he indicated that the

species was valid, but closely resembled A. planispinus, a larger species from the slopes of Mt.

Sumaco. I agree with this conclusion and, as indicated by the list below, have found a good series

of specimens in the various museums. Rivero

mentioned that the posterior margin of the frontoparietal and the upper margin of the suprascapulas

are raised and prominent in some of the syntypes,

but this is not evident in the series I have examined

except in individuals either dried or preserved in

a solution with too high an alcohol content.

Little is known about the ecology of this species.

A specimen collected by W. E. Duellman was taken

on the margin of a small stream during daylight

hours.

SPECIMENS EXAMINED.—ECUADOR: Napo Province; El Chaco, Upper Rio Quijos, 1454 m (USNM

193541-42). Pastaza Province; Abitagua, 1200 m

(UMMZ 90367), Baiios (FMNH 28079), Hacienda

San Francisco, on Bafios-Mera trail (FMNH 36657,

152136), Mapoto (FMNH 36687), Puyo (FMNH

28078), Sarajacu (AMNH 33891-92), southern slope

of Cordillera Du£ above Rfo Coca, 1150 m

(UKMNH 122544), Rio Azuela, 1740 m (UKMNH

142957-64).

Atelopus planispinus Espada

FIGURE 29

Atelopus planispina Espada, 1875:148, pi. 2: figs. 1-2 [type

locality: "San Jose de Moti, Ecuador"].

Atelopus longirrostris [sic].—Espada, 1875: pi. 2: fig. 1 [see

below, in "Remarks"].

Phryniscus cruciger.—Boulenger, 1882a: 154 [a questioned synonymy].

A.[telopus] cruciger Nieden, 1926:84 [a questioned synonymy].

A.[tetopus] planispinus.—Rivero, 1963:107.

SYNTYPES.—Twenty-eight specimens, MM 212,

Museo Nacional de Ciencias Naturales, Madrid,

Spain.

DESCRIPTION.—Snout from above with rounded,

protrusive tip, with sharp angle at nostril; canthus

rostralis from nostril to angle of eye forms straight

line angling outward from anterior to posterior;

no heavy fleshy margin on canthus. Dorsum of

head flat, in same plane as broad, flat dorsum of

body; bony ridge from corner of eye to end of

skull above tympanum, ending in boss; head

shagreened with tiny spines, heavier on eyelids.

Snout from side overhangs lower jaw, ending in

heavy point, which may be bent somewhat downward, i.e., "hooked," nostril laterally directed,

below canthus, loreal region vertical, does not form

concave area. Eye slightly overhung by moderately

fleshy margin of eyelid. Upper lip and head behind

eye spinose and pustulose. Entire body finely

speckled with tiny spinules visible under high

power of microscope; on sides and in groin spines

are much more prominent and pustules are also

present.

Forelimb slender, elongate, shagreened with

spicules and spines. When forelimb is pulled back

along body, palm reaches vent. Hand fleshy; first

digit enclosed in fleshiness except for its tip; other

digits free or with slightly fleshy border, not

webbed. Second digit shorter than fourth. Subarticular tubercles absent or very ill defined, larger

palmar tubercle, no tubercle at all at base of first

digit.

Hind limb slender, covered with spicules and

scattered spines, not pustulate. Tibiotarsal articulation reaches to points between nostril and pos-
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 29. A representative individual, AMNH 22198, of A. planispinus Espada.

terior margin o£ eye when carried forward along

body. Heels overlap when tibiofibulae are parallel

to femora at right angles to body. Entire foot

fleshy, webbing almost entirely encloses first and

second toes; webbing extends to tip of other toes

as fringe, but is deeply incised between them. Subarticular tubercles weakly defined, outer metatarsal

tubercle forms rounded hump, inner metatarsal

tubercle obscured by fleshiness of first toe.

COLOR IN ALCOHOL.—Dorsal ground color light

tan, with highly vermiculated markings of light to

dark brown. Dorsolateral region usually solid

ground color with scattered small dark spots, rest

of dorsum very irregular and sometimes so heavily

marked with dark brown that ground color forms

isolated spots. Lower sides, chin, chest, and venter

usually without spots, yellowish white (one juvenile has brown markings on chin and throat).

Ground color of limbs tan, with dark brown bars

or vermiculations which do not extend onto

yellowish white lower surfaces of limb. Outer

margins of soles of feet dark brown, inner parts

yellowish white.

SYNTYPES.—The 28 syntypes in the Madrid Musum are badly faded, and a few appear to have

been dried. Most of them have lost all signs of

pattern, but one small male still retains most of

its color, and it is possible to see enough parts of

the pattern on other individuals to be sure that

the more recent specimens identified here as planispinus belong to the same species. Insofar as they

can be determined on the type series, the characters given above for the species as a whole are

accurate for the types. Characters such as spicules

and spines, or degree of fleshiness, could not be

determined on the syntype.

REMARKS.—The type locality for this species,

given as "San Jos£ de Mod" by Espada, is apparently the town in Napo Province now called San

Jose Nuevo. The United States Board on Geographic Names in their volume on Ecuador (1957:

155) gave a reference to San Jose de Mote, with a

cross reference to San Jose" Nuevo, at 0°26'S,

77°20/W, which is on the lower northwestern slopes

of Mount Sumaco, the only place from whence I

have seen specimens. The town itself is quite low,
NUMBER 145 41

and it is unlikely that the type series originated

there. I suspect the 28 specimens came from the

slopes of Mount Sumaco, as do the AMNH specimens. I doubt also the reference to the "top" of

Sumaco for AMNH specimens, since that is at 3900

meters, an altitude higher than any other species

of Atelopus in Ecuador except ignescens, and much

higher than any other species in the longirostris

group, which, as pointed out above, tends to be

found at lower, warmer altitudes.

Espada (1875, pi. 2: figs. 1, 2) illustrated what

he identified in the legends for the Plate 2 varieties

of A. longirrostris (sic), but did not present a figure

of his own new taxon, "A. planispina." If one reads

the section termed "Observ." on page 154 of

Espada's text, one discovers that Espada considered longirostris and planispinus to be closely related, and in fact he thought of the two samples

he had as variants of longirostris for some time,

until he decided that "son verdadermente dos

especes, muy afines si, pero distantas." Since his

figure 1 on Plate 2 is the same taxon I am calling

planispinus on the basis of the AMNH specimens

from Mount Sumaco, I believe that the plate legend

did not get changed when Espada decided to give

the new taxon the name "planispina" rather than

to continue to regard it as a variety of longirostris.

The species shown in Plate 2; figure 2, in Espada,

is probably actually A. longirostris, although the

figure is not detailed enough to be certain. Espada

did have specimens of longirostris from Nanegal in

his collection (1875:159).

This species and A. halihelos are apparently restricted tc isolated mountain masses to the east

of the main Andean chain in Ecuador. It is unlikely

that either of them are found in the lowland areas

surrounding these masses, because other wideranging species of Atelopus are found there.

SPECIMENS EXAMINED.—ECUADOR: Napo Province; San Jose de Moti (MM 212-syntypes), Top of

Sumaco Mountain (AMNH 22193, 22195-97,

22199-200, USNM 193960), side of Sumaco Mountain (AMNH 22300-301).

Atelopus pulcher pulcher (Boulenger)

FIGURE 30

Phryniscus pulcher Boulenger, 1882:154, pi. IS: fig. 2 [type

locality: "Chyavetas, Peru" perhaps the same as Chayahuitas, Peru, according to Rivero, 1968:19)].

I -•'

FIGURE 30.—A syntype, BMNH 1947.2.14.80, of A. pulcher Boulenger.
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Atelopus pulcher.—Boulenger, 1894:375.—Peracca, 1904:22.—

Rivero, 1963:107.

Atetopus spumarius.—Rivero, 1968:19.

Atelopus spumarius spumarius.—Duellman and Lynch, 1969:

232.

SYNTYPES.—Four individuals in the British Museum (Natural History), and USNM 193574.

DESCRIPTION.—Snout from above rounded from

nostril to nostril, canthus forms a straight line from

nostril to eye, little or no fleshy area on snout

region or on canthus. Dorsum of head flat, smooth,

slight knob at corner of skull. Snout from side very

slightly protrusive beyond upper lip and lower

jaw; nostril directed laterally. Loreal region vertical; upper eyelid slightly fleshy, overhangs eye;

upper lip not fleshy; temporal and tympanic areas

smooth or very lightly wrinkled and shagreened.

No tubercular areas on head, body, or limbs.

Dorsum and sides of body lightly wrinkled, belly

wrinkling somewhat more pronounced.

Forelimb slender and slightly wrinkled, forearm usually thicker than humeral region. When

lorelimb is drawn back along body, palm reaches

thigh. Palm of hand fleshy; first digit distinct, with

tip of digit free of fleshy webbing; other digits entirely free of webbing; subarticular tubercles very

indistinct; palmar tubercle very large, rounded.

Hind limb slender, smooth or finely wrinkled,

under surfaces of thighs below anus with markedly

wrinkled and somewhat pustulate area, usually

distinctively colored. Tibiotarsal articulation

reaches points between anterior and posterior margins of eye; heels touch or overlap when tibiofibulae are parallel to femora at right angles to

body. Entire foot somewhat fleshy, with first digit

entirely hidden in webbing, second with only its

tip protrusive, other three digits webbed to tip, on

fourth chiefly as fringe. Subarticular swellings present; outer metatarsal tubercle present but small.

COLOR IN ALCOHOL.—Dorsal ground color greenish yellow, with dark brown vermiculations and

spotting more or less symmetrically distributed on

head, back and sides; upper lip yellow, with a fine

brown line along margin. Belly and chin yellow,

with small rounded brown spots primarily located

posteriorly. Limbs barred above with dark brown,

yellow below with brown spotting. Soles of feet and

area just below anus flesh colored.

COLOR IN LIFE.—Dorsal ground color yellowish

green, with jet black spotting and striping. Soles

of feet bright red, with area on venter that is

flesh colored in alcohol also bright red. In some

individuals the entire ventral surface is red.

REMARKS.—Rivero (1968:19) discussed at length

the two taxa A. pulcher Boulenger and A. spumarius Cope, and finally concluded, with doubts

based on Cope's inadequate description of his type,

that they were synonymous. I have not accepted

this action in my work for the following reasons.

First, A. pulcher Boulenger is a well-defined, easily

recognized species, with a good figure accompanying the original description, and with type material

available for comparisons. As a consequence, the

name has been used and the species recognized by

all workers in eastern Peru and Ecuador since

Boulenger's Catalogues were published. Second,

Cope's species is inadequately described; no figure

was published to show what in the world he meant

when he wrote about "aggregated annuli of greenish yellow . . . the appearance of green foam," and

the holotype has never been located. The substitution of a poorly described, inadequately characterized taxonomic name for a well-described, clearly

defined one with type material available, and which

has been in wide use for 80 years, seems to me to

be an inadvisable taxonomic practice, and I hope

we can return herewith to the widely recognized

name with only a minimum of use of the substitution. This does not seem to me to require more

than a recognition by herpetologists that A.

spumarius is clearly a dubious name until such a

time as the holotype is rediscovered, and, as such,

should not be applied to a known species until

the doubts are resolved. This would require that

the subspecies described by Rivero (1968:23) be

known as Atelopus pulcher andinus Rivero.

Rivero has also discussed (1968:20) my views

that high altitude records for this species are undoubtedly incorrect. He pointed out that he could

recognize differences between specimens supposedly from Riobamba, a city in the inter-Andean

highlands at 2800 meters, and the lowland material

from the Upano Basin. Savage (1960:14) has discussed the validity of Riobamba as the source of

certain AMNH specimens, and I follow him in

not accepting Riobamba as a valid locality for any

lowland species until verification in the form of

newly collected specimens has been made. The

AMNH specimens were received from H. Feyer,

and his "Riobamba" records of lowland species
NUMBER 145 43

have been causing problems in Ecuadorian herpetology for years. It seems clear that Feyer, with

his residence in Riobamba, purchased specimens

from any seller, including mule drivers coming up

from the Amazonian lowlands on their regular

trips, carrying a few frogs and snakes along to sell

to the "gringo," who then used "Riobamba" as

the origin of the specimens.

The specimens mentioned by Rivero (1968:20)

from Chancha, Chimborazo Province, are from

Normandia, in Morona-Santiago Province. This

locality is discussed by Savage (1960:13), who located it at 1300 meters on the trail between Macas

and Riobamba. This is within the known range of

the species.

Duellman and Lynch (1969:232) described and

figured a single presumed tadpole of this species,

which was collected in the Rio Puyo in 1969. They

followed Rivero in the use of spumarius for the

species, as did McDiarmid (1971:39).

The species Atelopus tricolor Boulenger (1902:

397) is close to A. pulcher in its characters. I have

had an opportunity to examine the syntypes, however, and they appear to me to represent a distinct

taxon. One of the syntypes is now in the USNM

collection, through the kindness of the British

Museum (Natural History) (USNM 193575).

Rivero (1968:19) has discussed the spelling of

the type locality in detail, and concluded that it

was very likely that the Peruvian village called

Chayahuitas, at 5°5'S, 76°10'W, was the original

source of the type material. Since Chyavetas has

never been located, his argument has considerable

validity, and I have indicated that the village he

located is probably the legitimate type locality.

SPECIMENS EXAMINED.—ECUADOR: (AMNH

17433-38, 17447). Morona-Santiago Province; Chancha, Normandia (AMNH 16695-712), Cutucii Cordillera (AMNH 33913-15), Macas (AMNH 21492-

94, USNM 65487-90), Miazal (GOV S940, JAP

3843), Limon (CAS 134202-03, JAP 6966, 7520,

UMMZ 131022-23, USNM 193479-90). Pastaza

Province; "Bafios and Canelos" (AMNH 16713-

15), Canelos (MCZ 17927-28), Puyo (FMNH

28078), 2l/2 km southeast of Puyo (JAP 1937-38),

Rio Oglan (GOV S939), Veracruz (JAP 6105,

UKMNH 120480-85).

PERU: Chayahuitas (BMNH 1947.2.14.80,

USNM 193574, syntypes of A. pulcher).

Present Position of Taxa Formerly Assigned to

Atelopus In Ecuador

There is a single species originally described in

the genus Atelopus from Ecuador that is no longer

considered to belong to that genus. Atelopus festae

Peracca is now considered to belong to the genus

Rhamphophryne (Trueb, 1971:22). A second species, Atelopus minutus Melin, although not originally described from Ecuador, has been collected

in the country several times. It was recently transferred to the genus Dendrophryniscus by McDiarmid (1971:40).

Incertae Sedis

Atelopus longirostris marmorata Werner

This taxon was described by Werner in 1910,

page 600, from "Ecuador," without more detailed

information on the type locality. He noted that the

females in his collection appeared different from

the males of longirostris, and used the trinomen

above for them. Nieden (1926:81) included the

taxon in the synonymy of Atelopus longirostris, and

the few citations since that time (Rivero, 1963;

Cochran and Goin, 1970) accepted Nieden's action

as having done away with the taxon. Careful examination of all of Nieden's work in this volume of

Das Tierreich, however, shows that he included

all trinomials in existence prior to his work as

synonyms of a binomial, and then mentioned those

he considered to be recognizable in the text below

the synonymy I find only three subspecies mentioned in the entire volume, all in the text material.

One of these is marmorata. It is, therefore, not particularly valid to cite Nieden as the authority for the

synonymy of marmorata with longirostris. I think

it entirely possible that Werner had specimens of

a completely different species in hand. Since elegans occurs on the western slopes in Ecuador, and

may actually be sympatric with longirostris in part

of its range, I would suggest that Werner may not

have recognized specimens of elegans among his

longirostris material. Unfortunately, the type

specimens were destroyed during World War II,

and marmorata becomes a nomen dubium.
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Zoogeography and Evolution

Rivero (1963:112) has remarked, "Frogs of the

genus Atelopus seem to be very sensitive to environmental factors." This is certainly true of

Ecuadorian species, particularly at altitudes above

the subtropical zone. The Ecuadorian terrain

makes exploration by herpetologists difficult, and

not too much work has been done on the outward

facing slopes of the Andes as a consequence. Access

to areas between 1500 and 3300 meters has always

been and still generally is available only on foot

or horseback, although road construction on both

east and west slopes continues to make more areas

accessible by car. Most of my personal collecting

efforts in Ecuador have been focussed on these

slopes. I have spent comparatively little time in

the Pacific lowlands, and even less in the Amazonian rain forest. Herpetologically, the latter has been

the most popular and thoroughly explored area in

the country, and, while it continues to produce

both novelties and surprises, it still does not hold

the same promise for investigation of ecological and

evolutionary phenomena as do the Andean slopes.

In 1958-1959 I made a series of seven transects

from the inter-Andean valleys to the Pacific lowlands, and in 1962 my group made five transects on

the eastern slopes, usually, although not always,

penetrating to the Amazonian lowlands. Such transects perforce follow available routes, and of the

total of twelve transects, five were by mule and

on foot, six were by road in truck or Landrover,

and one was by rail. Since the routes taken were

either mule trails several hundred years old, or

roads that follow such old trails and replace them,

it is to be expected that each trail would be separated by some distance and usually some fairly substantial physical or physiographic barrier from the

next trail, since arrieros follow paths of least resistance, and do not develop and follow duplicate

trails ending at the same place very often (except

as smugglers). The physiographic barriers are deep

river valleys, sheer cliffs, or too heavily dissected

slopes, all of which can and do act as biological

barriers to species movement equally well. In addition, rain shadow slopes, where the vegetation and

moisture available change radically within a few

thousand feet, occur on both east and west slopes,

and represent a major biotic barrier to horizontal

migration of mountainside species. Since these various types of barriers are common, and are most

often encountered between the traditionally used

trails, one would expect each transect to produce

different taxa, each evolving in an isolation produced by the barrier of altitude on one hand, and

the many horizontal barriers mentioned above on

the other hand. This proved to be particularly true

of frogs of the genus Atelopus and lizards of the

genus Proctoporus. Since representatives of both

genera are fairly often found even if the total collecting time available is slight, they become good

indicators of evolutionary patterns, effects of isolation, adaptive radiation, and other phenomena

often associated with the study of insular faunas. In

point of fact, mountains such as the Andes of

Ecuador can be considered "continental islands,"

and many of the phenomena long discussed by

students of insular biology are equally observable

there, with the added advantage that the mountains

are not dependent upon waifs and drifters for replenishment of the supply of evolutionary materials.

The taxa in the genus Atelopus lend themselves

to the study of these phenomena because of the sensitivity to environmental factors already noted by

Rivero. But, although they are sensitive to, and

often limited by, very slight environmental

changes, they are also capable of adaptation and divergence, which permits them to conquer new

environments. Suitable habitat for the taxa in the

lowlands o.f Amazonian Ecuador as well as in the

lower altitudes on the Pacific side appears to be

continuous, with few barriers to gene flow, and

lowland species tend to have a considerable latitudinal range. A. pulcher ranges throughout the

southern half of Amazonian Ecuador and a considerable distance in Peru, from whence it was originally described. It stops at about the Rio Pastaza,

and is replaced by A. palmatus, the northern limit

of which is currently unknown. A. elegans is a

lowland species on the west side of the Andes in

Ecuador, and is found north well into Colombia

and south to the point where the wet areas are

replaced by the dry coastal scrub, under the influence of the Humboldt Current. Figure 31 is an

attempt to show how the situation changes when

the species found on the slopes at slightly higher

altitudes are examined, and the opportunities for

horizontal migration are diminished by the multiplicity of barriers. The extremely narrow distribu-
NUMBER 145 45

tion of the species found at altitudes above 2000

meters is in part an artifact of the collecting opportunities, which are restricted to a few hundred

yards laterally to the trails, but it is fairly certain

that there is a high degree of endemism on the

higher slopes, and it is very likely that the endemic

species are totally allopatric. Finally, after an altitude sufficient to permit the inter-Andean highlands to be reached, the picture is reversed. Th e

highly successful species A. ignescens is capable of

living at very high altitudes under most inhospitable conditions. As a result, it is the most widespread amphibian species in the Andean highlands,

ranging from well into Colombia practically to the

Peruvian border. Th e ability to live in paramo

and subparamo climates has eliminated all physical barriers to movement by the species, and it is

ubiquitous over a considerable range. It is the only

species that may in several cases be sympatric with

those species with restricted ranges on the outer

slopes. I have found A. ignescens within 2 to 3

kilometers of populations of A. pachydermus, and

it appears that the same is true for A. bomolochos,

much farther to the south. A. ignescens is, of course,

sympatric with A. bufoniformis, which has been

taken as high as 3400 meters, and is known from

both slopes of the Andes, but only in the most

northern provinces of Ecuador. A. bufoniformis is

the end point in the evolutionary sequence discussed below.

While the evolutionary sequences are not entirely clear, and the evidence used here is primarily

morphological and zoogeographic and thus could

be easily disproven by data from cytological, gecc

Ul

4000

3500

3000

2500

2000

1500

1000

500

0

K -r

V

G

f- INTO^-

.1 I

- PERU **-

1

-•INTO

- • COLOMBIA

4--I7

INTO

- • COLOMBIA

5°S 4°S 3°S 2°S

LATITUDE

0*

A ARTHURI

B BALIOS

C BOMOLOCHOS

D BOULENGERI

E BUFONIFORMIS

F ELEGANS

G HALIHELOS

H IGNESCENS

I LONGIROSTRIS

J MINOOENSIS

K NEPIOZOMUS

L PACHYDERMUS

M PALMATUS

N PLANISPINUS

0 PULCHER

FIGURE 31.—A diagrammatic representation of the known distribution patterns of the species of

Atelopus in Ecuador. Each square (or line) is based on known extremes, and therefore may

well include areas from whence a particular species is not known. In each species a complete

rectangle is shown, although this is not easy to discern where two species have common limits.

Dashed lines indicate uncertainty, usually explained in the text.
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

netic, biochemical, and other more modern tests,

it still seems legitimate to construct hypotheses

concerning what has happened to Atelopus phylogeny in Ecuador. The potential absurdity and essential sterility of such hypotheses can only become

evident after considerable research on the taxa

concerned, research which I do not anticipate being done for many years—and which might even

verify my guesses here.

In any event, the evidence derived from an examination of the distribution and morphology of

the two groups of species indicates the direction

that evolution has taken. With regard to distribution, the taxa of the longirostris group are almost

entirely low altitude species. The only exception

may be A. planispinus, which is found on Mount

Sumaco, which reaches an altitude of 3900 meters.

The type locality for the species, at the foot of the

mountain, is at 500 meters, and specimens in the

American Museum of Natural History are recorded

as coming from the top of Sumaco, which gives the

species a possible altitudinal range of 500 to 3900

meters, a most unlikely situation. Since this taxon

is part of the longirostris group, and since it has

not been taken elsewhere in the Amazonian lowlands, as would be expected if it actually occurred

at San Jose Nuevo (the modern name for San

Jose de Moti, or Mote), I predict it will eventually be found to occupy the slopes of Sumaco

at lower to intermediate altitudes (perhaps 1000 to

2000 meters) and probably on the eastern slope

only (because of habitat change on the back or

western side of the mountain). Other than A.

planispinus, the taxa in the longirostris group do

not reach altitudes above 2000 meters, and enjoy

their greatest abundance below 1000 meters.

The taxa of the ignescens group, on the other

hand, are almost all high altitude forms. The one

exception, A. mindoensis, is an enigma. As I point

out frequently in this paper, this taxon is intermediate between the two groups in many ways.

For example, it tends to fall halfway between the

groups in almost all of the analyses of proportional

measurements. Leaving that species to one side,

the remaining members of the igenscens group all

show a common zoogeographic pattern. They are

very restricted horizontally, and appear to be

equally restricted vertically to a fairly narrow altitudinal band. They have not, and I believe they

are unable to, spread widely on a horizontal plane,

although the horizontal distribution of each is

certainly greater than my data show. The vertical

restrictions are for the most part real, and I think

the various taxa show sharp response to and are

closely confined by the changing conditions which

accompany increasing altitude. All of these species

are living in a comparatively young area, geologically speaking. The surrounding lowlands are

much older, and it seems plausible that the ancestral species in the genus Atelopus are the lowland taxa, and the highland taxa have been repeatedly derived from the lowland stocks. I would

further suggest that each of the highland taxa

(with the probable exception of A. ignescens and

the possible exception of A. bufoniformis) is the

product of a separate, successful invasion of the

highlands by lowland stock. This, of course, means

that the ignescens group is clearly not monophyletic, but simply represents a series of separately

derived species all differentiating from the ancestral

stock along similar morphological lines.

The morphology of the two groups of species

tends to verify the hypothesis. The adaptive

changes evolved in response to higher altitudes and

colder temperature include shorter, more stocky

bodies and a reduction of the limbs from long and

slender to short and stout. This suggests, of course,

the development of mechanisms to reduce total surface and concomitant heat loss. The highland species are much less likely than the lowland taxa

to be found out and actively moving about except

during breeding periods, when they become abundant.

While not as sharply distinctive between groups

as the body and limb configuration, the thickening

of the skin in the webbing between the digits, and

the overall fleshiness of the highland species in

general, may well be another modification toward

heat conservation. It reaches maximum development in A. bufoniformis, the taxon which has both

the maximum development of the characteristics

discussed here and reaches, with A. ignescens, maximum altitudes.

McDiarmid (1971:58) has discussed the evolutionary relationships of Atelopus with other genera

of the family Bufonidae, and demonstrated that

the genus is quite advanced rather than primitive

in a large number of its characteristics. He pointed

out several phenomena concerning the genus Ate-
NUMBER 145 47

lopus that are of interest in understanding evolutionary events in Ecuador. He stated:

With the initial uplifts of the Andes in Peru and Ecuador in

the Late Cretaceous ... . entire new areas became available

to and were exploited by Atelopus. The ancestral stock probably came from southeastern portions of the Guiana shield

where the more primitive Atelopus flavescens is found today.

However, it is possible that Atelopus initially invaded the

montane habitat in the southern Andes and moved north

along their eastern slope. It appears also that the higher

elevations have been invaded several times.

I would agree completely with the concept of

multiple invasion of the higher elevations, but

think it unlikely that there has been very much

movement north along the eastern slope of the

Andes from the south. The evidence presented

above seems to indicate fairly strongly that lateral

or horizontal movement is not a common phenomenon in Ecuador, probably as a consequence

of multiple barriers. McDiarmid (1971:58) has

himself provided an additional argument against

horizontal movement, for he said:

The apparent close association of species of Atelopus with

mountain streams has restricted species to particular drainage

systems. The resulting isolation (drainage system and mountain top) probably accounts for the striking differences among

many populations and probably has been an important factor

contributing to speciation.

The only species of the ignescens group which

has been able to break out of the limitations noted

above for other species is A. ignescens itself. It has

been a highly successful invader of the interAndean valleys in Ecuador, and its vertical range

is such that the nudos or knots between the valleys

are not a barrier to horizontal movement. The

species ranges from the southernmost provinces in

Ecuador well into Colombia, a horizontal range

almost as great as that of any lowland species.

Only A. bufoniformis shows greater differentiation

than A. ignescens along the morphological sequences to high altitude adaptation, as can be seen

in the graphs of proportional measurements. A.

ignescens represents an explosive movement taking

place, because its ancestral stock continued the upward expansion of its range until it spilled over the

wall of the Andes, opening up the whole interAndean plateau to settlement.

The success and widespread distribution of A.

ignescens leads one to a second look at the other

members of its group. It is entirely possible that

they represent repeated successful invasions of the

outer slopes by A. ignescens, adapting and differentiating in different areas. A. ignescens definitely

occurs on the eastern slope within a few kilometers

of A. pachydermus, and the relationship between

them seems to be close. The same is true for A.

bomolochos, which is equally close to A. ignescens

both geographically and morphologically. Why,

then, is it not possible that A. ignescens is the direct ancestor to these restricted populations?

Perhaps the strongest argument against this hypothetical ancestry is the fact that A. ignescens

tends to maintain its specific identity throughout

a considerable distance in the inter-Andean plateau. Although quite variable, and perhaps with

recognizeable subspecies in several places, it still

remains one species throughout, in my estimation.

It is uncertain how much contact there is between

populations separated by nudos, but I suspect there

is less contact between them than within such

populations, certainly a situation to encourage

local populational differentiation, if the species is

plastic enough to respond. I suggest that A. ignescens is a species well advanced evolutionary, and

highly adapted to a very inhospitable environment,

and therefore has comparatively little evolutionary

potential. The likelihood of such a species invading a series of isolated drainages on both sides of

the Andes of Ecuador and giving rise to such a

large number of very distinct species as are found

there seems very slight to me.

On the other hand, I must reverse that argument

with regard to A. bufoniformis. This species has

gone beyond A. ignescens in every one of the adaptive characteristics discussed here. It is clearly the

terminal species in the differentiation sequence.

It is difficult for me to visualize any other ancestor

for this species than A. ignescens, and this causes

some considerable discomfort. At one time I had

thought A. bufoniformis to be an inhabitant of

the high eastern slope, found slightly below the

range of A. ignescens. But material in the

UKMNH collections show it from 3400 meters, and

well into the higher parts of the inter-Andean Plateau. It does not seem to occur in the inter-Andean

valleys, although it has been found low enough on

the outer slopes that altitude would not be a barrier. It appears to have been a comparatively recent derivative of A. ignescens stock, and is perhaps

even more thoroughly adapted to the cold, wind-
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

swept, high altitude paramo than the parent stock.

It is probably in the process of expanding its range

at those high altitudes, but may face problems in

the fact that the habitat it utilizes is not sufficiently

continuous to permit it to reach all existent paramo.

Literature Cited

Acosta Sol is, M.

1944. Nuevas Contribuciones al Conocimiento de la Provincia de Esmeraldas. 606 pages. Quito: Editorial

Ecuador.

Andersson, Lars G.

1945. Batrachians from East Ecuador Collected 1937,

1938 by Wm. Clarke-Maclntyre and Rolf Blomberg.

Arkiv for Zoologi, 37A (2) :l-88.

Boettger, O.

1892. Katalog der Batrachier-Sammlung in Museum der

Senckenbergischen Naturforschenden Gesellschaft in

Frankfurt am Main. 73 pages. Frankfurt a. M., Germany.

Boulenger, George A.

1880. Reptiles et Batrachiens recueillis par M. £mile de

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1882a. Catalogue of the Batrachia Salientia s. Ecaudata in

the Collection of the British Museum. Second Edition, 495 pages, 30 plates. London: British Museum.

1882b. Account of the Reptiles and Batrachians Collected

by Mr. Edward Whymper in Ecuador in 1879-80.

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9:457^67.

1894. On the Genus Phryniscus of Wiegmann. Annals

and Magazine of Natural History, series 6, 14:374-

375.

1898. An Account of the Reptiles and Batrachians Collected by Mr. W. F. H. Rosenberg in Western

Ecuador. Proceedings of the Zoological Society of

London, 1898:107-126.

Cochran, Doris M., and Coleman J. Goin

1970. Frogs of Colombia. United States National Museum

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Cope, Edward D.

1867. On the Families of Raniform Anura. Journal of

the Academy of Natural Sciences of Philadelphia,

series 2, 6:189-206.

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Obtained by the Orton Expedition to Equador and

the Upper Amazon, with Notes on other Species.

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Cornalia, Emilio

1849. Vertebratorum Synopsis in Museo Mediolanense

Extantium quae per Novam Orbem Cajetanus Osculati Collegit Annuis 1846-47-48. Museo Mediolanense, 1849:304-315.

Duellman, W. E., and J. D. Lynch

1969. Descriptions of Atelopus Tadpoles and Their Relevance to Atelopodid Classification. Herpetologica,

25:231-240.

Dunn, Emmett R.

1931. New Frogs from Panama and Costa Rica. Occasional Papers of the Boston Society of Natural History,

5:385-401.

Espada, Marcos Jimenez de la

1875. Vertebrados del Viaje al Pacifico verificado de 1862

a 1865 por una Comision de Naturalistas enviada

pox el GoHerno Espanol. 208 pages, 6 plates.

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Guibe, Jean

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McDiarmid, Roy W.

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History, Science, 12:1-66.

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1926. Amphibia. Anura II. Engystomatidae. Das Tierreich, 49:1-110.

Noble, G. K.

1921. Five New Species of Salientia from South America.

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Orton, James

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Parker, H. W.

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1904. Viaggio del Dr. Enrico Festa nell'Ecuador a regioni

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NUMBER 145

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1889. Observations sur quelques Reptiles et Batraciens de

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1901. Ueber Reptilien und Batrachier aus Ecuador und

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