A Preliminary Analysis of the

Intergeneric Relationships of

the Frog Family Leptodactylidae

W. RONALD HEYER

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 199
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 19 9

A Preliminary Analysis of the

Intergeneric Relationships of

the Frog Family Leptodactylidae

W. Ronald Heyer

SMITHSONIAN INSTITUTION PRESS

City of Washington

1975
ABSTRACT

Heyer, W. Ronald. A Preliminary Analysis of the Intergeneric Relationships of

the Frog Family Leptodactylidae. Smithsonian Contributions to Zoology, number

199, 55 pages, appendix, 16 figures, 38 tables, 1975.—Thirty-seven characters of

external morphology, myology, osteology, life history, and chromosome morphology are studied. For each character, the evolutionary directions of changes of

states are inferred. This information is used to construct a phylogenetic hypothesis of the intergeneric relationships of the New World frog family Leptodactylidae. Five major groupings of leptodactylids are proposed: the telmatobines,

ceratophrines, leptodactylines, grypiscines, and eleutherodactylines. Formal recognition of these groupings is delayed until more information becomes available

which will likely modify the intra- and intergroup relationships. The phylogenetic

analysis demonstrates that the five groups are robust units, however.

Recognition of the five groups allows a reinterpretation of the historical

zoogeography of the family. The family Leptodactylidae had its origins in the

temperate beech forests of South America. The telmatobines represent a remnant

of the original leptodactylid stock, which has remained in the beech forests.

Two groups became adapted to drying conditions, the ceratophrines and leptodactylines. The grypiscines represent a forest-stream adaptational complex that

centered in southeastern Brazil. The eleutherodactylines were probably derived

from a grypiscine ancestor. Early attainment of direct development in the

eleutherodactylines was a preadaptation which resulted in an explosive radiation

of the Eleutherodactylus-complex, which is now represented by about 350 species

which occupy diverse environmental situations.

A leptodactylid-liopelmatid relationship is suggested, which has the advantages

of an in situ evolution of the leptodactylids rather than a migration from North

Temperate regions as previously proposed. An alternate leptodactylid-discoglossid

relationship argument which was based in large part on tadpole evidence is

countered by a consideration of the major functional adaptations of tadpoles.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5309. SERIES COVER

DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Hcyer, W. Ronald

A preliminary analysis of the intergeneric relationships of the frog family Leptodactylidae.

(Smithsonian contributions to zoology, no. 199)

Supt. of Docs, no.: SI 1.27 199

1. Leptodactylidae. 2. Reptiles—Evolution. I. Title. II. Series: Smithsonian Institution.

Smithsonian contributions to zoology, no. 199.

QL1.S54 no. 199 [QL668.E257] 591'.08c [597'.8] 74-31235
Contents

Page

Introduction 1

Methods and Materials 2

Character Selection 2

Study Group 2

Directional Criteria 2

Character Analysis 3

Analysis of Relationships 21

25 Operational Taxonomic Units 21

38 Operational Taxonomic Units 27

Additions, Modifications, and Comments on the Preferred Phylogeny .. . 32

Systematic Conclusions 34

The Preferred Phylogeny and Hennig's Sister-Group Concept 35

Comparisons with Other Schemes 35

Phylogenetic Content of the Characters Analyzed 36

Evolutionary Zoogeography 39

Historical Zoogeography 40

Within-Group Patterns 43

Origin and Evolution of Terrestriality 46

Appendix: Supplementary Data 48

Literature Cited 54

i n

A Preliminary Analysis of the

Intergeneric Relationships of

the Frog Family Leptodactylidae

W. Ronald Heyer

Introduction

Until recently, the family Leptodactylidae has

been an unstable category in frog classification. For

example, Noble (1931) placed the genera currently considered to form the family into two families, the Bufonidae and Brachycephalidae. Within

the family Bufonidae, Noble recognized seven subfamilies, of which three contained various genera

recognized in this paper. Subsequent taxonomic

accounts have been based either on a small set of

specific characters, a limited number of genera, or

both. The single exception is the work of Lynch

(1971, 1973a). The most significant change since

Noble has been the recognition of the family

Leptodactylidae as a group distinct from the

Bufonidae. The inclusion or exclusion in the family Leptodactylidae of the Australian and African

genera, and of such New World genera as Allophryne, Geobatrachus, Pseudis, Rhinoderma, and

Sminthillus has been in large part a matter of

preference, for, until recently, no comprehensive

review of the situation has been available. The

concept of the genus within the family has also

changed considerably from Boulenger's (1882) recognition of 34 genera to the 63 genera recognized

by Gorham (1966). Lynch (1971, 1973a) has made

a significant contribution to the systematics of the

Leptodactylidae with his recent review of the family

at the genus level.

W. Ronald Heyer, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution,

Washington, D. C. 20560.

The present work is an attempt to interpret and

extend data presented by Lynch (1971, 1973a),

and to use the analytical methodology of Hennig

(1966) to produce a phylogenetic hypothesis.

This phylogenetic approach has been applied to

one segment of the family, Lynch's (1971) subfamily Leptodactylinae (Heyer, 1974a). In that

study (Heyer, 1974a), certain differences with

Lynch's (1971) scheme were found with respect to

systematic conclusions. Specifically, the question

was raised whether some genera Lynch assigned to

the subfamily Leptodactylinae did not in fact have

closer relationships to genera in Lynch's subfamily

Telmatobiinae. This study was initiated to answer

that question. It soon became apparent that a preliminary analysis of the relationships among the

New World leptodactylid genera would be the best

approach. The present study must be preliminary

because total information is not available for some

rare, monotypic genera, and the range of variation

for certain large genera is not available at this time.

It is hoped that the character analysis section will

provide a base upon which additional data can be

added and analyzed as it becomes available.

ACKNOWLEDGMENTS.—For the loan of comparative material, I am grateful to William E. Duellman

and Joseph T. Collins, University of Kansas; John

W. Wright, Natural History Museum, Los Angeles

County; and Richard G. Zweifel and George R.

Foley, American Museum of Natural History.

The following assisted with the computer analysis: Joseph Felsenstein, University of Washington;

Charles D. Roberts, Smithsonian Institution; and

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Harold K. Voris, Field Museum of Natural History.

1 have liberally stolen ideas from the following

individuals who discussed various aspects of the

paper with me: Ronald I. Crombie, Smithsonian

Institution; Richard F. McGinnis, Pacific Lutheran

University; Jay M. Savage, University of Southern

California; Beryl Simpson, Smithsonian Institution;

Richard Wassersug, University of Chicago; David

B. Wake, University of California, Berkeley; and

George R. Zug, Smithsonian Institution.

David B. Wake and George R. Zug carefully

read the manuscript and offered many helpful

suggestions.

Methods and Materials

Briefly, character states are categorized for a suite

of characters from the study sample. The directionality of states is analyzed and the information from

derived states is used to generate possible phylogenetic trees.

CHARACTER SELECTION

Characters of myology, osteology, adult and larval

external morphology, and life history are sampled.

Samplings from a variety of systems produce a more

robust phylogeny than sampling of a single system

in leptodactylid frogs (Heyer, in prep.). Basically,

characters that have been used in previous systematic treatments are used. Information on character

states is taken from the literature (Lynch, 1971, and

sources cited therein. Heyer, 1974a) and from examination of specimens (Appendix).

STUDY GROUP

The family Leptodactylidae is used in the restricted sense, limited to New World leptodactyloids (Lynch, 1973a). The genus is the unit of

study. As many genera as possible are included. Inclusion of a genus depends on having complete

morphological information from at least one species. Some rare, monotypic genera were not included ilue to lack of material at this time.

The genera recognized by Lynch (1971) are

used with the exceptions of the addition of the

genera Adenomern (see Heyer, 1974a) and Vonzolinius (Heyer, 1974b). The genus Elentherodactylus is very large and the limits of variation

are not known. For purposes of this study, three

species of Eleutherodactylus are analyzed individually, one species each from the West Indies, Middle America, and South America.

DIRECTIONAL CRITERIA

The reasoning of Marx and Rabb (1970) as used

previously (Heyer, 1974a) is followed. As the sample for this study differs from the previous samples,

the criteria need to be restated.

I. OUTGROUP COMPARISONS (character state uniform in the outgroup).—To use this criterion, information is needed from a group of organisms

outside the study sample. The ideal outgroup

would be the ancestral stock to the Leptodactylidae.

Two closely related families are used as the major

outgroup to the Leptodactylidae, the Australian

family Myobatrachidae, and the African family

Heliophrynidae. If the distribution of states in

these outgroups does not allow directionality to be

assumed, other familial outgroups are used as appropriate.

A character state is presumed to be primitive if

it is found throughout the outgroup and derived if

unique or nearly so in the study sample.

II. OUTGROUP COMPARISONS (character state polymorphism in the outgroup).—A character state is

presumed to be primitive if it is widespread in the

outgroup and derived if unique or nearly so in the

study sample. As indicated previously, a large sample size of characters is needed to swamp the effect

of those possible rare instances where evolution

has not operated in a logical manner (Heyer,

1974a).

III. MORPHOLOGICAL SPECIALIZATION.—A character state is assumed to be derived if it is predominant in some adaptive specialization.

IV. ECOLOGICAL SPECIALIZATION.—A state is considered to be derived when it is relatively much

more abundant in taxa with a particular mode of

life than in all taxa. A state is considered primitive

if it has differential relative abundance among taxa

classified by adaptive zone.

At tliis time, only extreme adaptive categories

can be recognized due to the lack of information on

many genera. The following ecological categories

are considered: aquatic, fossorial, arboreal, and ter-
NUMBER 199

restrial (broad sense). Ecological categories for the

genera are presented in the appendix (Table A).

V. GEOGRAPHIC RESTRICTION.—A state is assumed to be derived if it is predominant in taxa

from a particular geographic area. The following

geographic areas appear to be important in leptodactylid frogs: Mexico; Middle America; west coast

lowlands of South America; northern Andes; southern Andes; Guyana Shield; Amazonia; southeastern

Brazil; Gran Chaco; West Indies. Geographic categories for the genera are presented in the appendix

(Table A). After the analysis was completed, R.

Crombie informed me that Telmatobius was also

found in the northern Andes. Its omission in the

character analysis section does not change any decisions. The corrected distribution was included

for the analysis which led to Figure 12.

The mechanics of sorting out the occurrence of

character states by genera among the outgroup,

ecological, and distributional categories was done

with E-Z sort cards. Data were gathered serially,

external morphology first, myology and life history

second, and osteology and karyotype last. Characters were analyzed when all available data for each

character grouping had been gathered. The apTABLE 1.—Character 1: Pupil shape (N = The total number

of genera which exhibit a given state. N may be less than

the sum of the numbers in the columns because a genus may

exhibit more than one ecological or geographic category for

a given state. The outgroup is comprised of the Australian

Myobatrachines and Cydoranines of the family Myobatrachidae and the African family Heliophrynidae. The ecological

categories are fossorial, terrestrial, aquatic, and arboreal.

The remaining categories are geographic categories for the

New World.)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

51

7

7

0

T

37

3

3

5

It

3

3

7

3

9

it

17

3

B

11

1

3

1

3

5

1

0

0

0

0

0

3

0

1

1

1

0

pendix (Table B) reflects the data on which the

character analysis was based. In some cases, more

information became available at a later time; this

information was included in the computer analysis

section. During the osteological analysis, I felt

Zachaenus should be split for further analysis; the

character analysis for bones alone reflects this division. Both subgroups of Zachaenus (sensu Lynch,

1971) have the same external morphology and

myological character states as coded for computer

analysis.

Character Analysis

1. PUPIL SHAPE.—State A: pupil round. State B:

pupil horizontal.

The distribution of states by genera among outgroup, ecological, and geographic categories is presented in Table 1.

State A is more widespread than state B in terms

of ecological and geographic categories. Except for

Hydrolaetare, all New World genera are from

southern South America. This could be interpreted

in two ways. First, because state B is relatively restricted to southern South America it could be

derived because it is a rather localized phenomenon. Second, it could be argued to be primitive because (1) state B is also found in Australian and

African genera, (2) the earliest leptodactylids occurred in southern South America, therefore (3)

TABLE 2.—Character 2: Tympanum visibility (see Table 1

and text for explanation)

State

H

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

28

0

It

0

3

2 0

1

2

3

2

2

0

3

2

8

2

8

2

B

It

0

1

0

1

2

1

0

0

0

0

01-1 0

1

1

2

0

C

It

0

0

1

1

3

0

1

2

2

1

1

1

1

1

1

1

0

D

19

6

It

0

It

11

1

0

0

0

0

0

2

0

0

1

7

0

E

6

1

1

0

2

It

1

0

0

0

1

1

3

0

1

1

0

0
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

state B is a primitive state that is still found in the

ancestral region of South America.

The application of criteria so far does not lead to

a clear-cut choice. In expanding the outgroup to

include other related families, vertical pupils (state

B) are found in the Pelobatidae, Pelodryadidae,

and some Hylidae. Thus, applying criterion II to

this larger outgroup, state B is assumed to be the

primitive state. Lynch (1973a) also argued that

vertical pupils are primitive. For purposes of

coding for computer analysis, numerical categorization of the states is preferable. Thus, state A = state

0, state B = state 1. The direction of change of character states is:

TABLE 3.—Character 3: Male thumb (see Table 1 and text

for explanation)

2. TYMPANUM VISIBILITY.—State A: tympanum

well developed, easily seen externally. State B:

tympanum partially concealed, but still visible externally. State C: intrageneric variability, some species with state A, others with state B. State D:

tympanum completely hidden, may be absent. State

E: intrageneric variability, some species with state

C, others with state D.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 2. State A is broadly distributed

among ecological categories and geographic areas

in the New World. States B and C are found in few

genera with no real patterns of distribution. State

D is widespread among the Australian leptodactyloids, with a slight trend of occurrence in fossorial

genera and a distinct trend of occurrence only in

southern South America. State E is similar in its

distribution to state D.

State D might be considered primitive on the

basis of criteria I and II. States B-E might be considered derived on the basis of criterion III, with

the hidden ear correlated with a fossorial ecological

adaptation. States B, D, and E are derived according to criterion V, as the states are relatively restricted in geographic occurrence. I choose criteria

III and IV in this case, as the trend toward earlessness has been shown to correlate with higher altitudes (e.g., McDiarmid, 1971, for bufonids) and is

apparent in other burrowing frogs (e.g., Microhylidae). As states B, C, and E are represented by

few genera and represent intrageneric variation in

part, the states are combined for purposes of further

analysis. New state 0=old state A; new state 1 =old

State

N

Hyobatrachines

Cyeloranlnes

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

29

5

2

1

2

21

0

. 3

3

2

1

2

1

0

7

1

8

2

B

22

2

6

0

6

14

2

0

1

1

2

1

5

2

3

k

6

0

c

3

0

1

0

1

1

1

0

0

0

0

0

2

0

0

0

0

0

D

3

0

0

0

0

2

0

0

0

0

0

0

1

0

0

0

2

0

E

3

0

1

0

1

2

0

0

1

1

1

0

0

1

1

1

2

1

F

1

0

0

0

0

0

1

0

0

0

0

0

1

0

0

0

0

0

states B and C, genera with the tympanum paritally

concealed at least in some member species; new

state 2 = old states D and E, tympanum hidden, at

least in some member species. The direction of

change of character states is:

3. MALE THUMB.—State A: male thumb lacking

either a nuptial pad or spines. State B: nuptial

pad present. State C: intrageneric variation, some

species with state A, others with state B. State D:

spines present. State E: intrageneric variation, some

species with state A, others with state D. State F:

intrageneric variation, some species with state B,

others with state D.

The distribution of states among outgroup, ecological, and geographic categories is presented in

Table 3. State A has the most general distribution.

Criteria I and II are not applicable, as the outgroup is well represented with both states B and C.

Criterion IV appears to be applicable to state B, as

several fossorial and one-half of the aquatic genera

have this state. Criterion V does not seem to apply.

Morphological specialization, criterion III, suggests that state A is derived because some sort of

nuptial asperity is commonly found in forms that

breed in water. The nuptial asperity aids the male

to hold onto the female during amplexus. Application of the criteria so far does not yield consistent

results. Part of the problem may be due to the

amount of intrageneric variation which led to the
NUMBER 199

recognition of states C, E, and F, each of which is

represented by few genera. The situation is resolved

by enlarging the outgroup. Members of the families

Bufonidae and Hylidae often have nuptial asperities. Therefore I consider the presence of nuptial asperities to be the primitive state. The amount

and type of intrageneric variation observed (Table

3) necessitates recognition of only three states:

state 0 = old states B, D, and F, some sort of nuptial

asperities uniformly presented; state l=old states

C, E, nuptial asperities absent in some member

species; state 2 = old state A, no nuptial asperities

in any member species. Two trends are apparent:

(1) spines are probably a derived condition over a

pad; (2) development of spines and loss of nuptial

aspersities have occurred several times in the leptodactyloid frogs. Because of this, the data must be

reduced to the new states recognized if the same

directional criteria are to apply equally for the entire study sample. The direction of change of character states is:

4. BODY GLANDS (eight extreme states are recognized).—State A: no well-defined parotoid, inguinal, or dorsolateral folds. State B: well-defined

parotoid glands present. State C: intrageneric variation, some species with state A, others with state B.

State D: well-defined inguinal glands present. State

E: intrageneric variation, some species with state A,

TABLE 4.—Character 4: Body glands (see Table 1 and text

for explanation)

TABLE 5.—Character 5: Toe disks (see Table 1 and text

for explanation)

State

H

Myobatrachines

Cycloranines

Hellophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andee

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

1.6

5

8

1

8

30

k

1

1

1

2

2

7

1

7

3

lU

2

B

1*

1

2

0

1

2

0

0

0

0

0

0

1

0

0

0

0

0

c

1

0

0

0

1

0

0

0

0

0

0

0

0

0

0

1

1

0

D

3

1

1

01-1 3

0

1

1

1

0

0

0

0

1

0

1

0

E

2

0

0

0

0

1

0

1

1

0

0

0

1

1

0

0

1

0

F

2

H 0

0

1

1

0

0

1

1

1

1

0

0

1

1

0

0

G

2

0

0

0

0

2

0

0

0

0

0

0

1

0

1

0

0

0

H

1

0

0

0

0

1

0

0

1

1

1

0

0

1

1

1

1

1

State

H

Myobatra chine s

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

kk

7

9

0

11

26

k

0

3

2

3

3

9

2

6

6

12

0

B

5

0

0

1

0

k

0

0

0

0

0

0

1

0

0

0

2

1

c

1

0

0

0

0

1

0

0

0

0

0

0

0

0

1

0

1

0

D

6

0

0

0

0

3

0

3

2

2

1

0

0

0

1

0

0

1

E

k

0

0

0

0

1*

0

0

0

0

0

0

0

1

1

0

3

0

r

I

0

0

0

0

1

0

0

0

0

0

0

0

0

1

0

0

0

others with state D. State F: intrageneric variation,

some species with state C, others with state D. State

G: well-defined dorsolateral folds. State H: intrageneric variation, some species with state A, others

with state G.

The distribution of states by genera among outgroup, ecological, and geographic categories is presented in Table 4. On the basis of criteria II, IV,

and V, state A is assumed to be the primitive state

and states B-H are assumed derived. Because so few

genera are represented among states B-H, the states

should be combined in pan. The following states

can be combined with no loss of information: B

with C, D with E, and G with H. A problem arises

with state F. Clearly, state F could be derived from

two routes, either from states B and C or from

states D and E. For coding purposes, it seems best

to double code those genera having state F so they

could be derived from either route. Each of the

types of body glands appears to be an independent

derivation. The recombined states are: new state

0=old state A; new state 1 =old states B, C, and F,

parotoid glands present in some or all member

species; new state 2=old states D, E, and F, inguinal

glands present in some or all member species; new

state 3 = old states G and H, dorsolateral folds

present in some or all member species. The directions of change of character states are:
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

5. TOE DISKS.—State A: no well-defined disks.

State B: disks on toes. State C: intrageneric variation, some species with state A, others with state

B. State D: toes disked with circumferential groove.

State E: toes disked with dorsal scutes. State F: toes

disked with 3-5 dorsal longitudinal grooves. Leptodactylus wagneri is the only species in the genus

that demonstrates intraspecific variation in having

states 0 or 1; the genus is coded as state 0.

The distribution of states by genera among outgroup, ecological, and geographical categories is

presented in Table 5.

Application of criterion I using the Australian

leptodactyloids as the outgroup indicates state A to

be primitive. Criteria IV and V also support state

A as primitive because the state is broadly distributed among fossorial, terrestrial, and aquatic zones

and among most geographic areas in the New

World. Also, all arboreal genera have toe disks with

circumferential grooves, indicating that state D is

derived. Because both states C and F are represented by single genera, the states should be recoded for further analysis. States B and C can be

combined without loss of information. State F is a

morphologically unique and distinctive state

among the study sample. As such, it does not yield

information on common ancestries. If state A is

primitive, then states B and C are derived and

states D and E are independent derivations of a

combination of states B and C. Into this scheme,

TABLE 7.—Character 7: Outer metatarsal tubercle

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cycloranines

Beliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

2 0

8

10

0

9

12

1

0

0

0

0

0

2

0

0

1

0

0

B

kh

1

1

1

It

30

3

3

it

l»

k

3

8

3

11

5

18

3

state F would have to be coded along with states

B and C into a single state.

The new coding is: new state 0 = old state A, no

toe disks; new state l=old states B, C, and E, toes

disked in all or some of the member species, not

with circumferential grooves or dorsal scutes; new

state 2= old state D, toe disks with circumferential

grooves; new state 3=old state E, toe disks with

dorsal scutes. The directions of change of character

states are:

TABLE 6.—Character 6: Tarsal decoration (see Table 1 and

text for explanation)

State

If

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

Jorth Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasll

West Indies

A

29

k

8

0

u

19

2

3

3

1

0

2

3

0

2

0

8

2

B

19

1

2

1

6

1 0

2

0

1

2

2

0

1*

2

6

6

1

C

1

0

0

0

0

1

0

0

0

0

1

0

0

0

0

0

0

0

D

2

0

0

0

0

2

0

0

0

0

0

0

0

0

2

1

1

0

E

1

0

0

0

1

1

0

0

1

1

1

1

0

0

1

1

0

0

F

3

0

0

0

0

3

0

0

0

0

0

0

0

1

0

0

3

0

6. TARSAL DECORATION.—State A: no folds, flaps,

or tubercles. State B: tarsal fold. State C: tarsal

tubercle. State D: tarsal fold and tubercle. State E:

intrageneric variation, some species with state B,

others with state C, others with state D. State F:

extensive tarsal flap.

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 6. Criterion I clearly indicates that states

C-F are derived. Criterion V further suggests that

state F is derived. Criteria I and V do not distinguish between states A and B. Criterion IV suggests

in part that state A is specialized, as all arboreal

genera lack any tarsal modifications. States A and

B are both found in bufonids, hylids, and discoglossids; state A characterizes the pelobatids.

Thus there does not appear to be any logical way of

determining whether state A or B is primitive. The
NUMBER 199

character is treated in a conservative manner, combining both states A and B as primitive. The redescribed character states, combining states A and B,

and states C, D, and E because of unique taxa are:

state 0: tarsus with fold or without modifications;

state 1: tarsus with tubercle, at least in some species; state 2: tarsus with extensive flap. The directions of change of character states are:

7. OUTER METATARSAL TUBERCLE.—State A: absent. State B: present.

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 7. Criterion II suggests that state A is

primitive; criterion IV suggests that state B is derived, as all arboreal taxa have state B; criterion

V suggests that state A is derived, as it is restricted

to southern South America. The restricted area is

in the presumed ancestral home of the leptodactylids; therefore, criterion II is not negated and state

A is assumed the primitive state. For further coding purposes, A=0, B=l. The direction of change

of character states is:

8. INNER METATARSAL TUBERCLE.—State A: normal or pointed. State B: a cornified spade. State

C: intrageneric variation, some species with state

A, others with state B.

The distribution of states among the outgroup,

ecological, and geographic categories is presented

TABLE 8.—Character 8: Inner metatarsal tubercle

(see Table 1 and text for explanation)

TABLE 9.—Character 9: Toe webbing (see Table 1 and

text for explanation)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

1*8

5

5

1

0

36

it

3

It

3

2

2

9

2

9

2

15

3

B

7

0

3

0

7

0

0

0

0

0

1

0

0

0

1

3

1

0

c

6

2

1

0

It

2

0

0

1

1

1

1

1

1

1

1

2

0

State

H

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

17

1

6

1

8

3

h

0

0

0

1

0

It

0

2

2

2

0

B

It

0

1

0

1

u0

0

0

1

0

0

1

0

0

0

1

0

c

26

3

3

0

1

21

0

2

3

1

2

3

3

0

7

2

7

2

D

9

2

1

0

1

8

0

0

0

0

0

0

0

1

1

0

5

0

E

It

1

0

0

0

3

0

1

2

2I-I 0

1

2

1

1

2

1

F

1

0

0

0

1

0

0

0

0

0

0

0

0

0

0

1

1

0

in Table 8. Criteria I and II do not appear to apply to this character; the development of a spade

is a morphological specialization, thus criterion III

indicates that state B is derived; criterion IV also

indicates that states B and C are derived, as the

states are found in fossorial genera. State C is intermediate between states A and B. For further

coding purposes, A = 0, B=l, C = 2. The direction

of change of states is:

9. TOE WEBBING.—State A: webbing present.

State B: webbing present or absent. State C: no

web. State D: lateral toe fringe present. State E:

intrageneric variation, some species with state C,

others with state D. State F: intrageneric variation,

some species with state A, others with state E.

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 9. Criteria I and II are not applicable;

criterion IV suggests that state C is derived, as the

arboreal genera have state C; criterion V indicates

that state D is derived, as the state is predominant

in southeastern Brazil. Members of the bufonids,

hylids, ascaphids, discoglossids, and pelobatids commonly have webbing; therefore, state A is assumed

the primitive state. The number of states can be

reduced without loss of information as follows:

state 0: toes webbed; state 1: toes free in some or all

species; state 2: toes fringed in some or all species.

State 2 is morphologically intermediate between
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 10.—Character 10: Life history (see Table 1 and

text for explanation)

State

N

Kyobatrachines

Cycloranines

Beliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

23

0

T

1

8

15

0

0

l

l

2

0

5

2

3

3

8

1

B

IT

3

6

0

5

9

3

0

l

l

l

l

3r-l 2

3

It

0

C

23

2

It

0

5

16

2

0

2

2

2

1

6

2

5

3

10

0

D

8

0

5

1

5

3

1

0

0

0

1

0

1

0

1

1

0

0

E

5

1

0

0

1

It

0

0

0

0

0

0

1

0

0

1

2

0

p

9

0

6

0

3

8

0

0

2

2

2

1

0

1

3

2

2

1

G

1

0

0

0

0

1

0

0

0

0

0

0

1

1

0

0

1

0

H

1 0

0

0

0

0

5

1

3

3

2

0

0

0

0

1

0

3

2

states 0 and 1 and is assumed to be phylogenetically

intermediate. The direction of change of character

states is:

10. LIFE HISTORY.—The coding of life history

categories poses certain problems. Several distinct

states are evident that are part of the same complex

in part and mutually exclusive in part. One could

therefore recognize one character with extreme

states or break up the information into two or

more characters. The danger with this latter course

is that certain information would be overemphasized as it would appear in two or more characters.

The best course appears to be to recognize the

maximum number of discrete states, such that certain genera will have more than one state, then

upon analysis of distribution of states, recategorize

the states.

Eight states are recognized. State A: tadpole present, median vent. State B: tadpole present, dextral

vent. (Lynch, 1971, states on page 26 that Caudiverbera and Odontophrynus have dextral vents; in

the generic accounts, both are listed as median on

pages 115, 131. Examination of the only Odontrophrynus tadpole at hand (USNM 121324, O.

cultripes) indicates a dextral vent. For purposes

of coding, I assume Cnudiverbera also has a dextral

vent.) State C: tadpole present, mouthparts with

a dentical row formula of 2/3. State D: tadpole

present, mouthparts with a dentical row formula

greater than 2/3. State E: tadpole present, mouthparts with a dentical row formula less than 2/3.

State F: larvae present, eggs placed in foam nest.

State G: larvae present, eggs placed in foam nest

in some species, not in others. State H: no tadpole,

direct development from encapsulated egg. For

present purposes, state H is interpreted narrowly

and does not include cases where eggs hatch into

larvae which in turn metamorphose without feeding.

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 10. Criteria I and IV support the generally accepted hypothesis that direct development

is derived with respect to presence of a tadpole.

The only criterion that allows a distinction between whether a median or dextral vent is primitive is criterion IV; all the aquatic genera have

state B indicating it is derived. Similarly there are

no strong indications as to which type of denticle

formula is primitive or whether a foam nest is derived. Foam nests are not found in ascaphids,

pelobatids, bufonids, dendrobatids, centrolenids, or

most hylids. Thus, a foam nest is probably derived.

There is no evidence to suggest whether a foam

nest was an intermediate stage in the evolution of

direct development. The safest assumption to make

for present purposes is to recognize only two states:

state 0: tadpole present; state 1: direct developTABLE 11.—Character 11: Adductor mandibularis muscle

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cycloranines

He liophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

5

0

0

0

2

1

1

1

1

1

1

0

1

0

1

2

0

0

B

48

5

9

1

9

3^

3

2

I*

3

3

2

6

3

10

3

16

3
NUMBER 199

ment. The direction of change of character states

is:

11. ADDUCTOR MANDIBULARIS MUSCLE.—State A:

both adductor mandibulae posterior subexternus

and adductor mandibulae externus superficialis

present ("s + e" in Starrett's 1968 terminology). State

B: adductor mandibulae posterior subexternus

only present ("s" in Starrett's 1968 terminology).

Starrett (1968) listed Elosia ( — Hylodes) lateristrigata as having condition s + e. In examining two

other species of Elosia (Appendix 1), I found the

condition to be s; I then examined H. lateristrigata (USNM 101720) and also find it to have the

s only condition. Starrett probably examined an

incorrectly identified specimen. I find the condition

of Sminthillus to be state B, contrary to Starrett

(1968).

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 11. Criterion I indicates that state B is

derived; none of the other criteria appear useful

in determining which state is primitive. Starrett

(1968) argued that the s + e condition is primitive

to the s condition as a loss of a muscle slip is involved. As evolutionary trends often involve simplification of parts, her reasoning is followed. For

further coding purposes, A = 0, B= 1. The direction

of change of states is:

12. DEPRESSOR MANDIBULAE MUSCLE.—State A:

muscle originates from the dorsal fascia, squamosal

and otic region, and annulus tympanicus; the relative bulk of fibers may vary, but all three regions

are clearly involved. State B: muscle origin from

dorsal fascia and squamosal and otic region only,

as in state A, the relative bulk of the two slips

may vary. State C: intrageneric variation, some

species with state A, some species with state B.

State D: muscle origin from squamosal and otic

region only. State E: intrageneric variation, some

species with state B and some with state D. State

F: origin from squamosal and otic region and annulus tympanicus.

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 12. Clearly the primitive state is either

A or B. Application of criteria I, II, IV, and V do

not indicate which state is the primitive one. Starrett (1968) indicated that an origin from both the

dorsal fascia and the squamosal region was primitive to an origin from the squamosal region only.

She did not comment on the annulus tympanicus.

Morphologically, state A is a more generalized state

than B. In a previous study in which state A was

considered primitive, there was no evidence that

indicated otherwise (Heyer, 1974a). Therefore, I

consider state A to be primitive. As states D, E,

and F are unique to single genera of New World

leptodactylids, the states are redescribed for further

TABU: 12.—-Character 12: Depressor mandibulae muscle

(see Table 1 and text for explanation)

TABLE 13.—Character 13: Geniohyoideus medialis muscle

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Possorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

2 2

2

6

0

9

1 0

2

2

1

0

2

1

2

0

5

It

5

1

B

24

2

2

1

2

2 0

1

0

2

3

2

1

3

2

6

2

10

2

C

2

0

0

0

0

1

0

1

1

1

0

0

1

1

0

0

1

0

D

3

2

0

0

0

3

0

0

0

0

0

0

0

0

0

0

1

0

E

1

0

0

0

0

0

1

0

0

0

0

0

1

0

0

0

0

0

F

3

1

1

0

1

2

0

0

1

0

0

0

0

0

0

0

0

0

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

33

1

7

1

8

21

3

1

2

2

2

1

6

2

6

3

11

2

B

3

0

0

0

1

3

0

0

1

1

1

1

1

1

2

1

2

0

c

18

5

2

0

2

12

1

2

2

1

1

0

0

0

3

2

i.

1
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

purposes as follows: state 0: origin from dorsal

fascia, squamosal and otic region, and annulus

tympanicus or squamosal, otic region and annulus

tympanicus only; state 1: origin from dorsal fascia

and squamosal and otic region in some or all species; state 2: origin from squamosal and otic region

in some or all species. The direction of change of

character states is:

TABLE 15.—Character 15: Sternohyoideus muscle

(see Table 1 and text for explanation)

13. GENIOHYOIDEUS MEDIALIS MUSCLE.—State A:

muscle contiguous medially (Heyer, 1974a, fig. 2,

0). State B: muscle contiguous medially in some

species, separated medially in others. State C: muscle separated medially (Heyer, 1974a, fig. 2, 2).

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 13. Since state B is intermediate between

states A and C, the question is the direction of

change involved. Application of the criteria does

not give clear-cut indications of the direction of

change. Because the ecological and geographic categories of state A are more broadly represented, state

A is assumed the primitive state. For further coding purposes, A = 0, B=l, C = 2. The direction of

change of character states is:

0-»l-»2

14. ANTERIOR PETROHYOIDEUS MUSCLE.—State A:

the muscle inserts on the lateral edge of the hyoid

plate. State B: the muscle inserts on the ventral

body of the hyoid in some or all species.

TABLE 14.—Character 14: Anterior petrohyoideus muscle

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorlal

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

Worth Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

kk

1

7

1

8

27

1*

3

1*

3

3

1

7

3

8

k

15

2

B

1 0

5

2

0

2

9

0

0

1

1

1

1

0

0

3

2

2

0

State

H

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

31

0

5

1

7

20

2

2

k

3

2

0

5

3

5

5

13

1

B

11

0

0

0

0

6

2

1

0

0

1

1CM 0

3

0

3

0

c

10

6

It

0

2

8

0

0

0

0

0

0

0

0

0

0

0

0

0

1

0

0

0

0

1

0

0

0

0

0

0

0

0

1

0

1

0

E

1

0

0

0

1

1

0

0

1

1

1

1

0

0

1

1

0

0

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 14. Criteria IV and V indicate that state

A is primitive. State A is found in bufonids, hylids,

dendrobatids, and pelobatids, also indicating that

the state is primitive. For further coding purposes,

A=0, B= 1. The direction of change of states is:

15. STERNOHYOIDEUS MUSCLE INSERTION.—State

A: muscle insertion entirely near lateral edge of

hyoid body (Heyer, 1974a, fig. 1). State B: insertion with some fibers near lateral edge of hyoid

body, some fibers near midline of hyoid body

(Heyer, 1974a, fig. 1). State C: muscle insertion

of narrow band of fibers extending to midline of

hyoid body (Heyer, 1974a, fig. 1). State D: intrageneric variation, some species with state A, others

with state C. State E: intrageneric variation, some

species with state A, others with state D. Lepidobatrachus has a unique insertion pattern with two

distinct slips, one inserting near where the alary

process normally is, the other inserting near the

posterolateral process. Because both slips insert on

the lateral border, Lepidobatrachus is coded as

state A.

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 15. State C is confined to the Australian

outgroups. State B is not represented in any of the
11

TABLE 17.—Character 17: lliacus externus muscle

(see Table 1 and text for explanation)

FIGURE 1.—Omohyoideus character states.

outgroups and is considered derived on the basis

of criterion I. State A is generally found among

hylids, pelobatids, and dendrobatids (Trewavas,

1933), and is considered the primitive stage. As

states D and E are only found in single taxa, the

states are combined and redefined for further analysis as follows: state 0: muscle insertion near lateral

edge of hyoid body in all species or some species

with insertion near midline; state 1: muscle insertion with some fibers near midline and some fibers

near lateral edge in all species or some species with

insertion near midline. The direction of change of

states is:

16. OMOHYOIDEUS MUSCLE.—State A: muscle absent. State B: muscle insertion on hyoid body and

fascia between posteromedial and posterolateral

processes (Figure 1B). State C: muscle insertion

on hyoid body only (Figure lc). State D: intrageneric variation, some species with state B, some

with state C. State E: intrageneric variation, some

TABLE 16.—Character 16: Omohyoideus muscle

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

2 0

6

7

0

7

12

2

0

1

1

2

0

1

1

3

3

2

2

B

12

0

0

0

1

11

0

0

0

1

0

0

1

1

3

0

5

0

c

9

0

0

1

1

1*

1

1

0

0

1

0

1

0

2

1

1*

1

D

2

0

0

0

0

1

1

0

0

0

0

0

1

0

0

0

1

0

E

3

0

0

0

2

2

0

0

1

1

1

1

0

0

2

2

2

0

F

1

0

0

0

0

0

0

1

1

0

0

0

0

0

0

0

0

0

G

2

0

0

0

0

1

0

1

1

1

0

0

1

0

0

0

0

0

H

2

0

0

0

0

2

0

0

0

0

0

0

1

1

o

0

2

0

I

^

0

0

0

0

3

0

0

1

0

0

1

1

0

1

0

1

0

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

7

0

•5

0

5

3

0

0

0

0

0

0

0

0

0

1

1

c

B

23

2

3

1

6

15

2

0

1

1

2

1

2

2

k

3

13

0

c

5

0

1

0

1

3

2

0

0

0

0

0

1*

0

0

0

0

0

D

16

3

0

0

0

13

0

2

2

1

1

1

1

0

6

1

2

2

E

2

0

0

0

0

1

0

1

2

2

1

0

0

1

1

1

1

1

species with state A, others with state C. State F:

muscle insertion on edge of hyoid plate adjacent

the posteromedial process (Figure IF). State G:

intrageneric variation, some species with state A,

others with state F. State H: intrageneric variation,

some species with state C, others with state F.

State I: muscle inserts on edge of posterolateral

process (Figure li).

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 16. All Australian leptodactyloids examined have state A, but the other outgroup member, Heliophryne, has state C. The absence of the

muscle appears to be a morphological specialization; on the basis of criterion III, together with

the consensus that evolution often involves a simplification of parts, state A is considered derived.

The criteria allow no further distinctions to be

made. The states are combined and redescribed

for purposes of further analysis as follows: state

0: muscle present in all species; state 1: muscle

present in some species, absent in others; state 2:

muscle absent in all species. The direction of

change of states is:

0-M->2

17. ILIACUS EXTERNUS MUSCLE.—State A: muscle

extends less than one-half anteriad on ilium. State

B: muscle extends from one-half to three-fourths

anteriad on ilium. State C: intrageneric variation,
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

some species with state A, others with state B.

State D: muscle extends from three-fourths to full

length anteriad on ilium. State E: intrageneric variation, some species with state B, others with state

D.

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 17. Criteria I and II are not applicable

as the various states are spread throughout the outgroup. Criterion IV indicates that state D is derived as arboreal genera have that state. Criterion

IV appears to apply also to state A as several

fossorial genera have that state. Criterion V also

indicates that state A is derived as the state is restricted to southern South America. State B is

morphologically intermediate between states A

and D. As state E represents intrageneric variation

and is represented by two genera, it is combined

with state D. For further coding purposes, A = 0,

B=l, C=2, D = 3. The directions of change of

character states are:

18. TENSOR FASCIAE LATAE MUSCLE.—State A:

the muscle inserts posterior to the anterior extent

of the iliacus externus on the ilium (Figure 2A).

State B: muscle inserts at same level as anterior

extent of iliacus externus on the ilium (Figure

2B). State C: intrageneric variation, some species

with state A, others with state B. State D: muscle

inserts anterior to forward extent of iliacus externus

on the ilium (Figure 2D). State E: intrageneric

variation, some species with states A, B, or D. State

F: muscle inserts on anterior end of ilium immediately anterior to iliacus externus and the tensor

fasciae latae and the iliacus externus are contiguous for a considerable length (Figure 2F).

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 18. The broad distribution of state A

TABLE 18.—Character 18: Tensor fasciae latae

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

33

5

1

6

2k

2

1

2

1

2

0

3

3

6

4

\h

2

B

6

0

2

0

2

1

2

0

0

0

1

0

2

0

1

1

1

0

c

1

0

0

0

0

1

0

0

0

0

0

0

1

0

0

0

0

0

D

It

0

2

0

1

2

0

0

0

0

0

0

0

0

1

0

1

0

£

2

0

1

0

2

2

0

0

1

1

1

1

0

0

1

1

0

0

F

7

0

0

0

0

5

0

2

2

2

0

1

1

0

2

0

1

1

among the outgroup, ecological, and geographic

categories indicates that state A is the primitive

state. Because states C and E represent intrageneric

variation and are represented by single New World

genera, they are combined with states B and D,

respectively. State F is distinctive morphologically,

but could be derived from state D. However, I

think it is best to be conservative at this point

and not to assume that state F was derived from

state D. For coding purposes, a conservative interpretation is that state F is independently derived

from state A. For ease of further analysis, the states

are recoded as: state O = old state A; state l=old

states B and C; state 2 = old states D and E; state

3 = old state F. The directions of changes of states

are:

19. SEMITENDINOSUS MUSCLE.—State A: interior

and exterior heads about equal bulk, or exterior

A B D G

FIGURE 2.—Tensor fasciae latae character states. FIGURE 3.—Semitendinosus character states.
NUMBER 199 13

TABLE 19.—Character 19: Semitendinosus

(see Table 1 and text for explanation)

State

N

Myobatrachinee

Cycloranines

Heliophrynids

Fossorial

Torres triri 1

Aquatic

Arboreal

Mexico

Middle America

Went Court South America

North Andes

.South Andes

G.iiana .Shield

Amo zonifi

Chsco

Southeapt Brasil

West Indies

A

6

1

0

0

0

It

1

0

1

1

1

0

1

1

It

]

2

1

B

IT

2

8

1

6

10

0

1

1

1

0

0

0

0

1

1

3

0

c

1

o

0

0

1

0

0

0

0

0

0

0

0

0

0

1

1

0

D

16

0

1

0

3

10

2

1

1

0

1

1

5

2

1

1

8

0

E

1

0

0

0

0

0

1

0

0

0

0

0

1

0

0

0

0

0

F

1

0

0

0

1

1

0

0

1

1

1

1

0

0

1

1

0

0

G

8

0

0

0

0

8

0

0

1

1

1

0

0

0

It

1

3

0

H

3

2

0

0

0

2

0

1

0

0

0

0

0

0

0

0

0

1

head bulkier (Figure 3A) . State B: exterior head

smaller than and attached by a tendon to interior

head (Figure 3B) . State C: intrageneric variation,

some species with state A, others state B. State D:

interior and exterior portions about equal, exterior

portion attached by tendon to interior portion,

bulks of two portions displaced (Figure 3D) . State

E; intrageneric variation, some species state B,

others state D. State F: intrageneric variation, some

species with states A, B, or D. State G: exterior

head rudimentary, attached by tendon to interior

portion (Figure 3G). State H: exterior head absent.

The distribution of states among the outgroup,

ecological, and geographic categories is presented in

Table 19. Criterion II indicates that state B is the

primitive state. Criterion V indicates that state D

is primitive. In a previous study in which state A

was considered the primitive state (Heyer, 1974a),

character state D of the semitendinosus did not

predict the relationships as accepted. Therefore, in

this case, criterion V is followed and state D is considered the primitive state. All the states make a

morphological series which are assumed to correlate with evolutionary directions of change. As

states C, E, and F represent intrageneric variation

and are represented by single genera, the states are

recoded as follows: new state 0 = old state D; new

state 1 =old states B and E; new state 2 = old states

A, C, and F; new state 3 = old states G and H. The

directions of change of character states are:

20. ADDUCTOR LONGUS MUSCLE.—State A: muscle

well developed, insertion is on or near the knee,

usually visible superficially. State B: muscle poorly

developed, inserts entirely on adductor magnus

muscle, covered entirely by sartorius muscle. State

C: intrageneric variation, some species with state

A, others with state B. State D: muscle absent.

State E: intrageneric variation, some species with

state A, others state D.

The distribution of states among the outgroup,

ecological, and geographic categories is presented

in Table 20. Criterion II suggests that state A is

primitive. State A is also commonly found in

bufonids, hylids, centrolenids, and dendrobatids

(Dunlap, 1960) and is assumed the primitive state.

As states C and E represent intrageneric variation

and are found in few genera, the states are combined and renumbered as follows: state 0=old state

A; state 1 = states B and C; state 2 = states D and E.

The direction of change of character states is:

21. QUADRATOJUGAL.—State A: quadratojugal

present, contacting maxilla. State B: quadratojugal

present, not contacting maxilla. State C: quadratojugal absent. State D: intrageneric variation, some

TABLE 20.—Character 20: Adductor longus muscle

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast: Brasil

nest Indies

A

27

5

5

0

It

17

2

2

2

2

1

0

2

2

6

11

2

B

17

0

3

0

It

12

0

1

2

1

2

1

2

0

It

It

]

c

It

0

1

0

1

2

2

0

0

0

0

0

1

0

0

1

D

it

0

0

1

1

3

0

0

0

3

0

0

0

o

x

1

E

1

0

0

0

1

1

0

0

!

1

1

1

0

0

i

1

o
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 21.—Character 21: Quadratojugal

(see Table 1 and text for explanation)

TABLE 22.—Character 22: Nasal contact with maxilla

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cyeloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

56

8

9

1

9

36

3

3

5

h

k

3

5

2

10

5

16

3

B

1

0

1

0

1

0

0

0

0

0

0

0

0

0

0

0

0

0

c

5

0

0

0

0

5

0

0

0

0

0

0

3

1

1

1

3

0

D

1

0

0

0

0

0

1

0

0

0

0

0

1

0

0

0

0

0

species with state A, others with state C. Lynch

(1971) incorrectly lists Notaden as having no quadratojugals (p. 38); the generic account correctly

gives the character state as state B as used here

(p. 82 and fig. 57).

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 21. Criteria II, IV, and V all

indicate that the primitive state is state A. As state

B is not found in a Neotropical genus, and state D

is represented by a single genus, the states can be

redescribed as follows: state 0: quadratojugal present, contacting maxilla; state 1: quadratojugal uniformly absent or absent in some species. The

direction of change of states is:

22. NASAL CONTACT WITH MAXILLA.—State A:

the nasal either contacts the maxilla or the two elements are in proximity to each other; I used this

state broadly, interpreting any nasal configuration

which appears to give strength to the nasal-maxillary region as state A. State B: intrageneric variation, some species with state A, others with state C.

State C: nasal widely separated from maxilla.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 22. Criteria II, IV, and V indicate that state A is primitive. As only a single Neotropical genus has state B, it is combined with

State

N

Myobatrachines

Cyeloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coatt South America

North Andes

South Andes

Guiana Shield

Eout.liea:-t Brae I 1

West Indies

A

<*5

2

9

1

9

29

3

3

3

3

b

1

I

I

( C

13

1 it

0 1

0 0

0 2

2 9

0 1

0 0

1

b

0

state C for further analysis. For further coding purposes, A = 0, B and C = 1. The direction of change of

character states is:

23. NASAL CONTACT WITH FRONTOPARIETAL.—

State A: nasals not in contact with frontoparietal.

State B: nasals in contact with frontoparietal. State

C: intrageneric variation, some species with state A,

others with state B. State D: nasals fused with

frontoparietal.

TABLE 23.—Character 23: Nasal contact with frontoparietal

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cyeloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

38

4

7

1

7

26

3

3

It

2

1

2

7

2

6

3

12

2

B

17

3

2

0

1

13

0

0

0

1

1

1

0

0

3

0

6

0

c

2

0

1

0

1

1

0

0

1

1

1

0

0

1

1

1

1

1

D

3

0

0

0

2

0

1

0

0

0

1

0

1

0

1

2

0

0
NUMBER 199 15

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 23. Criterion I indicates that

state D is derived; criteria IV and V indicate

state A is primitive. As only one neotropical genus

has state C, states B and C are combined for further

analysis. For further coding purposes, A=0, B+C=

1, D=2. The direction of change of character states

is:

24. EXTENT OF COVERING OF FONTANELLE BY THE

FRONTOPARIETALS.—State A: frontoparietals meet

medially, not exposing fontanelle; questionable

states are included in state A. State B: frontoparietals separated medially, exposing fontanelle. State

C: intrageneric variation, some species with state

A, others with state B.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 24. Criterion II suggests that

state B is primitive. As no Neotropical genus has

state C, it is not analyzed further. For further coding purposes, A=0, B = l. The direction of change

of character states is:

25. SQUAMOSAL.—State A: zygomatic ramus

slightly longer than, slightly shorter than, or equal

to otic ramus, neither ramus modified. State B: as

TABLE 24.—Character 24: Fontanelle

(see Table 1 and text for explanation)

TABLE 25.—Character 25: Squamosal

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

37

1

1*

0

6

25

2

2

k

3

It

3

3

2

11

6

lit

3

B

2h

6

6

1

5

15

2

1

1

1

0

0

6

1

0

0

5

0

c

1

1

0

0

0

1

0

0

0

0

0

0

0

0

0

0

0

0

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South An

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

erica

A

31

1

It

1

5

23

2

1

2

3

2

2

6

3

7

It

Ik

2

B

7

0

5

0

1

6

0

0

1

0

0

0

0

0

1

0

1

0

c

h

0

0

0

3

61

0

0

0

1

0

1

0

1

2

1

0

D

5

0

1

0

1

0

1

0

0

0

0

1

2

0

1

0

0

0

E

2

0

1

0

1

1

0

0

0

0

0

0

0

0

0

0

1

0

F

3

0

1

0

1

2

0

0

0

0

0

0

0

0

0

0

2

0

G

12

2

7

0

1

9

0

2

2

1

1

0

0

0

1

0

0

1

in state A with a definitely expanded otic plate.

State C: as in state B with the zygomatic ramus

articulating with the maxilla. State D: zygomatic

ramus much longer than otic ramus, neither modified. State E: as in state D with the zygomatic

ramus articulating with the maxilla. State F: as in

state D with an otic plate. State G: otic ramus much

longer than zygomatic ramus. Only extreme and

obvious modifications are recognized in states B-G;

for example, a small otic plate is coded as state A if

the two rami are about equal in length.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 25. Criteria IV and V indicate

that state A is primitive. A problem arises in attempting to determine the polarities of the states

with respect to state F. That is, state F could

either be derived through state B or state D. As few

genera are involved, it appears that the best present

solution is to combine several of the states, thereby

bypassing the problem. The new character states

are: new state 0=old states A and B; new state 1 =

old state C; new state 2=old states D, E, and F; new

state 3=old state G. The directions of changes of

character states are:

26. VOMERINE TEETH.—State A: teeth present.

State B: intrageneric variation, teeth present or

absent. State C: teeth absent.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 26.—Character 26: Vomerine teeth

(see Table 1 and text for explanation)

State

H

Myobatraehines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

U3

0

10

1

9

28

2

1

2

2

3

1

6

3

9

1*

15

2

B

6

1

0

0

1

It

1

1

2

2

1

2

3

0

1

1

0

0

c

Ik

7

0

0

1

10

1

1

1

0

0

0

1

0

1

1

it

1

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 26. Application of the criteria

does not indicate clearly which state is primitive,

although the more general distribution of state A

would suggest that this is primitive. This is also

supported by the fact that most frogs have vomerine teeth. For further coding purposes, A=0, B=l,

C = 2. The direction of change of states is:

TABLE 27.—Character 27: Median contact of vomers

(see Table 1 and text for explanation)

State

N

Hyobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

uo

8

5

1

3

29

3

3

3

3

2

2

6

1

5

2

10

2

B

1

0

0

0

0

1

0

0

0

0

0

0

0

0

1

0

1

0

c

21

0

5

0

8

11

1

0

2

1

2

1

3

2

5

It

8

1

27. MEDIAN CONTACT OF VOMERS.—State A: vomers not in medial contact. State B: intrageneric

variation, some species with contact, others without. State C: vomers in medial contact.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 27. Criteria II and IV indicate

that state A is primitive. As only one genus has

state B, it is combined with state C for further

analysis. For further coding purposes, A=0, B + C

= 1. The direction of change of character states is:

28. PROOTIC FUSION WITH FRONTOPARIETAL.—State

A: elements not fused. State B: elements fused.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 28. Criteria I, IV, and V indicate that state A is primitive. For further coding

purposes, A = 0, B = l. The direction of change of

character states is:

0-» 1

29. OCCIPITAL CONDYLES.—State A: condyles confluent. State B: condyles close to each other. State

C: condyles widely separated. State D: intrageneric

variation, some species with state B, others with

state C. Lynch (1971) states that the Grypiscini is

characterized in part in having the occipital condyles widely separated medially (p. 135). His figure

of Zachaenus parvulus substantiates this (p. 140,

fig. 91); however, his figure of Z. stejnegeri shows

TABLE 28.—Character 28: Prootic fused with frontoparietal

(see Table 1 and text for explanation)

State

N

Myobatraehines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

51

7

10

1

9

36

k

0

3

3

3

2

9

2

9

it

16

l

B

9

0

0

0

2

U

0

3

2

1

1

0

0

1

1

CM 3CM
NUMBER 199 17

TABLE 29.—Character 29: Occipital condyles

(see Table 1 and text for explanation)

TABLE 30.—Character 30: Anterior process of the hyale

(see Table 1 and text for explanation)

States

V

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

1

0

0

0

1

0

0

0

0

0

1

0

0

0

1

1

0

0

B

2 2

0

10

1

8

12

3

0

0

0

0

0

5

0

0

2

5

0

c

37

8

0

0

2

28

1

3

It

3

2

3

3

2

9

2

13

2

D

1

0

0

0

0

1

0

0

1

1

1

0

0

1

1

1

1

1

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

2 1

1

1»

1

U

13

1

3

3

2

1

1

1

0

It

1

5

2

B

1

0

0

0

0

1

0

0

0

0

0

0

1

0

0

0

0

0

c

32

It

5

0

7

22

3

0

2

2

3

1

5

3

7

5

13

1

the occipital condyles in medial proximity (p. 140,

fig. 92). I assume the figures are correct.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 29. Criterion II suggests that

state B is primitive; criterion V suggests that state C

is primitive. Lynch (1971, p. 53) indicated that the

primitive frog families, bufonids, and Rhinoderma

have state B. State B is considered the primitive

state. As few genera have either states A or D, the

states are combined for further analysis as: new

state 0=old states A and B; new state 1 =old states

C and D. The direction of change of character states

30. ANTERIOR PROCESS OF THE HYALE.—State A:

anterior process present; only well-defined processes

are included; anterior bumps or swellings of the

hyales are not considered as state A. State B: intrageneric variation, some species with state A, others

with state C. State C: anterior process absent.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 30. Application of the criteria

do not clearly indicate whether state A or C is the

primitive state. The anterior process of the hyale

represents a part of the hyoid arch that is lost in

frogs lacking the processes. State A is thus assumed

to be the primitive state. As only one genus has

state B, it can be combined with state C as a single

state. For further coding purposes, A = 0, B + C=l.

The direction of change of states is:

31. ALARY PROCESS OF THE HYOID.—State A: alary

process narrow, stalked. State B: alary process rudimentary. State C: no alary process. State D: alary

process short, not distinctly stalked. State E: alary

process broad and winglike. State F: intrageneric

variation, some species with state D, others with

TABLE 31.—Character 31: Alary process of the hyoid

(see Table 1 and text for explanation^

State

N

Myobatrachine s

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle Amsrica

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

35

0

7

1

7

2lt

3

2

3

2

1

1

6

3

6

2

13

3

B

3

0

1

0

0

2

0

1

1

1

1

0

0

0

0

0

0

0

c

5

0

0

0

2

2

0

0

0

0

1

0

0

0

1

2

3

0

D

1

0

0

0

0

0

1

0

0

0

0

0

1

0

0

0

0

0

E

9

5

1

0

2

7

0

0

1

1

1

1

0

0

3

2

1

0

F

1

0

0

0

0

1

0

0

0

0

0

0

0

0

1

0

1

0
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

state E. Lynch (1971) gives as a character state for

the Cycloraninae that the alary process has a narrow stalk (p. 75). Trewavas (1933) figures the

process of Mixophyes, a cydoranine, as being rudimentary (p. 438, fig. 28). I follow her figure.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 31. Criterion I indicates that

state C is derived. There is a morphological series

among the states, but knowing that state C is derived does not determine whether state A or state E

is primitive. Criteria IV and V indicate that state A

is primitive because of the general distribution

among categories. If state A is primitive, there

have been two morphological trends starting with a

B

TABLE 32.—Character 32: Posterior sternum

(see Table 1 and text for explanation)

o

FIGURE 4.—Representative sternal apparati (diagrammatic).

(AD, state A; EH, state B; I-L, state c; M-P, state D. A =

Odontophrynus, B = Notaden, c = Lepidobatrachus, D =

Helioporus, E =Batrachyla, F = Cycloramphus, c = Eleutherodactylus fleischmanni, H = Thoropa petropolitana, i =

Eleutherodactylus coqui, j = Syrrophus, K = Telmatobius,

L = Zachaenus parvulus, M = Edalorhina, N = Hydrolaetare, o = Limnomedusa, p = Thoropa miliaris.)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

Korth Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

25

5

8

1

10

13CVI 0

0

0

1

0

3

0

1

3

5

1

B

6

0

0

0

0

6

0

0

1

1

0

1

2

0

1

0

2

0

c

13

0

1

0

0

8

2

3

2

1

1

0

2

1

1

0

5

1

D

9

0

0

0

1

8

0

0

2CVI CVI 1

1

2

7

2

It

1

E

2

0

0

0

0

2

0

0

0

0

0

0

0

0

1

1

2

0

narrow, stalked alary process. As states D and F are

represented by single genera, they are combined

with other states and redescribed as: state 0: alary

process narrow, stalked; state 1: alary process rudimentary; state 2: no alary process; state 3: alary

process not stalked, usually broad and winglike.

The directions of change of states are:

32. POSTERIOR STERNUM.—The posterior sternum

has traditionally been given great taxonomic weight

in classification schemes of the leptodactylid genera.

Previously, I considered the variation encountered

to have more phylogenetic meaning than the recognition of only two states, that is, a cartilagenous

plate versus a bony style (1974a). In order to more

objectively categorize the variation encountered,

diagrams of each of the sternal apparati were

drawn on 6i/2- X 7y2-cm cards. At least one card

was prepared for each genus. If the posterior

sternum showed variation within a genus, more

cards were prepared. The name of each genus was

written on the back of each card. The cards were

arranged alphabetically and numbered consecutively on the side with the diagram. I then sorted

the cards into similar piles, using only the numbers

as identifications for which card was represented in

which pile. After I had determined four basic

states, I gave the cards to my coworkers to sort.

They arrived at three states. They agreed that it
NUMBER 199 19

was reasonable to split one of their states into the

two I recognized.

Five states are recognized (Figure 4): State A:

the posterior sternum a simple cartilagenous plate;

the sides of the plate broaden posteriorly to the

attachment to the pectoral girdle; a posterior bifurcation may be present or absent. State B: the

posterior sternum is cartilagenous; the sides of the

sternum are either parallel or they narrow posterior

to the attachment to the pectoral girdle; a xiphisternum is often developed, which may or may not

have a posterior bifurcation. State C: similar to

state B with deposition of mineral in the mesosternal area; a xiphisternum is always differentiated.

State D: posterior sternum differentiated into a

bony style for the mesosternum and a cartilagenous

xiphisternum. State E: intrageneric variation, some

species with state B, others with state D. The recognition of four major states differs from Lynch

(1971), most notably in that some of the genera

exhibiting state C as defined here were included

either in Lynch's cartilagenous state or in Lynch's

bony style state.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 32. Criteria I and II clearly

indicate that state A is primitive. Criterion IV

further indicates that state C is derived, as all

arboreal genera have that state. One could make a

morphological series starting with state A—»B—»

C—»D. However, the intrageneric variation represented by state E indicates that state D has been

directly derived from state B. For further coding

purposes, A = 0, B = l, C = 2, D = 3, E=4. The directions of changes of states are:

33. RELATIONSHIP OF TRANSVERSE PROCESS OF

LAST PRESACRAL VERTEBRA TO SACRAL VERTEBRA.—

State A: last presacral vertebra about same width

as sacrum. State B: last presacral vertebra much

shorter than sacrum.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 33. Criterion II suggests that

state B is primitive, whereas criterion V suggests that

state A is primitive. Trueb (1973), on the basis of

the distribution of state B among the more primitive families of frogs, considered the state primitive. State B is considered primitive. For further

coding purposes, A=0, B = l. The change of direction of character states is:

34. SACRAL DIAPOPHYSES.—State A: sacral diapophyses expanded. State B: intrageneric variation,

some species with state A, others with state C.

State C: sacral diapophyses rounded.

The distribution of states by genera among the

TABLE 33.—Character 33: Last presacral vertebral width

(see Table 1 and text for explanation)

TABLE 34.—Character 34: Sacral diapophyses

(see Table I and text for explanation)

States

N

Myobatrachines

Cycloraniaes

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

38

0

3

0CVI

28

2

3

5

k

3CVI 7

3

10

3

15

3

B

21

6

6

1

8

11

1

0

0

0

1

0

2

0

1

3

k

0

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

North Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

3fc

8

10

0

10

3

0

0

0

2

0

7

1

3

3

6

0

B

2

0

0

0

1

2

0

0

1

1

1

1

0

0

1

1

1

0

c

27

0

0

1

0

20

1

3

3

1

2

3

2

8

2

12

3
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

outgroup, ecological, and geographic categories is

represented in Table 34. State B is intermediate

between states A and C; the question is whether

state A or C is primitive. Criterion II indicates that

state A is primitive; criterion IV suggests state C is

derived, as all arboreal genera have state C, while

state A has a broader ecological representation.

The bufonids, hylids, and centrolenids have state

A, indicating the state is primitive. As only two

genera demonstrate state B, old states B and C are

combined into a single state. For further coding

purposes, A=0, B + C=l. The direction of change

of states is:

35. TERMINAL PHALANGES.—State A: terminal

phalanges simple, knobbed, or claw-shaped. State

B: terminal phalanges T-shaped.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 35. Criteria II, IV, and V indicate that state A is primitive. For further coding

purposes, A = 0, B = 1. The direction of change of

states is:

36. DORSAL CREST OF THE ILIUM.—State A: no

dorsal crest. State B: well-defined dorsal crest

present.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 36. Criterion II suggests that

TABLE 35.—Character 35: Terminal phalanges

(see Table 1 and text for explanation)

TABLE 36.—Character 36: Dorsal crest of the ilium

(see Table 1 and text for explanation)

State

H

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

Horth Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

kk

7

10

0

11

27

It

0

3

2

3

3

9

2

7

6

12

1

B

19

1

0

1

0

15

0

3

2

2

1

0

1

1

k

0

7

2

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

Horth Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

20

6

6

l

9

12

0

0

0

0

1

0

3

l

l

3

3

0

B

36

0

It

0

1

27

It

3

5

it

3

2

It

2

10

3

15

3

state A is primitive, whereas criterion V suggests

that state B is primitive. Trueb (1973, p. 100) indicates that primitive frog families have state A.

State A is considered the primitive state. For further coding purposes, A=0, B = l. The direction of

change of character states is:

37. DIPLOID CHROMOSOME NUMBER.—State A: intrageneric variation of 26 and more than 26. State

B: 26. State C: intrageneric variation of 24 or 26.

TABLE 37.—Character 37: Diploid chromosome number

(see Table 1 and text for explanation)

State

N

Myobatrachines

Cycloranines

Heliophrynids

Fossorial

Terrestrial

Aquatic

Arboreal

Mexico

Middle America

West Coast South America

Horth Andes

South Andes

Guiana Shield

Amazonia

Chaco

Southeast Brasil

West Indies

A

2

0

0

0

1

0

0

1

1

1

1

0

0

0

1

1

0

0

B

13

0

0

1i-i

10

1

1

0

0

0

0

It

1

0

1

6

1

c

3

0

0

0

0

3

0

0

0

0

0

0

1

0

1

0

2

0

D

5

3

2

0

1

5

0

0

0

0

0

0

0

0

0

0

0

0

E

1

0

0

0

0

0

1

0

0

0

0

0

1

0

0

0

0

0

F

1

0

0

0

1

0

0

0

0

0

0

0

0

0

0

1

1

0

G

9

0

0

0

2

7

0

1

It

2

2

1

1

2

5

2

It

1

H

1

0

0

0

0

1

0

0

0

0

0

0

0

0

1

1

1

0

I

2

0

0

0

0

2

0

0

0

1

0

0

0

0

1

0

0

0
NUMBER 199 21

State D: 24. State E: intrageneric variation of 26

(or more) or 22. State F: intrageneric variation of

24 or 22. State G: 22. State H: intrageneric variation of 22 or less. State I: less than 22.

The distribution of states by genera among the

outgroup, ecological, and geographic categories is

presented in Table 37. The criteria indicate that

either state B or state G is the primitive state.

Criterion II indicates that between states B and

G, B is primitive. As this corresponds with the consensus of opinion among chromosome workers, state

B is considered to be the primitive state. As one

state occurs only in the outgroup, and three states

are represented by single genera, the states are redefined: state 0: 26; state 1: intrageneric variation

of 26 and more than 26; state 2: intrageneric variation of some combination of 26 (or more), 24, 22;

state 3: 22; State 4: less than 22, at least in some

species. The directions of change of character states

are:

1 *- 0 -»2 -> 3 -» 4

Analysis of Relationships

25 OPERATIONAL TAXONOMIC UNITS

Complete data are available for 25 genera (Appendix: Table C). The relationships among these

genera are analyzed first, with two purposes in mind.

The first is to compare different analytic methods

so that a choice, when made, will be based on a

full data set. The second is that once an analytic

method is chosen as best, the results based on the

full data set can act more or less as a standard

for analysis of the larger data set in which some

data observations are not known at this time.

RELATIONSHIPS BASED ON PRIMITIVE AND DERIVED

STATES.—On philosophical grounds, I am opposed

to deducing phylogenies based in part on primitive

character states (see next section). The results of

an analysis based on primitive and derived states

is included for two reasons: (1) to compare with

results based only on derived states to see how the

results differ, and (2) for interest, as this method

resembles the way one mentally deduces relationships, that is, one mentally compares overall similarity.

The results of an unweighted pair-group method

of clustering using simple matching coefficients is

presented in Figure 5.

RELATIONSHIPS BASED ON DERIVED STATES.—One

of the principles Hennig (1966) outlined is proposing relationships based only upon derived states.

The reasoning for this is straightforward. Primitive

states merely indicate that an ancestral character

has not changed and a taxon with a primitive state

has not diverged from the ancestor with respect to

that state. Taxa sharing clusters of derived states,

on the other hand, indicate common derivations

from an ancestral condition and are indicative of

relationships. As taxa represent collections of primitive and derived character states, it is logical to

base relationships only upon the derived states. Apparently, some workers equate the practice of using

only derived states to analyze relationships with

the practice of considering primitive states to be

useless and discarding them altogether. Such is

not the case. Primitive states are important in the

process of analyzing characters; after all, one needs

to know what a primitive state is in order to know

what a derived state is for any character. Also, it

may be very informative to know which taxa have

changed the least from the presumed ancestral

stock, which means looking at those taxa that have

the greatest number of primitive states (for an

external example, see Rabb and Marx, 1973).

ADENOMERA

BATRACHYLA

CROSSODACTYLUS

HYLODES

MEGAELOSIA

TH0R0PA

CYCLORAMPHUS

Z. PARVULUS

LEPTODACTYLUS

CAUDIVERBERA

CERATOPHRYS

LEPIDOBATRACHUS

PROCERATOPHRYS

0DONTOPHRYNUS

EDALORHINA

E. COQUI

E. FLEISCHMANKI

HYLACTOPHRYNE

T0M0DACTYLUS

SYRROPHUS

EUPSOPHUS

TELMATOBIUS

PHYSALAEMUS

PLEURODEMA

PSEUDOPALUDICOLA

FIGURE 5.—Predicted phylogenetic relationships of 25 OTUs

using an unweighted pair-group method of clustering using

simple matching coefficients. (Both primitive and derived

states are included in the analysis.)
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Using the data matrix (Appendix: Table C),

two different analytic assumptions may be made.

The first is to base the relationships only upon

the derived states exhibited by the taxa, ignoring

possible ancestral but derived states. This assumes

that the state exhibited was derived directly from

the primitive state. As one does not know for certain the ancestral states of a given character due

to the incompleteness of the fossil record, this may

appear to be a reasonable assumption. A result of

an unweighted, single linkage method using a simple matching coefficient of only derived states is

presented in Figure 6.

A second analytic assumption is the inclusion of

possible ancestral but derived states. In other words,

where there is more than one derived state for a

given character, the information from the character

state trees (as analyzed in the character analysis

section) is included. This assumes that there was

some organism ancestral to the taxon that had the

ancestral but derived state. The analytic method

used was the combinatorial method developed by

Felsenstein and Sharrock. One of the options of

the combinatorial method is to include the information from the character state trees. The combinatorial method locates all nonredundant monothetic

character and taxa subsets and prints them out. A

monothetic cluster possesses a unique set of states

which is both sufficient and necessary for membership in the cluster. It is termed monothetic because

the defining set of character states is unique. A

redundant combination is one completely contained within a larger combination sharing the

same character states. A number of phylogenies

can then be constructed from the clusters. The

phylogeny of Figure 7 was constructed by maximizing the number of states at each cluster point.

This was done by finding the two taxa that shared

the most derived states and then finding the taxon

or taxa which shared the most derived states with

the initial two, etc. This is essentially sister-group

formation in terms of Hennigian terminology (see

later section for an explanation of sister-groups).

The combinatorial program recodes the derived

states in numerical sequence (Table 38).

As the phylogenies of Figures 6 and 7 differ, a

choice needs to be made as to which has the greater

probability of being correct.

First, it is interesting to note that the following

clusters appear in all three phylogenies: (1) the

two Eleutherodactylus, Hylactophryne, Syrrophus,

and Tomodactylus; (2) Crossodactylus, Hylodes,

and Megaelosia; (3) Eupsophus and Telmatobius;

(4) Ceratophrys and Lepidobatrachus, and (5)

Odontophrynus and Proceratophrys. These are robust clusters, independent of the three analytic

methods used. The reason for this is that each of

these clusters share such a large number of derived

ADENOMERA

PHYSALAEMUS

LEPTODACTYLUS

CYCLORAMPHUS

E. COQUI

E. FLEISCHMANNI

SYRROPHUS

TOMODACTYLUS

HYLACTOPHRYNE

PLEURODEMA

PSEUDOPALUDICOLA

Z. PARVULUS

CROSSODACTYLUS

HYLODES

MEGAELOSIA

EDALORHINA

THOROPA

EUPSOPHUS

TELMATOBIUS

BATRACHYLA

ODONTOPHRYNUS

PROCERATOPHRYS

CERATOPHRYS

LEPIDOBATRACHUS

CAUDIVERBERA

FICURE 6.—Predicted phylogenctic relationships of 2.r> OTUs using an unweighted, single linkage

method using a simple matching coefficient. (Derived states only are included in the analysis.)
NUMBER 199 23

states that they still cluster when primitive states

are included.

Certain character states that were not included

in the analysis can be used to determine whether

the phylogeny represented in Figure 6 or 7 is the

more probable. The placement of Cycloramphus

and Zachaenus parvulus differ in Figures 6 and 7,

Lynch (1971) indicates that these two taxa share

the derived state of a ventral phlange on the body

of the hyoid. Only Hydrolaetare also has this state.

Further, Cycloramphus and Zachaenus parvulus

share a common, derived life history pattern of

having large eggs which hatch as late larvae and

live in wet leaves. The additional evidence strongly

suggests that Cycloramphus and Zachaenus parvulus

are closely related. This close relationship is best

expressed in the phylogeny of Figure 7. For that

reason, the methodology used to arrive at the phylogeny represented in Figure 7 is chosen over the

methodology used to arrive at the phylogeny represented in Figure 6.

As mentioned, a number of phylogenies can be

constructed from the cluster information that was

used to construct the phylogeny of Figure 7. Some

of the possible alternatives should be mentioned,

but first a general limitation of the method should

be pointed out. The end point clusters, which are

based upon clusters of many character states, are

robust. The basal clusters are not robust because

they are based upon very few character states. For

taxa which branch out basally, such as Caudiverbera, which has the fewest derived character

states of the genera analyzed, the proposed relationships may well be due to convergence because

so few states are involved. Character analysis may

indicate whether the clusters appear to be reflective

of relationships or convergence, but the very

method itself cannot clearly discern basal relationships when the clusters are based on few characters.

There are three other alternative clusterings of

Adenomera, Edalorhina, Leptodactylus, Physalaemus, and Pseudopaludicola. Each of the four patterns has certain advantages, which will not be

detailed, but the important point is that the closest

relationships of each of these taxa appears to be

in the cluster of all five taxa. The reason for not

detailing the alternative clusters is that the addition of certain genera in the next stage of analysis

TABLE 38.—Characterjstate directory

Character Number

Character State Number

Transition to State

Character Number

Character State Number

Transition to State

Character Number

Character State Number

Transition to State

Character Number

Character State Number

Transition to State

Character Number

Character State Number

Transition to State

Character Number

Character State Number

Transition to State

Character Number

Character State Number

Transition to State

1

0

11

5

3

21

12

1

31

17

3

Ul

22

1

51

28

1

6l

33

0

2

2

1

12

6

l

22

12

2

32

18

1

1*2

23

1

52

29

1

62

1

3

2

2

13

2

23

13

1

33

18

2

1*3

23

2

53

30

1

63

35

1

1*

3

1

l l

7

1

Sk

13

2

31*

18

3

i*l*

21*

0

31

1

6k

36

1

5

2

15

8

1

25

11.

1

35

19

1

1*5

25

1

55

31

2

65

37

1

I

1

16

8

2

26

15

1

36

1?

2

1*6

25

2

56

31

3

66

37

2

7

1*

2

17

9

1

27

16

1

37

19

3

25

3

57

32

1

67

37

3

3

h

18

9

2

28

16

2

38

20

1

1*8

26

1

58

32

2

68

37

I

9

5

1

19

10

1

29

17

0

39

20

2

1*9

26

2

59

32

3

10

5

2

20

11

1

30

17

2

40

21

1

50

27

1

60

32

1*

may modify and clarify the clustering pattern. The

only other arrangement that gives a significantly

different pattern from the figured phylogeny (Figure 7) is with the placement of the genus Pleurodema. At the level shown in Figure 7, Pleurodema

shares 10 states with Batrachyla. Pleurodema shares

17 states with Physalaemus. Physalaemus shares 22

states with Adenomera, 20 states with Pseudopaludicola, 19 states with Edalorhina, and 18 states

with Leptodactylus. Pleurodema shares 14 states

with Adenomera and Leptodactylus, 13 states with

Pseudopaludicola and Edalorhina. The best cluster

which fits Pleurodema, together with any combination of the other four genera, is a cluster of 10

states including Pleurodema, Adenomera, Edalorhina, Physalaemus, and Pseudopaludicola. This

leaves out Leptodactylus, which makes a better

fitting unit with Adenomera, Edalorhina, Physalaemus, and Pseudopaludicola than does Pleurodema. At a cluster of 10 states, Pleurodema fits in

with Batrachyla, as figured (Figure 7). Pleurodema

clusters with Adenomera, Edalorhina, Leptodactylus, Physalaemus, and Pseudopaludicola at a level

of 9 shared states. This is the same number of

states that the cluster of Adenomera, Edalorhina,

Leptodactylus, Physalaemus, and Pseudopaludicola

shares with the cluster of the two species of Eleutherodactylus, Hylactophryne, Syrrophus, and To-
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

SYRROPHUS TOMODACTYLUS ELEUTHERODACTYLUS ELEUTHEROOACTYLUS

2,20,21,27, 17,38,49,67 COQUI FLEISCHMANNI

35,54,65 / 20,26,51,58 21,34,38,42,67,68

,23,24,34, 17,37,44

)

*9,(10),52,61,63 HYLACTOPHRYNE

17,20,37,38,

50,67

4,5,(19),31,62

CROSSODACTYLUS HYLODES

40,48,49 38

\ /

41,51,52 MEGAELOSIA

V 4,5,35,46

(11),(13),44,

58,62 THOROPA

35,38,41,

52,60

9,61,63

CYCLORAMPHUS ZACHAENUS

7 PARVULUS

\ 5,46,58,61

2,3,4,17,42,

44,50,52,62

20,21,53.

EUPSOPHUS TELMATOBIUS

17,18,27 26,35,40,41

46,50,61,64

2,3,30,32,38,

48,58,66 _

•"»•- /

BATRACHYLA PLEURODEMA

9,17,30,38, 2,7,16,23,

39,41,63 50,59,60,

N. 66,67

18,40,52,61

20,21,53"
NUMBER 199 25

PSEUDOPALUDICOLA PHYSALAEMUS

4,5,24,31,37,40, 7,16,21,27,

41,49,68 32,(33),35, 26,27,56;

36,38,39,53, 63

59

ADENOMERA LEPTODACTYLUS

5,9,17,23, (8),41,42,67

2,12,17,23,25,

48,56,67

20,60'

\

4,21,31,35,

36,50,53,59

EDALORHINA

7,12,17,23,

24,27,28,37,

38,59,67

ODONTOPHRYNUS PROCERATOPHRYS CERATOPHRYS LEPIDOBATRACHUS

(6),15,18,27, 4,5,38,42,45,67 1,2,3,14,32, 23,24,(29),30,

35,36 > 65 v 35,39

1,2,3,14,16,20, 15,16,27.28.38,50, CAUDIVERBERA

50,66 ^ 51.(50,(55) 32,35,36,56,64

42,(43),45'

44,53

FIGURE 7.—Predicted phylogenetic relationships of 25 OTUs using the combinatorial method.

(Derived states only are included in the analysis; numbers in parentheses are unique state

appearances in the phylogeny.)
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ELEUTHERODACTYLUS ELEUTHERODACTYLUS

NIGROVITTATUS COQUI

23,24,34,38,42, 51

47 V . /

^ S . / ELEUTHEROOACTYLUS

20,26,58 FLEISCHMANNI

< 21,34,38,42

9,10,63 HYLACTOPHRYNE

, 20,38,50

SYRROPHUS TOMODACTYLUS

2,20,21,27, 17,38,44,49

35>5 \ /

7,1°,57,58 SMINTHILLUS

17,20,21,26,

27,28,38,44,

49

9,23,24,34,

47,48,51,63

17,37**4,57

EUPARKERELLA HOLOADEN

22,26,38,39, 32,41

42,44,48,49,

51,63

21,23,24,54,55 NICEFORONIA

\ 31,34,38,44,48,

?'

2,3,17,20,26.

"(19)

TELMATOBIUS EUPSOPHUS

26,35,40,41, 17,18,27

46,50,61,64 1

32,57,58 BATRACHOPHRYNUS

4,22,27,28,44,

49,61,62,64

ISCHNOCNEMA

17,20,34,37,

38,44,50,57

I

1,4,5,18,52,61,62

(NO)

BATRACHYLA

9,17,18,39,

40,41,52,57,

61,63 \

2,3,48

30,38 PLEURODEMA

\ 2,7,16,18,23,40,

\ 50,52,57,59,60,61

1,20,21,53.

(NO)

ODONTOPHRYNUS PROCERATOPHRYS

(6),15,18,27 , 4,5,38,42,45

x /

1,2,3,14,16,20,50

CERATOPHRYS LEPIDOBATRACHUS

1,2,3,14,32 23,24,29,30,35,39

15,16,27,28,38, CAUOIVERBERA

50,51,54,55 32,35,36,56,64

.42,(43),45

(NO
NUMBER 199 27

ADENOMERA LITHODYTES

23,25,27, 8,11,14

36,56 /

17,26,50 VANZOLINIUS HYDROLAETARE LEPTODACTYLUS

\ 36,42 5,26,32,33, 1,8,18,41

/ 46,56 /

26,

\

4,21,31,35,53,59

BARYCHOLOS EDALORHINA

4,5,9,10, 7,27,28,59,

26,31,42, 60

47,54,58,

6 3

 \

1,12,17,18,

23,24,37,38.

"44.52.61

(NO)

PARATELMATOBIUS

1,2,23,24,26,27,

28,52,58

\ /

36,42,50

PHYSALAEMUS PSEUDOPALUDICOLA

7,16,21,27, 4,5,24,31,37,40,

32,33,35,36, 41,49

38,39,53,59, / CROSSODACTYLUS HYLODES

bl V ' 40,48,49 38

\ /

MEGAELOSIA

4,5,35,46

41,51,52

CYCLORAMPHUS ZACHAENUS

7 PARVULUS

N . 5,46,58,61

2,3,4,17,42,

44,50,52,62.

11,(13),44, THOROPA

58,62 35,38,41,

\ 52,60

\ /

9,61,63

(NO)

LIMNOMEDUSA

29,30,32,33,

37,38,41,54,

55,59,60,61,

62

FIGURE 8.—Predicted phylogenetic relationships of 38 OTUs using the combinatorial method.

(Derived states only are included in the analysis; numbers in parentheses are unique state

appearances in the phytogeny; "NO" in parentheses indicates a nonmonothetic cluster; see text.)

modactylus. Thus, there does not appear to be any

reason for preferring the placement of Pleurodema

in either of the two possible places in the phylogeny.

Analysis of the character states involved in the alternative clusterings might give support for choosing one location over the other, but that is deferred

until the additional genera have been analyzed.

38 OPERATIONAL TAXONOMIC UNITS

A computer run of the combinatorial method

was attempted using 39 taxa and 37 characters.

The 39 taxa represent those for which a full set of

morphological information was available. The additional 14 taxa lack either information on life

history, karyotype, or both (Appendix: Table C).

The program had a preset limit of 2000 clusters,

which was exceeded. After two attempts at sufficiently reducing the data matrix to accommodate

the program, it ran with 28 OTUs and 35 characters.

The limit of the program was then increased to

3000 clusters. The full data set of 39 OTUs and

37 characters exceeded this new limit. When the

matrix was reduced to 33 OTUs and 36 characters,
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the program ran. In this last run, the taxa and

character that were deleted were done so for the

following reasons. The taxa removed were Caudiverbera, Ceratophrys, Lepidobatrachus, Odontophrynus, and Proceratophrys. In the intermediate

run of 28 OTUs and 35 characters, all generic units

that had not been included in the initial run of

25 OTUs and 37 characters were represented. Also

included in the intermediate run were Caudiverbera, Ceratophrys, and Odontophrynus (Lepidobatrachus and Proceratophrys were deleted). Ceratophrys and Odontophrynus came out closely related

to each other and not related with the other taxa;

Caudiverbera was isolated at the base of the phylogeny; the other taxa were all related with the

remaining taxa. In other words, the new OTUs

being analyzed all had closer relationships with

taxa other than with Ceratophrys, Odontophrynus,

and Caudiverbera. As the cluster of the five genera

Caudiverbera, Ceratophrys, Lepidobatrachus,

Odontophrynus, and Proceratophrys remained unchanged from the initial run, these genera were

removed from the data matrix of the final run.

The character removed was character 37, the karyotype, as several of the taxa lacked information on

this character. The only other character for which

information was missing for some taxa was character 10, life history. For purposes of analysis, no

information for character 10 was entered as a 9

in the data matrix and the program treated that

character as having the primitive state.

A new phylogeny was constructed which included

all 38 taxa. The phylogeny of Figure 8 represents

the initial maximization of clusters combining the

information for Caudiverbera, Ceratophrys, Lepidobatrachus, Odontophrynus, and Proceratophrys

from the initial run and all other taxa from the

last run. Two kinds of clustering information are

indicated, in addition to the numbers and kinds

of characters represented by each cluster. Character

states that appear but once in the phylogeny are

indicated, as well as clusters that are not monothetic.

Several changes are possible in the phylogeny

figured (Figure 8). In choosing the phylogeny

which has the greater probability of reflecting relationships, the following criteria were used: (1) a

decrease in the number of convergent states, (2) an

increase in the number of monothetic clusters,

(3) an increase in the number of unique state appearances within the tree, (4) maximizing the

number of derived states in any clusters.

The cluster of Ceratophrys and Lepidobatrachus

can be joined with the cluster of Odontophrynus

and Proceratophrys at a level of four shared states.

At the lower level of two shared states, Caudiverbera is linked in. This change results in the addition

of one convergence of states, the loss of a unique

appearance of a state, and no change in the number of monothetic clusters. The changes appear to

weaken the phylogeny.

In the sample Hylactophryne and Ischnocnema

share more states with each other than with any

other taxa. The reason that they are separated in

the phylogeny of Figure 8 is that Ischnocnema is

excluded from intermediate clusters as it does not

have state 19. State 19 is a terrestrial life history.

The information for this state is not known for

Ischnocnema with certainty, although it is reasonable to assume that it might have a terrestrial life

history. If a terrestrial life history is assumed for

Ischnocnema and it is placed next to Hylactophryne

in the phylogeny, and that is the only change made,

the phylogeny of Figure 8 is improved by the loss

of six convergent states; the number of unique state

appearances is unchanged; and as a change in the

data is being assumed, the printout cannot be used

to determine whether a change in monothetic clusters results. Due to the expense of the computer

runs, having the information for monothetic clusters is not worth the cost. As the purpose of this

entire analysis is to deduce the probable relationships based on available data, the best fit of data

assumes Ischnocnema to have a terrestrial life history. This assumption is made, and the resultant

changes incorporated into the phylogeny of Figure

9.

Niceforonia can be moved from the cluster shown

on Figure 8 to a cluster of states shared with the

cluster of Hylactophryne and Ischnocnema. This relocation improves the phylogeny by reducing three

convergences; no changes in the appearance of

unique states, and because of the presumed change

in coding for Ischnocnema, the situation for monothetic cluster changes is not known. As the relocation represents improvement, the changes are

incorporated into the phylogeny of Figure 9.

The two taxa that share the most derived states

with each other in the total data set are Barycholos

and Eleutherodactylus nigrovittatus. The reason
NUMBER 199 29

that they do not appear together in the phylogeny

of Figure 8 is that Barycholos will not cluster with

any combination of other taxa that combine with

E. nigrovittatus because Barycholos lacks states 19

and 62. Character State 19 is a terrestrial life history

and the reproductive pattern is not known for

certainty for Barycholos; the program treated no

information as a primitive state. Barycholos is

suspected of having direct development, due to the

large size of the few nonpigmented eggs found in

females (Heyer, 1969a). Character state 62 is

rounded sacral diapophyses. Other than states 19

and 62, Barycholos shares every character state with

E. nigrovittatus represented by all ancestral clusters. As state 62 appears to be the only real state

excluding Barycholos from the eleutherodactylines,

I reexamined the state in a cleared and stained

skeleton of Barycholos (USNM-GOV 8015). The

tips of the diapophyses are more heavily stained,

giving an illusion of distal expansion, but the diapophyses are rounded, not expanded as previously

coded. To check this new reassessment of coding

objectively, I laid out a number of leptodactylid

skeletal preparations showing expanded and

rounded conditions, including Barycholos, and

asked my colleagues to group them into the two

states. Barycholos was consistently grouped with

those other skeleta having rounded sacral diapophyses. When character state 62 is corrected for in

Barycholos, and character state 19 is assumed, the

transfer of Barycholos next to E. nigrovittatus results in the following changes. The phylogeny is

improved by the removal of 11 convergences. No

changes are made in the appearance of unique

states. The monothetic cluster situation for the

eleutherodactylines cannot be determined because

of changes of data assumptions, but the removal of

Barycholos from Edalorhina results in the addition

of the monothetic cluster joining Edalorhina with

Physalaemus, Pseudopaludicola, etc. The evidence

clearly suggests that Barycholos is most closely related to E. nigrovittatus of those taxa included in

the analysis. This relationship is depicted in the

phylogeny of Figure 9.

The placement of Pleurodema in the phylogeny

of Figure 8 has been discussed previously. As in

Figure 8, Pleurodema can be placed with another

assemblage of genera. Pleurodema shares the most

states with Physalaemus. If this pair is chosen, a

monothetic cluster of 11 states joins Adenomera,

Leptodactylus, Lithodytes, Physalaemus, Pleurodema, and Vanzolinius (as in Figure 9). Another

monothetic cluster of six states includes Edalorhina, Limnomedusa, Pseudopaludicola and Thoropa with the above six genera. Edalorhina shares

a monothetic cluster of 15 states with Pseudopaludicola, and Limnomedusa shares a monothetic

cluster of 11 states with Thoropa. With these

changes, the best fit basally is for Hydrolaetare

to join with the cluster at five states and Paratelmatobius to come in at three shared states. The

results of these changes are a loss of one convergence, the addition of the appearance of one unique

state, and the addition of three monothetic

clusters. As this change involves the movement of

Thoropa, further evaluation is needed before a

decision can be reached. If Thoropa is taken from

this new clustering and returned to the cluster of

Crossodactylus, Hylodes, and Megaelosia, but

within the framework of the new placement of

Pleurodema, the following result: an improvement

by the loss of five convergences; no change in the

number of appearances of unique states; but a

loss of three monothetic clusters ensues. Thoropa

fits in both places at 11 shared states. With both

changes, it would appear that there is an overall

loss of six convergences with no changes in number of appearances of unique states or monothetic

clusters. The monothetic cluster situation is improved, however, because a nonmonothetic cluster

is added to a region of the phylogeny already

characterized by nonmonothetic clusters, while two

other lineages are improved with the addition of

monothetic clusters. In addition, all genera which

are known to have a foam nest are united with

the new placement of Pleurodema. The replacement

of Thoropa would be improved if it had state 62,

rounded sacral diapophyses. As I had already made

an error with this character in Barycholos, I rechecked the condition in Thoropa: the sacral diapophyses are definitely expanded in Thoropa

miliaris (USNM 97765). The overall evidence suggests that the relationships of Pleurodema and

Thoropa are best expressed as shown in the phylogeny of Figure 9.

Within the framework of the above changes as

shown in Figure 9, Edalorhina can be paired with

Limnomedusa at 10 shared states and Hydrolaetare can be paired with Pseudopaludicola at 13

states. This has the disadvantage of adding four
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ELEUTHERODACTYLUS BARYCHOLOS

NIGROVITTATUS 12,54

HYLACTOPHRYNE ISCHNOCNEMA

/ ELEUTHERODACTYLUS

23,24,38,42,47 COQUI

\ ; i

20,26,58 ELEUTHERODACTYLUS

FLEISCHMANNI

37,50 NICEFORONIA

\ 2,3

20,38

EFORONIA \

,26,31,34,48 n )

21,34,38,42

SYRROPHUS TOMODACTYLUS

2,20,21,27, 17,38,44,49

3 5 ' \ /

7,10,57,58 SMINTHILLUS

^,20,21,26,

^,28,38,44,

",5 7

q 10 91 <17 fiq EUPARKERELLA HOLOADEN

9,io,3i,37,bJ 22,26,38,39, 32,41

42,44,48,49,

51,63

2,3,20,21,23,

24,26,54,55

/

\ 2 \ p

9,23,24,31,34,

47,48,51,63

CROSSOOACTYLUS HYLODES

40,48,49 38

41,51,52 MEGAELOSIA

< 4,5,35,46

CYCLORAMPHUS ZACHAENUS Il,(l3),44 , THOROPA

7 PARVULUS 58,62 35,38,41,

\ 5,46,58,61 \ ^2,60

2,3,4,17,42, 9,61,63

44 50 52 62 ^ \u°) PARATELMATOBIUS

' ^-v ^ ^ ^ 2,23,24,26,27,

18.21.53

(NO)

TELMATOBIUS EUPSOPHUS

26,35,40,41, 17,18,27

46,50,61,64 /

32,57,58 BATRACHOPHRYNUS

BATRACHYLA

9,17,18,39,

40,41,52,57,

61,63 v

20,21,30,38,53

52,61,62
NUMBER 199 31

ADENOMERA LITHODYTES

23,25,27, 8,11,14

36,56v /

\ / PLEURODEMA PHYSALAEMUS

17,26,50 VANZOLINIUS 40 12,17,25,27,

\ 36,42 \ 32,33,35,36,

\ / \ 38,39,44,48,

5,9,63 LEPTODACTYLUS \ 56,62,64

8,36,41,42,50 \ /

\ / 2,7,16,23

4,31,35,^,62,64

1,18,52.

EDALORHINA PSEUOOPALUOICOLA

7,27,28,38, 2,4,5,25,31,40,

59 \ 41,48,49,56,62

1,12,17,18,23,24,

37,44,52,64

'21,53,59'

"20,57,(60),61

LIMNOMEDUSA

18,29,30,37,

38,41,54,55

HYOROLAETARE

4,5,26,31,35,

36,42,44,46,

\ 50,52,56

32,33,62,64

(NO)

ODONTOPHRYNUS

(6),15,18,27,

3 5 '3

\ /

1,2,3,14,16,20,50

PROCERATOPHRYS CERATOPHRYS LEPIOOBATRACHUS

4,5,38,42,45 1,2,3,14,32 23,24,29,30,35,39

/ \ /

15,16,27,28,38, CAUDIVERBERA

50,51,54,55 32,35,36,56,64

42,(43),45

FIGURE 9.—Preferred predicted phylogenetic relationships of 38 OTUs using the combinatorial

method. (Derived states only are included in the analysis; numbers in parentheses are unique

state appearances in the phylogeny; "NO" in parentheses indicates a nonmonothetic cluster;

see text.)
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

convergences, but the advantage of adding one

monothetic cluster. It is difficult to choose among

these conflicting indications, so the relationships

are left as shown in the phylogeny of Figure 9.

With all of the above changes that have been incorporated into the phylogeny of Figure 9, Paratelmatobius now fits best with the line leading to

Crossodactylus, etc., as it shares five states in this

assemblage rather than three states with the other

lineage. No other changes result from this relocation.

The relationships shown in Figure 9 represent

what I believe are the best fit of the data with the

probable relationships among those taxa included

in the analysis.

Additions, Modifications, and Comments on the

Preferred Phylogeny

Considerable data were missing for certain genera,

and intrageneric samples were small for some genera

analyzed. The reasons data are missing are essentially two: (1) some genera are known from one

or a very few specimens; in such cases, there is

not enough material available to make muscle

dissections or skeletal preparations; (2) several

genera have not been systematically reviewed

recently and the species content of the genera is

not known. To undertake reviews of these genera

would take many years; for example, I have been

systematically reviewing the genus Leptodactylus

since 1965 and am about halfway completed. Possible modifications of the phylogeny with the

inclusion of more genera or more samples of some

genera are now discussed alphabetically by genus.

Amblyphrynus: The known derived states for

Amblyphrynns are (derived states for which no

information is known for the character in parentheses): 1, 4, 5, 14, 17, 18, (19-39), 42, 44, 46, 50,

(51), 52 (53-60), 61, 62, (64). Of these states known,

Amblyphrynus shares most with the eleutherodactyline genera and certainly fits well into the basal

cluster of characters shared by all eleutherodactyline genera. See "Eleutherodactylus" for futher

comments.

Barycholos: Lynch (1973b) suggested that Leptodactylus mantipus Boulenger, known only from

the holotype, should be included in the genus

Barycholos. The single most important character

state linking these two taxa, according to Lynch,

was the presence of a sternal style which was bifurcate posteriorly. I had examined the holotype

in 1969, at which time the pectoral girdle region

had not been dissected. Since I use different character states for the sternum than those Lynch

recognizes in part, I asked Alice G. C. Grandison

to examine the sternum of the holotype and to

compare it against my four categories. She kindly

responded with a drawing which clearly indicates

that according to shape, it is either state 1 or 2

of character 32. Grandison further stated (pers.

comm.) that "there is no ossification. It is a broad

plate, bifurcated posteriorly and I'd have thought

cartilagenous." Thus mantipus has state 1 of

character 32 (character state 57 as it appears in the

phylogenies). Combining my observations with

these provided by Lynch (1973b), the known derived states for mantipus are: 1, (4-5), 10, 14, 17, 18,

(19-47), (50-56), 57 (61-64). For the known characters, mantipus seems to fall well within the

eleutherodactylines, near the cluster including

Barycholos, Eleutherodactylus, Hylactophryne,

Ischnocema, and Niceforonia. The differences

between mantipus and Barycholos pulcher listed by

Lynch (1973b) in combination with a different

sternal apparatus suggest that the two taxa belong

to separate genera. For the present, the relationships would best be expressed by considering

mantipus an Eleutherodactylus, rather than belonging to either Barycholos or Leptodactylus.

Batrachophrynus: Lynch (1971, p. 123) was of

the opinion that the relationships between Batrachophrynus and Telmatobius were very close but

that the relationships were difficult to assess because

so few species have been studied. This study was

also based on limited species samplings of these

two genera.

Crossodactylodes: The known (and unknown)

derived states for Crossodactylodes are: 1, 3, 9, 14,

17, 18 (19-39), 42, 44, 48, 49, 52 (53-60), 61, 63 (64).

These states indicate relationships with Cycloramphus, Thoropa, and Zachaenus. Also see "Zachaenus."

Eleutherodactylus: The sample of Eleutherodactylus used in this analysis comprises less than 1

percent of the known species. How much of the

variation within Eleutherodactylus was included in

this study is unknown. Within the limits of this

study, two alternate conclusions may be drawn

with respect to Eleutherodactylus: (1) Barycholos
NUMBER 199 33

should be included in Eleutherodactylus, Eleutherodactylus would then be monophyletic, or; (2)

Eleutherodactylus as presently understood is polyphyletic. I think the latter conclusion is the correct

one. Once the species groups of Eleutherodactylus

are reasonably well defined and their relationships

analyzed within the eleutherodactyline complex,

the relationships as represented within the phylogeny of Figure 9 may be considerably altered. For

example, I think it likely that there are some

Middle American species groups of Eleutherodactylus that are more closely related to Syrrophus

and Tomodactylus than other Eleutherodactylus.

This would then bring Syrrophus and Tomodactylus closer with the species of Eleutherodactylus analyzed in this study and bring Sminthillus in at a

much lower branching level. When the relationships of the Eleutherodactylus species groups are

analyzed, the relationships of Amblyphrynus and

Eleutherodactylus mantipus will also be clarified.

One thing is clear: while there may be considerable

reshuffling within the eleutherodactylines as more

units are added for analysis, the integrity of the

eleutherodactylines as a unit will remain. From

the limited character sampling of Eleutherodactylus that I have done, together with what is reported

in the literature, there is little doubt that the

relationships of Eleutherodactylus as presently understood would be much better expressed if

Eleutherodactylus were partitioned into several

genera.

Hylorina: The known (and unknown) derived

states for Hylorina are: 8, (12-13), 14, 17, 18, (20-

39), 40, 50, (53-60), 61, (64). None of the known

derived states fit any of the basal clusters of

the phylogeny of Figure 9. For the few known

derived characters, Hylorina shares most with

Batrachyla. Such a relationship would bring the

cluster of genera including Batrachyla at a lower

level to the presumed common ancestor than indicated in Figure 9, as Hylorina does not have State 1.

Insuetophrynus: The known (and unknown)

derived states for Insuetophrynus are: 1, 2, (12-13),

14, (17-47), (50-56), (61), 62, (64). With so few

known derived states, the relationships could lie

most anywhere on the phylogeny of Figure 9.

Based on intuition, I would place Insuetophrynus

with Batrachyla, Batrachophrynus, etc.

Macrogenioglottus: Reig (1972) discussed the

relationships of the monotypic Macrogenioglottus

and proposed a new family for it. Lynch (1971)

treated the taxon as a synonym of the genus

Odontophrynus. I have not examined any specimens, but the data provided by Reig (1972) indicates that Macrogenioglottus has the following

known (and unknown) derived states: 1, 2, (4-

5), 14, 15, 16, 18, (23-24), (26), 29, 30, 35, 36,

42, 44, 51, 53, 54, 55, (64). Two things are clear

within the context of the phylogeny of Figure 9.

First, the relationships lie with the cluster represented by Caudiverbera, Ceratophrys, Lepidobatrachus, Odontophrynus, and Proceratophrys. Second, Macrogenioglottus and Odontophrynus each

have a cluster of character states not shared with

the other. The differences are great enough that

for present analytic purposes, the relationships

would be obscured by considering Macrogenioglottus and Odontophrynus as congeneric. Macrogenioglottus shares the following states with the

following combinations of taxa: 11 each with

Ceratophrys and Lepidobactrachus, 10 with Odontophrynus, 7 with Proceratophrys, 5 with Caudiverbera; 6 with a duster including both Ceratophrys

and Lepidobatrachus, 4 with a cluster including

Odontophrynus and Proceratophrys, and 1 with a

cluster including Caudiverbera, Ceratophrys, and

Lepidobatrachus. The data indicate that Macrogenioglottus has its closest relationships to Ceratophrys, Lepidobatrachus, Odontophrynus, and Proceratophrys, and that its inclusion tightens up this

cluster and excludes Caudiverbera. It is pointless to

speculate further on the exact relationships of

Macrogenioglottus until (1) the full data set becomes available for Macrogenioglottus, and, more

importantly, (2) more species are examined, especially of Odontophrynus and Ceratophrys including

what many authors consider the distinct genus

Chacophrys.

Physalaemus: The five species of Physalaemus

used in this analysis were chosen to get a cross

section of the genus as defined by Lynch (1971).

Physalaemus has one of the highest proportions

of character state variability within the entire

sample. It may be that when additional samples

of more species of Physalaemus are analyzed, the

relationships would be best expressed by partitioning the genus. If partitioned, the relationships

expressed within the phylogeny of Figure 9 would

probably change little, however, because the

Physalaemus-complex is likely a monothetic group.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pleurodema: As with Physalaemus, the sample

used in this analysis was small. A greater understanding of the variation within the genus as

presently conceived may lead to partitioning of

the genus, which could have an effect on the phylogeny represented in Figure 9. The discontinuities

in geographic distribution and life history, with

some species having foam nests and others not,

are particularly suggestive that the genus is polyphyletic.

Scythrophrys: Lynch (1971) proposed a new

genus for Zachaenus sawayae Cochran. So few derived character states are known for the only

specimen so far known that its relationships cannot be determined with any assurance. It probably

is most related to the Zachaenus, Thoropa, and

Paratelmatobius cluster. See also "Zachaenus."

Telmatobufo: The known (and unknown) derived character states for Telmatobufo are: 2, 3, 4,

6, (12-13), (20-39), (41-43), 46, (52-60), 61, (64).

Telmatobufo shares the most states with Batrachophrynus. Assuming that Telmatobufo has its

closest relationships with Batrachyla, Batrachophrynus, etc., the inclusion of Telmatobufo would

result in the entire cluster being independently

derived from the presumed ancestral stock, as

Telmatobufo has neither states 1 nor 14.

Zachaenus: In the middle of the character

analysis, I thought it best to separate the data on

Z. parvulus from the data on Z. stejnegeri. As the

limits of the combinatorial program were always

approached or exceeded with the data samples, the

data for Z. stejnegeri were never included. The derived states for stejnegeri are: 1, 2, 3, 4, 5, 14, 17, 18,

20, 21, 42, 44, 46, 50, 53, 61, 62, 64. The most

states are shared with Z. parvulus, but the states

that are not shared, 52 and 58, would considerably

modify the relationships among Cycloramphus,

Crossodactylus, Hylodes, Megaelosia, Thoropa,

Paratelmatobius, and Zachaenus as expressed in

the phylogeny of Figure 9. Two conclusions may

be drawn. First, characters 52 and 58, having

to do with the separation of the occipital condyles and the type of sternum are fundamental

characters within the total phylogeny. Thus the

relationships as presently understood would best be

expressed by generically separating stejnegeri from

parvulus. Lynch (1971) was the first author to

suggest combining the taxa in the same genus.

I think the data presented here warrant removing

Craspedoglossa from the synonymy of Zachaenus.

The second conclusion is that the relationships

within this total assemblage are unclear, as indicated in part by the several nonmonothetic clusters

in the phylogeny of Figure 9 for this group. I

think the relationships among this group will be

modified considerably as more information is

available on the variability and state content of

Craspedoglossa, Crossodactylodes, Scythrophrys,

Thoropa, and Zachaenus. The group itself will

remain intact as all genera have a number of derived states in common.

Systematic Conclusions

As is clear from the previous section, this study

is a preliminary analysis due to the lack of knowledge of states for some genera and intrageneric

variability for other genera. The combined information on relationships clearly indicates five major

groupings within the family. I think the groupings

will remain intact as more information becomes

available, but that the relationships within and

among groupings will be subject to change with

the addition of more information. The limits of

this study do not clearly demonstrate that the five

groups constitute a monophyletic group. Thus, the

relationships among the five groups themselves can

only be outlined in broadest terms for the present.

See "Historical Zoogeography" for further comment. Due to likely future modifications, the proposal of these five categories as formal taxonomic

units is premature. Rather, I prefer to consider

these units as informal for the present. The units

and their components follow in alphabetical

arrangement.

1. CERATROPHRINES

Ceratophrys

Lepidobatrachus

Macrogen ioglottus

Odontophrynus

Proceratophrys

2. ELEUTHERODACTYLINES

Amblyphrynus

Barycholos

Eleutherodactylus-complex

Euparkerella

Holoaden

Hylactophryne

Ischnocnema

Niceforonia

Sminthillus

Syrrophus

Tomodactylus

3. GRYPISCINES

Craspedoglossa

Crossodacty lodes

Crossodactylus

Cycloramphus

Hylodes

Megaelosia

Paratelmatobius

Scythrophrys

Thoropa

Zachaenus

4. LEPTODACTYLINES

Adenomera

Edalorhina
NUMBER 199 35

Hydrolaetare

Leptodactylus

Limnomedusa

Lithodytes

Physalaemus

Pleurodema

Pseudopaludicola

Vanzolinius

5. TELMATOBINES

Batrachophrynus

Batrachyla

Caudiverbera

Eupsophus

Hylorina

Insuetophrynus

Telmatobius

Telmatobufo

The Preferred Phylogeny and

Hennig's Sister-Group Concept

In addition to the concept of deducing relationships on the basis of shared, derived clusters of

states, Hennig (1966) advocated the formation of

sister-groups. Sister-groups require involvement of

at least two characters, with reciprocal representation of primitive and derived states. Hennig (1966)

proposed that this operational definition of sistergroups indicates a common point of evolutionary

divergence. The phylogeny of Figure 9 was not

constructed with sister-group formation as a criterion. The location of sister-groups within the

preferred phylogeny is discussed by group.

For the Telmatobines represented in Figure 9,

all clusters represent sister-groups; this is also true

for the ceratophrine genera of Figure 9.

Within the grypiscines, Thoropa does not form a

sister-group with Crossodactylus, Hylodes and Megaelosia. Otherwise, all other groups are sistergroups.

Within the leptodactylines, Limnomedusa and

Hydrolaetare do not form a sister-group with

Adenomera, Lithodytes, Vanzolinius, Leptodactylus, Pleurodema, and Physalaemus. All other clusters yield sister-groups. With a slight rearrangement,

all cluster groups can be sister-groups within the

leptodactylines. Limnomedusa forms a sister-group

with Adenomera, Lithodytes, Vanzolinius, Leptodactylus, Pleurodema, and Physalaemus. Next, Hydrolaetare forms a sister-group with Adenomera,

Lithodytes, Vanzolinius, Leptodactylus, Pleurodema, Physalaemus, and Limnomedusa.

Within the eleutherodactylines, there are several

instances of clusters which do not represent sistergroups. The best rearrangement of taxa yields two

major groups. Group 1 is represented by Eleutherodactylus nigrovittatus and Barycholos add

Eleutherodactylus coqui add Eleutherodactylus

fleischmanni add Hylactophryne and Ischnocnema

add Niceforonia. Group 2 is represented by Syrrophus and Tomodactylus add Sminthillus add

Euparkerella and Holoaden. All clusters within the

two groups are sister-groups; groups 1 and 2 are

not sister-groups.

Among the five major groups, only the eleutherodactylines and leptodactylines form sister-groups

with one another.

The sister-group concept allows an assessment of

weaknesses in the phylogeny. In this case, those

areas where sister-groups cannot be formed with

the available data are within the grypiscines and

eleutherodactylines and among the five major

groups. These are the weakest parts of the phylogeny in terms of the data. In terms of the grypiscines and eleutherodactylines, the weakness of proposed relationships will be corrected only with

gathering more data. Gathering more data may not

help to resolve the relationships among the five

major groups because of the extreme likelihood of

convergent data not allowing sister-group formation at that level. The best test for the validity of

the five proposed groupings at present is to see if

the five groupings make sense biogeographically.

Comparison with Other Schemes

The only major difference between the phylogenies of Figures 7 and 9 is the placement of Pleurodema, which has already been discussed. The

similarities mean that the deletion of the karyotype

character in the analysis leading to the phylogeny

of Figure 9 did not affect the overall analysis of

relationships.

Lynch (1971) has been the first since Boulenger

(1882) to systematically review the entire family

Leptodactylidae. Lynch (1971) provides a historical review of all the suprageneric classifications

proposed for various sections of the family. The

interested reader is referred to Lynch's review for

a more complete comparison with the informal

classification proposed herein. My informal groups

are generally equivalent to certain of Lynch's subfamilies and tribes with certain exceptions as explained below.

CERATOPHRINES.—Several South American workers (e.g., Reig, 1972, and sources cited therein)

consider the ceratophrines to represent a distinct

family. These workers consider the ceratophrines

to have the same generic composition as recognized
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in this study. Lynch (1971) restricted his subfamily

Ceratophryinae to include only the recent genera

Ceratophrys and Lepidobatrachus. Lynch (1971)

placed the genera Odonlophrynus and Proceratophrys as a tribe into the subfamily Telmatobiinae.

The results of this analysis indicate that the relationships of Odontophrynus and Proceratophrys lie

with Ceratophrys and Lepidobatrachus, not with

the genera Lynch placed in the subfamily Telmatobiinae. This study does not provide evidence either

for or against recognition of the ceratophrines as

a distinct family. Such an analysis would need to

compare the five groups recognized here with other

family groupings generally recognized to be related to the leptodactylids.

ELEUTHERODACTYLINES.—This grouping is equivalent to Lynch's (1971) tribe Eleutherodactylini of

the subfamily Telmatobiiaae. The only difference

is that this study indicates that the genus Barycholos is an eleutherodactyline rather than a member of the subfamily Leptodactylinae as suggested

by Heyer (1969a) and Lynch (1971).

Grypiscines.—The grypiscines as recognized

herein are composed of members of four different

groupings proposed by Lynch (1971). The members of Lynch's subfamily Elosiinae and tribe

Grypiscini of the subfamily Telmatobiinae are completely included. In addition, the results of this

study indicate that Paralelmatobius is a grypiscine,

rather than a member of the (subfamily Leptodactylinae as proposed by Lynch (1971) and Thoropa

is a grypiscine, rather than a member of the tribe

Alsodini of the subfamily Telmatobiinae.

LKI'TODACTYLINES.—This grouping is equivalent

to Lynch's (1971) subfamily Leptodactylinae except for Lynch's inclusion of Barycholos, an eleutherodactyline, and Paralelmatobius, a grypiscine, as

discussed previously.

TKI.MATOBINKS.—This group is equivalent to a

combination of Lynch's tribes Alsodini and Telmatobiini ol the subfamily Telmatobiinae with the

exception of Thoropa, which Lynch (1971) considered an alsodine, but is here considered a

grypiscine as discussed above.

The study was initiated because of uncertain

relationships of some genera from a previous analysis (Heyer, 1971a). The previous study concluded

that there were two major clusterings of leptodactylines: (1) Adcnomcra, Lcptodactylus, Lithodyles,

and Vanzolinius, and (2) Physalacmus, Plrurodema, and Pseudopaludicola. The present study

agrees with the first cluster completely, but indicates that Pseudopaludicola is not as closely related

to Physalaemus as indicated in the previous study.

Examination of additional species of Pseudopaludicola, Physalaemus, and Pleurodema is needed to

clarify the relationships among these genera. The

previous analysis could not distinguish whether

Barycholos, Edalorhina, Hydrolaetare, Limnomedusa, or Paratelmatobius were basally related with

the other leptodactylines or had closer relationships

elsewhere in the family. This study indicates that

Edalorhina, Hydrolaetare, and Limnomedusa are

related to the other leptodactylines, but that Barycholos is an eleutherodactyline and that Paratelmatobius is a grypiscine.

Phylogenetic Content of the Characters Analyzed

Once the best phylogeny is chosen, the behavior

of the individual states can be evaluated within

the phylogeny. Certain states are better than others

in producing any given phylogeny. Those states

that form large clusters in the phylogeny contain

more phylogenetic information than those that do

not, in terms of that given phylogeny. The analysis of characters of a given phylogeny will only

have as much meaning as the degree of reflection

the phylogeny is to reality. Such an analysis assumes

that all states are adaptive, but that some states

contain more phylogenetic information than others.

The particular phylogeny analyzed is identical

to Figure 9 except that Caudiverbera is independently derived from a common ancestor as is the

cluster of Balrachyla, Batrachophrynus, Eupsophus,

and Tehnatobius. This particular phylogenetic arrangement maximizes monothetic clusters and numbers of states within clusters. To facilitate comparison of characters, each state used in the phylogeny

was evaluated with respect to its clustering behavior

within the phylogeny as follows. The number of

times a particular state actually appears in the phylogeny is divided by the total number of times the

state could appear in the phylogeny. The lower

the number, the better the state behaves in forming clusters. For example, if a state appears once

in a cluster ancestral to five taxa inclusively, the

state has a value of 1 (a single appearance in the

phylogeny, although the state is shared by all five

taxa above the cluster in which the state appears)
NUMBER 199 37

divided by 5 (the total number of times the state

could appear) equals 0.20. Alternatively, if another

state appears in a phylogeny five times, but in each

case the appearance is in an end point taxon, the

state has a value of 5 divided by 5 equals 1. Thus a

state with a value of one has no phylogenetic information. All states are analyzed in this fashion.

When a given character is represented by more

than one state, an average value of the component

states is used as the value for that character. The

values for each character were plotted on a graph

by occurrence. There are no clear groupings, but

the following characterizations appear reasonable.

The best characters in terms of clustering behavior

in the preferred phylogeny have values ranging

from 0.08 to 0.21. Good characters range in value

from 0.26 to 0.42. Average characters range from

0.49 to 0.72. Poor characters range from 0.77 to

1.00. The adjectives "best, good, average, poor"

as used throughout the remainder of this section

represent these value ranges. This method of analysis, while having the desirable trait of reproducibility has one minor drawback in terms of describing the phylogenetic value content of characters

within a phylogeny—there are two ways of arriving

at low numbers. The first is for the state to have

a unique appearance in the phylogeny and the

cluster in which it appears is ancestral to several

taxa. This is the best kind of character with respect to clustering behavior, and is referred to as

a Type I character for the best character category.

The second is for a given state to appear a low

number of times in the phylogeny, but each appearance is ancestral to several taxa. This is referred

to as a Type II best character. All good, average,

and poor characters are Type II. While the Type

II character state is descriptive of large clusters

and thus aids in the production of the phylogeny,

it is obviously not the same quality of state as the

Type I state. This problem is not present at the

other end of the spectrum. High values always represent low phylogenetic content.

The overall average clustering value for the 64

characters used to build the phylogeny of Figure

9 is 0.54. The average value for external characters

is 0.48, for muscle characters the average value is

0.61, and for skeletal characters the average value

is 0.54.

The only best Type I character is character 10,

life history.

There are four best Type II characters: pupil

shape, outer metatarsal tubercle, anterior process

of the hyale, last presacral vertebral width.

There are 11 good characters: male thumb, toe

disks, toe webbing, adductor mandibularis muscle,

omohyoideus muscle, fontanelle, occipital condyles,

posterior sternum, sacral diapophyses, terminal

phalanges, dorsal crest of ilium.

There are 13 average characters: tympanum visibility, tarsal decoration, inner tarsal tubercle, depressor mandibulae muscle, geniohyoideus medialis

muscle, sternohyoideus muscle, iliacus externus

muscle, tensor fasciae latae, semitendinous muscle,

nasal contact with frontoparietal, squamosal, median contact of vomers, prootic fused with frontoparietal.

There are 7 poor characters: body glands, anterior petrohyoideus muscle, adductor longus muscle, quadratojugal, nasal contact with maxilla,

vomerine teeth, alary process of the hyoid.

The diploid chromosome number was not included in the data set from which the phylogeny of

Figure 9 was produced. The clustering index value

for the character derived from the phylogeny of

Figure 7 is 0.79, indicating that overall, the diploid

number has little phylogenetic information. State

66 of the character, variation of the 2N=26, 24, 22

(which in the tree will also include the derivable

states of 67, 2N=22 and state 68, 2N= less than 22)

has a value of 0.27. Thus, within the character,

one of the states does have good phylogenetic information, although the state appears four times

in the tree. It is interesting to note that if the

phylogenies of Figures 7 and 9 are reasonably correct, the diploid number of 22 has arisen more than

once in the family. Also, it would be much more

parsimonious to assume that the diploid numbers

of 24 and 26, found in Adenomera, represent a

secondary reversion to a higher diploid number

from an immediately ancestral condition of 22,

rather than the retention of the primitive state.

Further kinds of data are needed to determine the

status of the Adenomera karyotype. The conclusion seems warranted that caution must be used in

proposing relationships at the generic level based

only on the karyotype, reinforcing the same conclusion of a study done at a lower level of analysis

(Heyer and Diment, 1974).

The variation of clustering values of individual

states is similar to the average of the states for all
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

polytypic characters except for the character, posterior sternum. State 60, the presence of a bony

style in at least some species, has a clustering value

of 0.18 and appears twice in the phylogeny of Figure 9, once at the cluster basal to the leptodactylines, and once at the end point Thoropa. In the

initial character analysis, two species of Thoropa

were examined for the sternal apparatus. In one,

Thoropa petropolitana (USNM 164135), the sternum is clearly the same as state 1 of character 32,

or state 57 as it appears in the phylogenies: the

sternum is cartilagenous, the sides are parallel

proximally, and the distal portion is expanded and

bifurcate (Figure 4H). The posterior sternum of

Thoropa miliaris (USNM 97765), is quite different in appearance. Functionally, the apparatus is

a sternal style, and this is how it was coded in the

analysis (Figure 4P). However, there is a major

difference between the sternal style of Thoropa

miliaris and the styles of the leptodactylines. The

styles of the leptodactylines are bony and appear

whitish in dissection. The style of Thoropa miliaris

is composed of calcified cartilage and appears

brownish in dissection. A further differentiation

is that the posterior sternal apparatus of the leptodactylines is composed of two distinct parts, a

bony mesosternum and a cartilagenous xiphisterum.

The sternum of Thoropa miliaris grades from a

bony style proximally to a cartilagenous bifurcate

xiphisternum distally. Thus, while the styles of

Thoropa miliaris and the leptodactylines are functionally the same, all evidence indicates that they

are not phylogenetically the same. With the sternal

apparatus of Thoropa miliaris recoded as state 2

of character 32 or state 58 as it appears in the

phylogenies, the relationships of Thoropa become

clearer. Thoropa had been placed in a cluster with

the leptodactylines previously at one point in the

analysis of relationships. It is now clear that this

clustering was made possible by the incorrect coding of the sternum in Thoropa, and that as suggested in Figure 9, Thoropa is a grypiscine, not a

leptodactyline. Further, state 60 is now a unique

state in the phylogeny of Figure 9, appearing only

in the basal cluster of the leptodactylines.

I am actually surprised by the high number of

convergences that appear in the phylogenies. I had

hoped that the data would not have as much noise

in it as apparently it does. Nevertheless, I am convinced that the methodology used to analyze these

data resulted in a phylogeny which reflects the relationships reasonably well. Perhaps one of the most

critical areas of the phylogeny is in the clusters

which define each of the five informal groups.

Overall, these five clusters are sound. The cluster

joining the ceratophrines is a monothetic cluster,

although it does not contain any unique states. The

telmatobines are represented by a monothetic cluster in the phylogeny of Figure 9, but with the addition of the other genera, there is no cluster that

defines the group. As discussed in the zoogeography

section, there is valid reason for not expecting the

telmatobines to share a cluster of derived states.

The eleutherodactylines have a basal, monothetic

cluster with a unique state, direct development.

The leptodactylines have a monothetic basal cluster with a unique state involving the sternal style.

The grypiscines are the weakest group represented

in the phylogeny of Figure 9, as they do not share a

monothetic cluster. In fact, the relationships of the

grypiscines in Figure 9 could as well be expressed

with the leptodactylines as they share the states 14,

20, 57. From evidence of evolution and zoogeography, however, to be discussed below, I think the

strongest case is for the closest relationships of the

grypiscines to lie with the eleutherodactylines

rather than the leptodactylines.

It is apparent from the nature of the data that

two requirements had to be met to arrive at what I

believe are meaningful conclusions. The first is

adequate sample size in terms of numbers of characters. I think with the diversity represented by

the genera of the family, the number of convergences in any data set is going to be high. To overcome this, the sample size of characters must be

fairly large, otherwise the noise due to convergences

may well override the phylogenetic information

present. I think the number of characters used

herein is on the low end of the sufficient number.

The second is adequate sample size in terms of

numbers of taxa analyzed. For instance, if Eleutherodactylus nigrovittatus had not been included

in the sample, I would most likely have included

Barycholos in the leptodactylines rather than the

eleutherodactylines. It is for this reason that I

think there will be modifications of the intragroup

relationships as more information becomes available.
NUMBER 199 39

Evolutionary Zoogeography

The emphasis of this discussion will be at the

level of the five major informal groupings for the

obvious reason that the intragroup details are not

worked out at present.

In formulating hypotheses on the evolutionary

zoogeography of leptodactylid frogs, the basic data

used are radiation centers and the broad ecological

adaptations exhibited by the genera. The following

radiation centers and broad ecological adaptations

appear to represent the most basic units below

which subdivision is not desirable. In part, the

radiation centers and ecological adaptations overlap.

RADIATION CENTERS.—A radiation center is a

geographic area that contains at least one endemic

genus of leptodactylid frog and is further set apart

by distinctive vegetation type or geographic relationships. The radiation centers correspond to the

geographic areas used in the initial character analysis with the exception of the Guiana Shield (discussion follows). The nine radiation centers, the

percentage of primitive states averaged for all

genera found in each center, the total number of

genera, and the number of endemic genera found

within each center are presented diagramatically

(Figure 10). As presently understood, there are no

genera endemic to Middle America. However, once

the relationships within the Eleutherodactyluscomplex become better understood, I believe there

will be at least one eleutherodactyline endemic to

this area. The absence of endemic leptodactylid

frog genera from the Guiana Shield is surprising at

first thought because the Guiana Shield is an ancient land mass that supports a number of endemic

frog genera. If the Guiana Shield region is thought

of in a broad sense, radiations of the following frog

families are found there at present: Allophrynidae,

Bufonidae, Centrolenidae, Dendrobatidae, and Hylidae (from Savage, 1973). It is therefore instructive that the Guiana Shield region does not

represent a radiation area for leptodactylid frogs.

The radiation centers compare favorably with

Miiller's distribution centers (1973). The radiation

centers recognized here encompass several of Miiller's distribution centers. This is to be expected, as

the unit of analysis for the radiation center is the

genus, while that of Miiller's distribution center

was the species and subspecies.

\

we

44.6

4

1

M

38.9

5

2

MA \

Wl

43.7

3

1

\ 37.8 \

\ 4 \

\ (1) \

\ \

NA

39.7

4

1

SA

61.1

10

7

A

44.9

II

5

SEB

52.7

20

14

C

51.0

6

1

FIGURE 10.—Leptodactylid radiation centers, (M = Mexico,

MA = Middle America, wi — West Indies, we = West coast

of South America, NA = northern Andes, SA = southern

Andes, A = Amazonia, SEB = Southeastern Brazil, c = Chaco.

Within each center, the upper number is the average percentage of primitive states, the middle number is the total

number of genera, and the lower number is the number of

endemic genera; also see text.)

ECOLOGICAL ADAPTATIONS.—The following broad

adaptations appear to be the major ones exhibited

by living leptodactylids: (1) adaptations associated

with temperate beech forest environments; (2)

adaptations associated with tropical and subtropical forests; (3) adaptations to tropical montane

environments; (4) arid adaptations, such as to the

Gran Chaco environment; and (5) savanna adaptations. The exact ecological adaptations are not
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

known for several of the genera: the assignments

are based on whatever information is available.

Data on radiation centers and ecological adaptations are summarized by groups.

TELMATOBINES.—Seven of the eight genera comprising the telmatobines are restricted to the beech

forests of the southern Andes. Telmatobius is found

in the northern and southern Andes. All adaptations within the group appear to be responses to

particular habitats within the beech forest ecosystem. Most notable are the trends toward adult

aquatic adaptations {Batrachophrynus, Caudiverbera, Telmatobius), stream adaptations (Telmatobufo), and toward terrestriality (Batrachyla).

CERATOPHRINES.—All genera are found in either

the Chaco and/or southeastern Brazil. In addition,

Ceratophrys is found in Amazonia and the east

coast of South America. The present ecological

adaptations are either adaptations to arid environments and/or forest environments. The overall

morphology of ceratophrines strongly suggests a

basic semifossorial adaptation. The semifossorial

adaptations would clearly have a selective advantage in arid environments and it seems most reasonable to assume that the basic adaptation of

ceratophrines was to an arid environment and that

the semifossorial adaptations allowed secondary access into forested situations.

LEPTODACTYLINES.—The 10 genera comprising

the leptodactylines are collectively widely distributed—they occur in every radiation center except

the northern Andean center. The genera which

demonstrate endemic patterns are found either in

Amazonia (Edalorhina, Hydrolaetare, Lithodytes,

Vanzolinius) or southeastern Brazil (Limnomedusd). All genera have ecological adaptations to

either forest or savanna environments. In sorting

out which adaptation is basic to leptodactylines,

forest or savanna, the restricted distribution patterns of the forest genera opposed to the widespread distribution patterns of the savanna genera

indicate that the forest adaptation set was primary,

the savanna adaptations secondary.

GRYPISCINES.—All 10 genera are limited to the

southeastern Brazil radiation center, although

Hylodes has been reported from the Guianas. Rivero (1968) described Elosia ditidensis from Mt.

Duida, Venezuela, as the only member of the genus

Elosia to occur outside of southeastern Brazil.

Lynch (1971) pointed out that Hylodes is the

proper generic name for Elosia. Rivero (1968) did

not examine the specimens for myological or osteological information. Clearly, he placed his new

species in Hylodes on the basis of dorsally divided

disks and fringed toes. Dorsally divided disks are

found in Hylodes, Crossodactylus, and Megaelosia

of the grypiscines, and Lithodytes of the leptodactylines, and I have observed the state in some

Eleutherodactylus of southeastern Brazil. Rivero

(1968) commented on the Eleutherodactylus-like

appearance of his new species. The evidence suggests to me that duidensis is a member of the

Eleutherodactylus-complex and not a Hylodes. All

grypiscines are found in forested habitats and further, eight of the genera are adapted to forest

stream life in one way or another. It is this overall

forest stream adaptational complex that convinces

me that the grypiscines are a natural unit.

ELEUTHERODACTYLINES.—The eleutherodactylines

are the most diverse of the groups. Members are

found in all radiation centers except the Chaco,

and some genera are endemic in five of them. The

major ecological adaptive types are to arid, forest,

and tropical montane environments. The ancestral

home and adaptive type of the eleutherodactylines

centers upon the origin of terrestriality. As argued

more fully later, I believe the origin to be in the

forests of southeastern Brazil.

HISTORICAL ZOOGEOGRAPHY

Savage (1973) summarized the major biogeographic patterns of living frog families, including

the family Leptodactylidae. Savage listed the following events as being of prime importance on

the diversification of leptodactylid frogs: the splitting up of Gondwanaland, first into east and west

portions in middle Jurassic, and a second splitting

of continents in the early Cretaceous; and the diversification of climatic and vegetation zones in the

Paleocene, including the appearance of xeric climates and vegetations. The most parsimonious

blend of radiation centers, ecological adaptive

types, continental changes, climatic and vegetational changes, and Savage's (1973) arguments is

represented in the diagram of Figure 11. The overall trend is clear: the family had its origin in the

beech forests of temperate South America, where a

remnant of that original stock still remains. Two

stocks became adapted to drying conditions, the
NUMBER 199 41

RECENT

PLEISTOCENE

PLIOCENE

MIOCENE

OLIGOCENE

EOCENE

PALEOCENE

LATE

CRETACEOUS MIDDLE

F V

\

NEOTROPICAL TERTIARY

GEOFLORA

(HYLIDS) ANTARCTO - TERTIARY

" ^ GEOFLORA

EARLY (RIFTS) HELIOPHRYNIDS LEPTODACTYLIDS

TO MYOBATRACHIDS.

MIDDLE (RIFT) \ PELODRYADIDS

EARLY t LIOPELMATO-LEPTOOACTYLOID

TRIASSIC LIOPELMATOID

FIGURE 11.—Proposed historical zoogeography of the five

leptodactylid groups, (A = arid, F = forest, M = montane,

s = savanna; also see text.)

ceratophrines and the leptodactylines. The grypiscines represent a forest-stream adaptation that centered in southeastern Brazil, and the eleutherodactylines most likely had their origin as an early

grypiscine stock. Before discussing each of the group

patterns in more detail, it is instructive to compare

and contrast the pattern proposed thus far (Figure

11) with previous proposals.

Vuilleumier (1968) was the first to point out

that the beech forest frog fauna of South America

was neither relictual nor depauperate but rather

was a consequence of a long and complex history.

Vuilleumier (1968) also suggested the frog fauna

of the beech forests was represented by four historical units. Lynch (1973a) contradicted the validity of Vuilleumier's four historical units and

stated that his ". . . analysis must be rejected because his conclusions are in part based upon the

erroneous conclusions of other authors" (p. 214).

Actually, Lynch concurred with Vuilleumier's major contention, as Lynch proposed that the origin

of the family was in the beech forests of temperate

South America.

All recent studies (Vuilleumier, 1968, Lynch,

1971, Savage, 1973) concur that the family originated in temperate South America. There is considerable difference of opinion as to how the

ancestral stock to the Leptodactylidae arrived in

its ancestral home. Lynch (1971) considered (1)

that the leptodactyloids were closely related to the

pelobatids, (2) that the pelobatids arose in the

Northern Hemisphere, and (3) that the leptodactylids represented a southward migration from

the Northern Hemisphere through the tropics to

the South Temperate Zone. Lynch (1971) based

the close pelobatid-leptodactylid relationships on

shared primitive states, a concept I philosophically

reject. As shown by Savage (1973) the pelobatids

and leptodactylids have little relationship in common.

Savage (1973) also proposed a holarctic migration for the leptodactylids, but from a discoglossoid

ancestor. As is clear from his diagrams (pp. 400-

401), deriving the leptodactylids in situ from a

leiopelmatid ancestor is a distinct possibility, in

fact, one that Savage has considered (pers. comm.).

The proposed leiopelmatid-leptodactylid relationship has a distinct zoogeographical advantage—it

does not require the unseemingly long migration

of a north temperate group across many thousands

of miles of tropics to the southern temperate region,

only to expand and differentiate northward again.

An in situ origin of leptodactyloids from a leiopelmatid ancestor also simplifies the zoogeographic

origin of the bufonids and ranoids (Savage, pers.

comm.). Savage (1973) based a northern leptodactyloid origin on tadpole evidence. Savage (1973)

found that Starrett's (1973) proposal of four suborders of frogs based upon larval characteristics

makes zoogeographic sense. Only two of Starrett's

(1973) tadpole types need be discussed for present

purposes. Both larval types have denticles and

beaks for scraping food sources from a substrate

and placing the foodstuff into temporary suspension. Both types of tadpoles differ from other tadpoles in: (1) feeding actions are more separated

from respiration, (2) presence of a long coiled gut

for algae feeding, (3) presence of extra jaw cartilages, muscles, and accessory mouth structures for
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

feeding, (4) smaller branchial chambers, and (5)

forelimbs that develop close to branchial chambers.

There is little doubt that both kinds of tadpoles

had an ancestral condition in which food was eaten

that was already in suspension in the water column.

The two tadpole types differ in that the lemmanuran tadpoles (Ascaphidae, Discoglossidae) have

separate branchial chambers with median external

openings, have forelimbs that develop close to the

branchial chambers, and have one more jaw muscle than the acosmanuran larvae (Pelobatidae,

Leptodactyloids, Bufonids, Ranoids), which have a

single branchial chamber with a sinistral external

opening and forelimbs that develop within branchial chambers. Starrett (1973) indicated and

Savage (1973) followed that the acosmanuran

type of tadpole is monophyletic. If it arose only

once from an ancestor with a lemmanuran type of

tadpole, then all Southern Hemisphere acosmanuran families had to have an ultimate origin in the

Northern Hemisphere. Obviously, Savage (1973)

was convinced that the acosmanuran tadpole is

monophyletic. One reason for suggesting the leiopelmatid-leptodactyloid relationship is to suggest

a reason to expect the acosmanuran type of tadpole to have originated twice.

As Savage (1973) points out, during the Jurassic,

the families of frogs had a distinctive distribution

pattern with respect to tadpole types. The North

and South Temperate zones were populated by

families with lemmanuran or acosmanuran types

of tadpoles, while all northern and southern tropical families had beakless larval types (Xenoanura

and Scoptanura of Starrett, 1973). The development of denticles and beaks and associated structures for scraping food into suspension doubtless

represents a considerable expenditure of energy.

This extra energy needed for a specialized feeding

apparatus is reflected in a longer metamorphic

time for denticled and beaked tadpoles as opposed

to beakless tadpoles (Heyer, 1973). If extra energy

is required for the specialized feeding apparatus of

the lemmanuran and acosmanuran tadpoles, then

there must be certain kinds of environments where

this kind of feeding apparatus has a distinct selective advantage. There appear to be two kinds of

habitats in which a scraping and chewing apparatus is required to exploit aquatic primary productivity. The first is stream environments, in which

the algae must be scraped off rocky substrates. The

second is temperate temporary or permanent ponds.

The water in such ponds is often clear; the primary

productivity is in the form of periphyton. In contrast, temporary tropical ponds in seasonal environments are usually turbid; there is much primary

productivity suspended in the water column. The

acosmanuran tadpole probably arose in temporary

temperate ponds, judging by the distribution of

larval types in the Jurassic. If this was the case, it

would make sense that similar selective forces, operating on similar sorts of beakless tadpoles continents apart, would result in similar adaptive kinds

of tadpoles. In other words, the acosmanuran tadpole may have arisen twice; once in the Pelobatids

in the temperate Northern Hemisphere, and a second time in the liopelmatids in the temperate

Southern Hemisphere. Unfortunately, living Liopelma are no help; all three living species have a

derived life history pattern of direct development

and no fossil liopelmatid larvae have been found.

The crux of the argument hinges, then, on whether

the pelobatid acosmanuran tadpole is really the

same as the leptodactyloid (bufonid, and ranoid)

acosmanuran tadpole. Starrett (1973) in her review

gives no indication with respect to resolution of

this question. I think that given the specialized

organism represented by a beakless xenoanuran

type of tadpole, there are an extremely limited

number of ways one can functionally mold such a

midwater suspension feeder into a scraping and

chewing feeder. If this is true, then one might not

expect to find great morphological differences

among acosmanuran types of tadpoles that had

been independently derived from xenoanuran sorts

of larvae. In comparing lemmanuran and acosmanuran types of larvae, it is obvious that there

are greater similarities that unite them and lesser

kinds of differences that differentiate them. I think

the characteristics that unite both kinds of larvae

and set them apart from other types are the minimum functional adaptations required for a scraping

and chewing existence. Further, the lemmanuran

type of tadpole might well be diphyletic, arising

once in the stream environment (Ascaphidae) and

once in the temperate pond environment (Discoglossidae). In summary, I would not expect

there to be great differences in functional morphology between pelobatid tadpoles and leptodactyloid tadpoles if they are diphyletic, given there

are such a limited number of ways, perhaps one, to
NUMBER 199 43

be a really efficient scraping and chewing feeder

based on a beakless, xenoanuran sort of tadpole

type. Rather, if there are consistent small differences in the way the single branchial chamber is

constructed or in how the forelimbs develop within

the branchial chambers of pelobatid tadpoles vs.

leptodactyloid, bufonid, and ranoid tadpoles, I

would argue for a separate origin of the larvae. To

my knowledge, such data is not available at present,

or if available, certainly not summarized.

Savage (pers. comm.) indicates that even if the

lemmanuran tadpole is monophyletic, a southern

origin of the Pelobatidae is not unreasonable.

The proposed events depicted diagrammatically

on Figure 11 during Middle and Late Jurassic and

Early Cretaceous assume that with the advent of

continental rifting, the evolutionary stocks diverged

as units on each of the continents. This is a reasonable assumption for those leptodactyloid groups

not presently associated with beech forests. Evidence

is accumulating that the beech forests of Antarctica,

South America, and Australia were contiguous up

until Paleocene and/or Eocene (Frakes and Kemp,

1972; Foster, 1974). Once the relationships among

the Australian and South American leptodactyloids

are better understood, it is possible that beech

forest faunal components in Australia and South

America will be found to be more closely related

to each other than either are to other Leptodactyloid groups.

WITHIN-GROUP PATTERNS

TELMATOBINES (Figure 12).—The telmatobines

represent an ancient radiation that was essentially

confined to the south Andean radiation center. Only

one genus occurs in the north Andean radiation

center. The radiation has been moderate resulting

in 8 genera and about 46 species. By far the most

successful genus in terms of numbers of species

is Telmatobius, with about 30.

CERATOPHRINES (Figure 13).—The primary radiation of the ceratophrines was in the Chaco type

of habitat with a secondary radiation in southeastern Brazil. The genus Ceratophrys has spread to the

Amazon region and there is one species in the

west coast lowlands of South America. The presence

of Ceratophrys on the west coast of South America

is most likely due to its presence there prior to the

uplifting of the north Andes, rather than an inTELMATOBINES

NA

1

SA

<!

8

•>

FIGURE 12.—Proposed telmatobine radiation. (Number in

circle is number of endemic genera; numbers at bottom are

total number of genera; abbreviations as in legend for Figure

10.)

vasion after the Andes had been uplifted. The

ceratophrine radiation has been rather small, resulting in about 5 genera and 22 species.

GRYPISCINES (Figure 14).—The grypiscine radiation took place in southeastern Brazil where all

members are endemic. The overall adaptation of

the group is to the forest stream habitat. Within

this general adaptation framework, the most noticeable evolutionary trends are toward terrestriality

(direct development) (e.g., Zachaenus) and arboreality (Crossodactylodes). The grypiscine radiation is an old and moderate radiation, which resulted in about 10 genera and 37 species.
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

GRYPISCINES

FIGURE 13.—Proposed ceratophrine radiation. (Numbers in

circles are numbers of endemic genera; numbers at bottom

are total number of genera; solid circle = primary radiation;

dashed circle = secondary radiation; abbreviations as in

legend for Figure 10.)

LEPTODACTYLINES (Figure 15).—The primary

adaptation of the leptodactylines was to the Neotropical Tertiary Geoflora. The remnants of that

radiation are found in the derived forests of Amazonia and southeastern Brazil. The size of this primary radiation has been small, resulting in 6

genera and 11 species. The secondary radiation of

savanna adapted forms was much more successful

and recent, reflected in the fact that the four savanna genera are widespread, none are endemic

to a single radiation center, and they are represented by about 85 species. Representatives of the

FIGURE 14.—Proposed grypiscine radiation. (Number in circle

is number of endemic genera; number at bottom is total

number of genera; abbreviations as in legend for Figure 10.)

savanna genera were able to island hop to the West

Indies, but have not radiated there. Three species

of Leptodactylus are represented on the Greater

and Lesser West Indian islands, which are more

closely related to other South American species

than to each other. This would indicate that the

leptodactylines were the last leptodactylids to arrive

in the West Indies. All of the leptodactyline species

found in Mexico (2 Leptodactylus and 1 Physalaemus) and Middle America (5 Leptodactylus and

1 Physalaemus) are also found in South America.

This indicates that the leptodactylines arrived in

Middle America and Mexico relatively recently.

This agrees with Savage's (1966) scheme in which
NUMBER 199 45

LEPTODACTYLINES

FIGURE 15.—Proposed leptodactyline radiation. (Numbers in

circle are numbers of endemic genera; numbers at bottom

are total number of genera; solid circles = primary radiations, dashed circle = secondary radiation; abbreviations as

in legend for Figure 10; also see text.)

a South American element penetrated Middle America when the land bridge between Middle and South

America became reconnected in the Pliocene.

Closely tied in with the success of the savanna

adapted leptodactylines is the foam nest. Lynch

(1971:220) suggested that the leptodactyline foam

nest evolved in response to decreasing equabilities

during the Cretaceous. He indicated that the presence of the foam nest allowed the leptodactylines

to breed in more xeric environments. I have suggested that the foam nest in Leptodactylus was an

adaptive response to aridity (1969b, 1974a), but

that the foam nest in Adenomera could not be

accounted for as an adaptation to arid environments, as the genus is primarily distributed in wet

forest (Heyer, 1974a). For Adenomera, I suggested

that the foam nest was part of an adaptive complex leading to direct development. The results of

the present analysis indicate that the foam nest had

its evolutionary origin in wet forests and that its

original adaptive value was in response to the wet

forest environment, such as is demonstrated by the

genus Adenomera (Heyer, 1974a). This type of

foam nest was a preadaptation, then, for the group

of frogs which adapted to the drying environments

during late Paleocene and Miocene. In other words,

the key feature which allowed the savanna leptodactylines such success in adapting to the arid environment, as opposed to members of the other

major generic groupings, was the presence of the

foam nest, which was preadaptive to the savanna

environment.

ELEUTHERODACTYLINES (Figure 16).—The eleutherodactylines have undergone an explosive radiation, some of which may well be continuing at the

species level. The history of the eleutherodactylines begins as a grypiscine stock in the forests of

southeastern Brazil. The key feature which allowed

the explosive radiation was clearly the early evolution of direct development. The remnants of this

primary radiation remain in southeastern Brazil as

two endemic genera containing three species. Not

indicated on the diagram of Figure 16 is the diversity represented by Eleutherodactylus. The composite genus is found almost wherever eleutherodactylines are found. There is a secondary center

of radiation in Amazonia, most probably indicated

by at least one endemic genus. The minor radiation

of the northern Andes is clearly derived from an

Amazonian ancestral stock, and one of the northern

Andean genera has penetrated the northern end of

the southern Andes. A minor radiation also took

place on the west coast of South America. As in

the ceratophrines, the presence of eleutherodactylines on the west coast of South America is most

likely because they were there prior to the uplifting

of the Andes. There appear to be at least two radiations of the eleutherodactylines in Mexico and

Middle America. The first is represented by Hylactophryne and appears to be the earliest radiation.

The second radiation is represented by endemic

Mexican and Middle American Eleutherodactylus,
46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ELEUTHERODACTYLINES

FIGURE 16.—Proposed eleutherodactyline radiation. (Numbers in circles are numbers of endemic genera; numbers at

bottom are total number of genera; solid circle = primary

radiation; dashed circles = secondary radiations; abbreviations as in legend for Figure 10; also see text.)

Syrrophus, and Tomodactylus. Middle and South

America were isolated from each other from Paleocene through Miocene and were not connected

until Pliocene. The Pliocene is too late for the

beginning of the kinds of radiation found. This

indicates that the eleutherodactyline stock that gave

rise to the two radiations island hopped while Middle and South America were isolated. This is not

unreasonable, as eleutherodactylines are good island hoppers as evidenced by their occurrence on

almost all West Indian islands. The West Indian

eleutherodactyline fauna appears to be the result

of two invasions, an early one, represented by the

monotypic Sminthillus in Cuba. Sminthillus is

quite specialized; it may be that the ancestral stock

was also specialized, which prevented it from an

explosive radiation as in Eleutherodactylus. The

second, more recent invasion was by an Eleutherodactylus-complex member. Further study may indicate at least two invasions by Eleutherodactyluscomplex members. The explosive radiation of the

eleutherodactylines is with the Eleutherodactyluscomplex, which has about 350 species, most of

which are in the West Indies, Middle America, and

the slopes of the northern Andes.

ORIGIN AND EVOLUTION OF TERRESTRIALITY

The key to the spectacular success of the Eleutherodactylus-complex lies in the evolution of direct development, giving the frogs a completely

terrestrial life history. Only the broadest features

of this radiation can be outlined at this time due

to the poor understanding of the composition of

the Eleutherodactylus-complex.

In order to trace the origin of direct development,

it is necessary to comment on the adaptive significance of the complex life cycle in frogs. The following argument is heavily influenced by the work

of Richard Wassersug, who has discussed these

points with me. The basic adaptive significance of

the frog larvae is to exploit bursts of primary

productivity that occur in temporary ponds. This

is especially adaptive in seasonal tropical environments, and it appears that the origin of frogs was

in seasonal tropical situations. In a seasonal tropical environment, there are predictable physical

signals which frogs can interpret that will insure

reproductive success. For instance if a frog is buried

a certain distance under the soil surface during

the dry season, it takes a considerable rainstorm

for water to reach the underground point where

the frog is buried. Once water reaches the frog,

the frog can then burrow out of the soil and reproduce, and the chances are very good that the

same rainfall that was sufficient to initiate activity

of the frog will form ponds that will last long

enough for the larvae to metamorphose. Thus, in

seasonal tropical environments the complex life cycle is adaptive, both from an energy utilization

viewpoint and a predictive aspect that will insure

reproductive success.
NUMBER 199 47

The wet forests pose two kinds of problems to

the complex life cycle. The first is that there is

generally no primary productivity bloom in temporary wet forest ponds, because the release of organic matter by the vegetation is rather continual,

not seasonal. The second is that there are no real

signals coming from the physical environment that

will insure reproductive success. For instance, Inger

(1969) has shown that large thundershowers in

Borneo are random events. Thus, rainfall pattern

does not give any information which would indicate that breeding at one particular rainfall would

have a greater likelihood of raising larvae through

metamorphosis than any other rainfall. Thus the

complex life cycle, as adapted for tropical seasonal

environments, is not well adapted for the wet forest environment. I would think that a wet forest

environment in these terms could be defined as

an environment with a drier season lasting no more

than two or three months. Thus, any adaptation

toward a terrestrial life cycle would have a selective advantage in the wet forest environment.

This trend toward terrestriality is seen in the

present-day grypiscines, such as Zachaenus. It is interesting to note that in the case of the grypiscines,

terrestriality has come out of a basic forest-stream

adaptational complex. The crux of my argument

for proposing that the eleutherodactylines are an

early grypiscme derivative that attained direct development is based on the supposition that terrestriality originated in a wet forest habitat, and

the grypiscines, in such an environment, demonstrate repeated trends toward terrestriality. The

eleutherodactylines, once they had attained direct

development, were able to radiate into a wide variety of ecological zones.

The development of terrestriality in the family

Leptodactylidae contains two major features. The

first is that the eleutherodactylines, with direct development, have been able to invade ecological

adaptive zones unavailable to the other leptodactylid groups. The second point is that terrestriality

has been essentially reached within each of the

major groups at least once, with the exception of

ceratrophrines. It is interesting to speculate why

the development of terrestriality only led to a radiation in the case of the eleutherodactylines. The

telmatobine that approaches terrestriality is Batrachyla. Apparently it lacked the genetic flexibility

to move out of the beech forests, and terrestrial

possibilities within a beech forest are limited. The

grypiscine forms that are essentially terrestrial are

such things as Cycloramphus and Zachaenus. These

are results of continued selective advantages for

wet forest frogs, and terrestriality has been approached in this group after the eleutherodactyline

radiation occurred; thus there was no ecological

access for the terrestrial grypiscines. The same appears to be true for Adenomera, the terrestrial leptodactyline genus. Its development of terrestriality

as a wet forest inhabitant was preceded by the

eleutherodactyline radiation. An alternative explanation is that the key feature to the success of

eleutherodactylines was the encapsulated egg,

which is a more successful mode of terrestriality

than the modes seen in Batrachyla, Cycloramphus,

Zachaenus, and Adenomera, which do not have encapsulated eggs. This alternative explanation could

allow the evolution of terrestriality in the eleutherodactylines to have occurred after the origin

of terrestriality in the other groups.

Only with the eleutherodactylines did direct development arise in conjunction with a group

which had genetic plasticity and ecological and

evolutionary access to form a spectacular adaptive

radiation. Perhaps it is more than coincidental that

the eleutherodactylines form a parallel example

to the terrestrial plethodontids which, from a stream

adapted ancester which evolved direct development,

underwent an explosive adaptive radiation, much

of which was centered in the New World tropics

(Wake, 1966).
Appendix: Supplementary Data

SPECIMENS EXAMINED.—The following specimens

supplement the data given by Lynch (1971) and

Heyer (1974).

LEPTODACTYLIDAE

Batrachophrynus macrostomus, USNM (in National Museum of National History, Smithsonian Institution, under

catalog numbers of former United States National Museum) 118172 muscle dissection (M), KU (University of

Kansas, Museum of Natural History) 9817 dry skeleton (S).

Batrachophrynus patagonicus, USNM 154170 (M).

Batrachyla leptopus, USNM 196279 (M).

Batrachyla taeniata, USNM 196282 (M).

Caudiverbera caudiverbera, USNM 139788 (M).

Ceratophrys calcarata, USNM 146952 (M).

Crossodactylus gaudichaudii^USNM 164105 (M).

Crossodactylus dispar, USNM 129376 (M).

Cycloramphus fuliginosus, USNM 164121 (M).

Cycloramphus dubius, USNM 129376 (M).

Eleutherodactylus coqui, USNM 86565 (M), USNM 192321

cleared and stained skeleton (CS).

Eleutherodactylus fleischmanni, USNM 67333 (M), KU

68158 (S).

Eleutherodactylus nigrovittatus, USNM-JAP 9140 (M),

USNM-GOV 8108 (CS).

Euparkerella brasiliensis, USNM 196288 (M), KU 93192

(CS).

Eupsophus grayi, USNM 139791 (M).

Eupsophus nodosus, USNM 154203 (M).

Holoaden bradei, USNM 196287 (M), KU 107088 (CS).

Hylactophryne augusti, USNM 116420 (M), KU 56187 (CS).

Hylodes aspera, USNM 129155 (M).

Hylodes nasus, USNM 164114 (M).

Jschnocnema quixensis, USNM 194793 (M), KU 104388 (S).

Lepidobatrachus llanensis, USNM-WRH 1360 (M), KU

129706 (S).

Megaelosia goeldi, USNM 96763 (M).

Niceforonia festae, USNM 160957 (M).

Odontophrynus americanus, USNM 123400 (M).

Odontophrynus cultripes, USNM 81138 (M).

Paratelmatobius lutzi, KU 107089 (CS).

Proceratophrys appendiculatus, USNM 12814 (M).

Sminthillus limb at us, USNM 136090 (M), KU 68684 (CS).

Syrrophus campi, USNM 52373 (M).

Syrrophus leprus, USNM 114087 (M).

Telmatobius hauthali, USNM 93208 (M).

Telmatobius jelski, USNM 61162 (M).

Thoropa miliaris, USNM 97765 (M).

Thoropa petropolitana, USNM 164135 (M).

Tomodactylus angustidigitorum, LACM (Natural History

Museum, Los Angeles County) 25520 (M).

Tomodactylus nitidus, USNM 114098 (M).

Zachaenus parvulus, USNM 164154 (M).

Zachaenus stejnegeri, USNM 164116 (M).

HELIOPHRYNIDAE

Heliophryne natalensis, KU 105925 (CS).

Heliophryne purcelli, USNM 162428 (M).

MYOBATRACHIDAE

Adelotus brevis, AMNH (American Museum of Natural

History) 59096 (M), KU 147213 (S).

Crinia signifera, USNM 167710 (M), KU 56245 (CS).

Cyclorana australis, USNM 128237 (M).

Glauertia orientalis, AMNH 67394 (M), AMNH 128276 (M).

Heleioporus albopunctatus, USNM 84127 (M).

Kyarranus sphagnicola, AMNH 60707 (M), KU 110331 (CS).

Lechriodus melanopyga, USNM 195572 (M).

Limnodynastes dorsalis, USNM 118774 (M).

Limnodynastes peroni, USNM 167709 (M).

Mixophyes jasciolatus, KU 147227 (S).

Myobatrachus gouldi, AMNH 46052 (M), KU 110333 (CS).

Neobatrachus pictus, AMNH 59114 (M).

Notaden nichollsi, AMNH 67178 (M), KU 93582 (CS).

Philoria frosti, AMNH 67357 (M), KU 50699 (CS).

Pseudophryne bibroni, USNM 167712 (M), KU 83588 (CS).

Taudactylus acutirostris, KU 124233 (CS).

Uperoleia marmorata, AMNH 60643 (M).

Uperoleia rugosa, KU 109861 (CS).

48
NUMBER 199 49

Genera

UTTODACTYLUftE

Amblyphrynus

Barycholos

Batrachophrynus

Batrachyla

Oaudlverbera

Ceratophrys

Croaaodactylodes....

Croesodactyl.ua

Cycloraaphus

Edalorhlna•••••••.. .

E. coqui

E. flelschmanni

E. nlgrovlttatus

Eaparkerella

Eusophus

Holoaden

Hydrolaetare

Hylactophryne

Hylodes

Hylorlna

Insuetophrynus

Ischnocneaa

Lepldobatrachus

Leptodactylua

Lionomedusa

Llthodytes

Megaelosla

Odontophrynus

Paratelmatoblus

Fhysalaemua

TABLE A.—Ecological and geographical categories for the study sample

Table A. — Ecological and geographical categories for the study sample

Middle West Coast northern Southern Guiana S.E. West

Fossorlal Terrestrial Aquatic Arboreal Africa Australia Mexico America S. America Andes Andes Shield Amazonia Chaco Brasil Indies

(x)
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE A.—Ecological and geographical categories for the study sample (continued)

Genera

Pleurodeaa

Procerataphrys..

Pseudopaludlcoli

Sminthillus

Syrrophua

Telnatoblus

Telaatobof o....

Thoropa

Itonodactylus...

Vantollnius

Zachaenus

CYCLORATDIAB

Adelotas

Cyclorana

He lloporus

Kyarranus

Lechrlodus

Liooodynastes..

Mlxophyea

Heobatrachus...

NotadeD

Philorla

MTOBATHACHIHAE

A«sa

Crlnla

Olauertia

Metacrlnla

Myobatnchus

Paeudophryne...,

Taudactylua....

Uperolla

HELZCPSIXHZM2

Helloptaryn*....

Middle West Coast Rorthern Southern Guiana S.E. West.

Foaaorlal Terrestrial Aquatic Arboreal Africa Australia Mexico Aaerica S. Aaerlca Andes Andes Shield Anazonla Chaco Brasll InJics
NUMBER 199 51

TABLE B.—Generic character states (arranged by character number) used in character analysis

Genus

Adenoraera

Aoblyphryniis

Barycholos

Bstrcichophryniis

Batrachyla

Caudiverbera

Ceratophrys

Crossodactylodes

Crossodactylus

Cycloranphus

Edalorhlna

E. coqui

E. flelschmannl

Euparkerella

Eupsophus

Boloaden

Hydrolaetare

Hylactophryne

Hylodes

Hylorlna

Insuetophrynus

lachnocnema

Lepldobatrachus

Leptodactylua

LlnzionedUBa

Llthodytes

Megaelosla

Odontophrynua

Paratelnatobius

Phyaalaemus

Pleurodena

Procera tophrya

Pseudopaludlcola

Sulnthlllus

Syrrophus

Telmatobius

TelBBtobufo

Thoropa

Tooodactylus

Vanvnl ini IIR

VBDZOllIllUE

Z. parvulus

Z. stejnegerl

Genus

Adenoaera

Aablyphrynus

Barycboloa

Batrachophrynua

Batrachyla

Caudiverbera

Ceratophrys

Crossodactylodes

Croasodactylus

Cycloraapous

Edalorhlna

E. coqul

E. flelactaaannl

E. nlgrovlttatus

Euparkerella

Eupsophus

Holoaden

Hydrolaetare

Hylactophryne

Hylodes

Hylorlna

Insuetophrynus

Ischnocneaa

Lepldobatrachus

Leptodactylua

Llmnoaedusa

Llthodytes

Megaelosla

Viceforonia

Odontophrynua

ParatelaatobluB

Physalaesua

PleurodeaB

Procera tophrya

Pseudopaludlcola

Sninthlllus

Syrrophus

Telmatobius

Telnsatobufo

Thoropa

Tooodactylus

Vanzollnius

Z. parvulus

Z. stejnegerl

1

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F

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A

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F

F

B

A

F

A

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19

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D

A

D

D

D

G

I

G

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D

B

A

G

D

G

A

B

D

F

D

0

D

B

E

B

D

D

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37

C

B

B

A

B

B

G

B

I

C

G

B

B

B

C

B

G

C

I

B

F

G

G

B

A

E

B

G

G

B

20

A

B

C

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A

B

A

A

B

A

B

B

D

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C

B

B

A

A
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE B.—Generic character states (arranged by character number) used in character analysis

(continued)

Qeniu 10 13 it 18 19

Adelotus

Cyelorans

Helloporus

Kyarramis

Lechrlodus

Llmnodynastes

Mlxophyes

Neobatrachus

Notaden

Phllorla

Assa

Crinla

Glauertia

Metacrinla

Myobatrachus

Pseudophryne

Taudactylus

Upe rolls

Heliophryne

A ;

A t

B/D /

A /

A 1

A /

A 1

A /

B /

A /

A

A i

A

F

A

I B

B

I A

k A

V A

I A

I A

V A

I B

!, A

\ A

i A

B

A/B

A

A

A

A/B

A

A

A

A/B

A

A

A

A

A

B

B

A

A

C

B

B

A

A

C

A

A

A

A

A

A /

C

C

B

A

A

C

E

A

C

C

C

B/D

B/C/F

A/B/C

A/D A/B/C/D/F

A/F

B/C/F

A/D/F

B/D

A/B/D

A/D

A/F

B/C

B/C

B/E

A/D

A/D

A/F

Genus

Ade lotus

Cyclorana

Helioporus

Kyarranus

Lechriodus

Llonodynastes

Mixophyee

Neobatrackus

Notaden

Philorla

Assa

Crinla

Olauertla

Metacrinla

Myobatrachus

Pseudophryne

Taudactylus

Uperolla

Hellophryne

23 2k

A/H
NUMBER 199 53

TABLE C.—Generic character states (arranged by

Genus

^denomera

Anblyphrynus

Barycholos

Batrachophrynus

Batraehyla

Caudlverbera

Ceratophrys

Crossodactylodes

Crossodactylus

Cycloramphus

Edalorhina

E. coqul

E. flelschmannl

E. nigrovittatus

Euparkerella

Eupsophus

Holoaden

Hydrolaetare

Hylactophryne

Hy lodes

Hylorlna

Ineuetophrynus

Ischnocnema

Lepldobatrachus

Genus

Adenamera

Arablyphrynus

Barycholos

Batrachophrynus

Batraehyla

Caudlverbera

Ceratophrys

Crossodactylodes

Crossodactylus

Cycloramphus

Edalorhina

E. coqui

E. fleishmanni

E. nigrovittatus

Euparkerella

Eusophus

Holoaden

Hydrolaetare

Hylactophryne

Hylodes

Hylorina

Insuetophrynus

Ischnocnema

Lepidobatrachus

Genus

Leptodactylus

Limnomedusa

Lithodytes

Megaeloeia

JTiceforonia

Odontophrynus

Paratelmatobius

Physalaemus

Pleurodema

Proceratophrys

Pseudopaludicola

Smlnthillus

Syrrophus

Telmatobius

Telmatobufo

Thoropa

Tomodactylus

Vanzolinius

Z. parvulus

Z. stejnegerl

Genus

Leptodactylus

Limnonedusa

Lithodytes

Hegaelosia

Hicef ronla

Ode tophrynus

Paratelmatoblus

Physalaemus

Pleurodema

Proceratophrys

Pseudopaludicola

Sminthlllus

Syrrophus

Telmatoblus

Teliiatobufo

Thoropa

Tomodactylus

Vanzollnius

Zachaenus parvulus

Z. stejnegerl

1

0

0

0

0

1

0

0

0

0

0

0

0

0

0

0

0

1

0

0

1

0

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1

18

0

0

0

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3

0

1

1

2

0

0

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3

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0

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5

0

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0

1

0

0

1

3

0

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2

2

2

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0

0

0

0

3

0

0

0

0

22

0

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0

1

0

0

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1

0

0

0

0

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0

1

0

0

1

0

0

0

5

0

0

3

3

0

0

0

0

2

0

0

1

2

1

0

0

2 2

1

1

0

0

0

0

0

0

0

0

1

0

0

1

1

0

0

0

0

6

0

1

0

0

0

0

0

2

0

1

0

0

0

0

0

0

0

0

2

0

0

23

0

1

1

0

0

2

2

1

0

1

0

0

1

1

1

0

0

1

0

0

0

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23

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