Analysis of the Intcrgencric

Relationships of the Australian

Frog Family Myobatrachidae

W. RONALD HEYER

and

DAVID S. LIEM

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 233
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SMITHSONIA N CONTRIBUTION S T O ZOOLOGY • NUMBER 233

Analysis of the Intergeneric

Relationships of the Australian

Frog Family Myobatrachidae

W. Ronald Heyer

and David S. Liem

SMITHSONIAN INSTITUTION PRESS

City of Washington

1976
ABSTRACT

Heyer, W. Ronald, and David S. Liem. Analysis of the Intergeneric Relationships of the Australian Frog Family Myobatrachidae. Smithsonian Contributions

to Zoology, number 233, 29 pages, 28 figures, 2 tables, 1976.—Forty-two characters

of external morphology, myology, osteology, and life history 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 primarily Australian frog family Myobatrachidae. Three

major groupings of myobatrachids are proposed; these are accorded subfamilial

status. One of the subfamilies is defined as new. In addition, three genera are

proposed as new.

It is proposed that the ancestral myobatrachid stock was associated with the

Antarcto-Tertiary Geoflora. Several genera still demonstrate a distribution associated with this beech-podocarp vegetation. Major evolutionary events in the

family have been associated with stream adaptations, trends toward and including direct development and fossorial adaptations. Convergence in these events

with the leptodactylid frog fauna of southern South America is commented

upon.

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

in the Institution's annual report, Smithsonian Year. SERIES COVER DESICN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Heyer, W Ronald.

Analysis of the intergeneric relationships of the Australian frog family Myobatrachidae.

(Smithsonian contributions to zoology ; 233)

Bibliography: p.

1. Myobatrachidae. 2. Amphibians—Evolution. 3. Amphibians—Classification. 4. Amphibians

—Australia. I. Liem, David S., joint author. II. Title. III. Series: Smithsonian Institution.

Smithsonian contributions to zoology ; 233.

QL1.S54 no. 233 [QL668.E2615] 591'.08s [597'.8] 76-608054
Contents

Page

Introduction 1

Methods and Materials 1

Acknowledgments 2

Character Analysis 3

Analysis of Relationships 5

Sister Groups within the Phylogeny 10

Comparison with Previous Schemes 10

Systematic Conclusions 11

RHEOBATRACHINAE, new subfamily 11

MYOBATRACHINAE Schlegel, 1850 11

Crinia Tschudi, 1838 11

Geocrinia Blake, 1973 11

Pseudophryne Fitzinger, 1843 12

Ranidella Girard, 1853 12

Paracrinia, new genus 12

Australocrinia, new genus 12

Kankanophryne, new genus 12

Phyletic Character Analysis 12

Evolutionary Zoogeography 15

Generic Distribution Patterns 15

Evolutionary Trends within the Subfamilies 23

Subfamily RHEOBATRACHINAE 23

Subfamily MYOBATRACHINAE 23

Subfamily LIMNODYNASTINAE 24

Major Trends among Subfamilies 24

Comparisons with Previous Zoogeographic Schemes 25

Myobatrachid-Leptodactylid Comparisons 26

Literature Cited 28

in

Analysis of the Intergeneric

Relationships of the Australian

Frog Family Myobatrachidae

W. Ronald Heyer

and David S. Liem

Introduction

The intergeneric relationships of the Australian

frog family Myobatrachidae (Leptodactylidae of

most authors) have been analyzed recently. Lynch

(1971) used a standard systematic approach to the

problem while Liem (in press) used a numerical

taxonomy approach. Neither author used the

phylogenetic approach as outlined by Hennig

(1966). The purpose of this paper is to apply the

same methods of analysis of relationships to the

Australian myobatrachids as have been applied

recently to the Neotropical leptodactylids (Heyer,

1975a).

METHODS AND MATERIALS.—Character states are

categorized for a number 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.

The characters used are mostly those described

by Liem (in press). Liem's study included members of the family Pelodryadidae. Characters in

Liem's study which have uniform states in the

Myobatrachidae are not used. Some other of

Liem's characters, for which not enough comparable information is known in other frogs to deterW. Ronald Heyer, Division of Reptiles and Amphibians, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C.

20560. David S. Liem, Wildlife Branch, Department of

Natural Resources, P.O. Box 5140, Boroko, Papua, New

Guinea.

mine directionality of states, are also deleted.

The taxa used for analysis are those recognized

by Liem (in press) with certain exceptions. Blake

(1973) and Liem (in press) used similar numerical

techniques of analysis on some of the same genera

with conflicting systematic results. For purposes of

analysis, the smallest generic groupings are used

to test the hypotheses of Blake and Liem. In addition to the genera recognized by Liem (in press),

the following units are added: (1) Crinia georgiana, (2) Crinia haswelli, (3) Ranidella (as used

by Blake, 1973), and (4) Pseudophryne occidentalis.

The criteria used for determining directionality

of character states are the same as those used

previously (Heyer, 1975a). In those cases where

the character states of this study are the same as

those used in the leptodactylid study (Heyer,

1975a), the arguments for directionality are the

same and are not recounted here. This omission

is occasioned by the other study's having used the

myobatrachids as the major outgroup for determination of state directionality. A primitive state for

the leptodactylids can also be assumed to be the

primitive state for the myobatrachids if the

assumption is made that the myobatrachids and

leptodactylids had a common ancestor. The primitive state is always coded as zero in the character

analysis section.

Life history and geographic range data were

taken from the published literature (most of the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

papers included in "Literature Cited"), Ian R.

Straughan's unpublished dissertation (1966), and

personal observations (DSL).

ACKNOWLEDGMENTS.—We wish to thank Ian

Straughan, Southern California Edison Company

and George R. Zug, Smithsonian Institution, for

their careful reading of the manuscript.

Thanks are due also to Hobart M. Smith, University of Colorado, who originally suggested the

collaboration that produced this manuscript.

This work was supported by a Smithsonian

Research Foundation Award to the first author

and by ARGC grants (D70/17419 and D70/17559)

and the University of Queensland Research Allocation to the Department of Zoology to the second

author during his tenure at that Department.

Character Analysis

CHARACTER 1, maxillary teeth: state 0, maxillary

teeth present, pedicellate; state 1, maxillary teeth

present, fanglike, at least in some species; state 2,

maxillary teeth uniformly absent or absent in

some species.

Trueb (1973:79) indicated that states 1 and 2

are independently derived states from state 0. The

directions of change of character states are:

1 « 0 • 2

CHARACTER 2, nasal contact with frontoparietal:

state 0, nasals not in contact with frontoparietal;

state 1, nasals in contact with frontoparietal.

Heyer (1975a) indicated that state 0 is primitive.

CHARACTER 3, carotid canal: state 0, exposed

shallow channel or carotid canal absent; state 1,

exposed shallow canal present, at least in some

species; state 2, carotid canal present, at least in

some species.

Lynch (1971) suggested that the carotid canal

is formed by thickening of the frontoparietal

posteriorly and/or involving dermal coossification

resulting in the enclosure of the carotid artery in

a bony canal. Since the vast majority of frogs have

state 0, it is considered the primitive state (criterion II of Heyer, 1975a). The direction of change

of character states is:

0 • 1 • 2

CHARACTER 4, frontoparietal fontanelle: state 0,

frontoparietals separated medially, exposing fontanelle; state 1, frontoparietal fontanelle covered,

at least in some species.

Heyer (1975a) showed that state 0 is the primitive state.

CHARACTER 5, sphenethmoid: state 0, sphenethmoid entire; state 1, sphenethmoid divided, at

least in some species.

Trueb (1973) thought state 1 reflects reduced

ossification. Among the New World and

African leptodactyloids, only Euparkerella and

Tomodactylus have state 1 (Lynch, 1971). State

1 is considered the derived state (criterion II of

Heyer, 1975a).

CHARACTER 6, squamosal: state 0, zygomatic and

otic rami about equal length, both moderately

long; state 1, zygomatic ramus shorter than otic

ramus, at least in some species.

Heyer (1975a) showed state 0 to be primitive.

CHARACTER 7, vomer and vomerine teeth: state

0, vomer and vomerine teeth present uniformly;

state 1, intrageneric variation, all species with

vomer present, at least some species with vomerine

teeth absent; state 2, vomer present, vomerine

teeth absent uniformly; state 3, intrageneric variation, at least some species with vomer and vomerine teeth absent; state 4, vomer and vomerine

teeth uniformly absent.

Heyer (1975a) indicated state 0 to be the primitive state. The direction of change of character

states is:

CHARACTER 8, intervertebral disc: state 0, intervertebral disc fused to the posterior end of the

centrum (procoelous); state 1, intervertebral disc

free between centra of presacral vertebrae (amphicoelous).

Trueb (1973) considered amphicoely as the

probable ancestral frog condition, but thought it

likely that amphicoely in modern frogs is a

paedomorphic trait, hence derived. The African

leptodactyloid, Heleophryne, is amphicoelous, the

New World leptodactylids are uniformly precoelous. Thus, the closest outgroup members do

not clearly indicate which state is primitive.

Expansion of the outgroup clarifies the situation.

Bufonids, hylids, pseudids, dendrobatids, and

centrolenids are uniformly procoelous. State 0 is

considered the primitive state.

CHARACTER 9, cervical vertebrae: state 0, cervical
NUMBER 233

and second vertebrae free; state 1, cervical and

second vertebrae fused.

Trueb (1973) stated the fused condition to be

derived. Lynch (1971) found state 0 uniformly in

the African and Neotropical leptodactyloids. State

0 is considered the primitive state (criterion I of

Heyer, 1975a).

CHARACTER 10, cervical cotyles: state 0, cervical

cotyles narrowly separated; state 1, cervical cotyles

widely separated.

Heyer (1975a) indicated state 0 to be the primitive state.

CHARACTER 11, relationship of transverse process

of last presacral vertebra to sacral vertebra: state

0, last presacral vertebra noticeably shorter than

sacrum; state 1, last presacral vertebra about same

width as sacrum.

Heyer (1975a) indicated the primitive state to

be 0.

CHARACTER 12, sacral diapophyses: state 0, sacral

diapophyses expanded; state 1, sacral diapophyses

rounded uniformly, in at least some species.

Heyer (1975a) indicated the primitive state to

be 0.

CHARACTER 13, coccyx: state 0, prezygapophyses

and transverse processes present: state 1, prezygapophyses and transverse processes absent.

Trueb (1973) suggested state 0 to be primitive.

CHARACTER 14, dorsal crest of ilium: state 0, no

dorsal crest; state 1, well denned crest -present.

Heyer (1975a) indicated the primitive state to

be 0.

CHARACTER 15, carpals: state 0, first, second,

third carpale and naviculare free; state 1, at least

one element of state 0 demonstrating fusion.

Geocrinia victoriana is the only member of the

genus demonstrating state 1. Since all other species

have state 0, Geocrinia is coded as state 0. Trueb

(1973) and Liem (1970), following Howes and

Ridewood (1888), indicated state 0 to be the

primitive state.

CHARACTER 16, prehallux: state 0, prehallux of

1 to 3 segments with blunt or pointed distal segments; state 1, prehallux of one shovel shaped

segment, at least in some species.

Heyer (1975a) indicated state 0 to be primitive.

CHARACTER 17, cricoid: state 0, cricoid forming

a complete ring; state 1, cricoid not forming a

complete ring.

All Neotropical and African leptodactyloids

have state 0 (Lynch, 1971). State 0 is assumed the

primitive state (Criterion I of Heyer, 1975a).

CHARACTER 18, alary process of the hyoid: state

0, alary process broad and wing-like; state 1, alary

process stalked, distally expanded into rounded

or oval plates.

Heyer (1975a) considered state 1 to be the primitive state, but found it to be a poor predictor of

phylogenetic relationships. In a consideration of

the relationships among the species of the genus

Adenomera (Heyer, 1975b), the better assumption

appeared to be that state 0 is the primitive state.

State 0 is here considered the primitive state.

CHARACTER 19, anterior process of the hyale:

state 0, anterior process present; state 1, anterior

process absent.

Heyer (1975a) indicated state 0 to be primitive.

CHARACTER 20, depressor mandibulae muscle:

state 0, slip from dorsal fascia present; state 1,

intrageneric variation, some species with state 0,

others with state 2; state 2, no dorsal fascia slip.

The condition for Assa is not known and

assumed to be primitive for purposes of analysis.

Heyer (1975a) indicated that state 0 is primitive

relative to state 2 as described here. The direction

of change of character states is:

CHARACTER 21, adductor mandibularis muscle:

state 0, both adductor mandibulae posterior subexternus and adductor mandibulae externus superficialis present; state 1, adductor mandibulae

posterior subexternus only present; state 2, adductor mandibulae externus superficial only present.

Heyer (1975a) indicated state 0 is primitive.

Directions of change of character states are:

CHARACTER 22, supracoracoideus superficialis and

profundus muscles: state 0, muscles separate; state

1, muscles fused, at least in some species.

Jones (1933) demonstrated that state 0 is the

general state in frogs. State 0 is assumed the

primitive state (Criterion II of Heyer, 1975a).

CHARACTER 23, adductor longus muscle: state 0,

muscle inserting on knee; state 1, muscle inserting

on distal 1/j, to 1/3 of adductor magnus muscle or

absent.

Heyer (1975a) indicated that state 0 is primitive.

CHARACTER 24, semitendinosus muscle distal ten-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

don insertion: state 0, distal tendon insertion ventral to femoral tendon of gracilis muscle; state 1,

distal tendon insertion dorsal to femoral tendon

of gracilis muscle; state 2, distal tendon piercing

the gracilis muscle or femoral tendon of the

gracilis muscle.

State 0 is the common state among the frogs

considered to be related to, or primitive to, the

myobatrachids (Dunlap, 1960). State 0 is assumed

the primitive state (Criterion II of Heyer, 1975a).

The directions of changes of character states are:

Taiidactylus species demonstrate both states 1

and 2. As these states are coded as independently

derived, the genus is coded as having state 0 for

purposes of analysis.

CHARACTER 25, adductor magnus muscle: state 0,

accessory head absent; state 1, accessory head

present.

State 0 is found in the Ascaphidae, Discoglossidae, Liopelmatidae, Microhylidae, Pelobatidae,

Pipidae, and Rhinophrynidae; state 1 is the usual

state for the remaining families (Dunlap, 1960).

State 0 is assumed to be the primitive state

(Criterion II of Heyer, 1975a, for expanded outgroup).

CHARACTER 26, extensor cruris brevis muscle:

state 0, muscle as long as tibio-fibula; state 1,

muscle from y3 to 2/3 length of tibio-fibula.

The common condition among frogs is state 0

(Dunlap, 1960). State 0 is assumed the primitive

state (Criterion II of Heyer, 1975a).

CHARACTER 27, abductor brevis plantaris hallucis

muscle: state 0, muscle absent; state 1, muscle

present, no accessory slip; state 2, muscle and

accessory slip present.

The muscle is absent in Ascaphus, Leiopelma,

Pipa, and Rhinophrynus (Dunlap, 1960), suggesting that this is the primitive state (Criterion II of

Heyer, 1975a). Liem (in press) stated that an

accessory slip was essentially unique to certain

myobatrachids. The direction of change of character states is:

CHARACTER 28, Omohyoideus: state 0, muscle

present in all species; state 1, muscle present in

some species, absent in others; state 2, muscle

absent in all species.

Heyer (1975a) indicated the direction of change

of states is:

0 • 1 • 2

CHARACTER 29, geniohyoideus medialis muscle:

state 0, muscle contiguous medially; state 1, muscle

separated medially.

Heyer (1975a) indicated state 0 is the primitive

state.

CHARACTER 30, sternohyoideus ventralis muscle

insertion: state 0, muscle insertion near lateral

edge of hyoid body; state 1, muscle insertion

extending to midline of hyoid body.

Heyer (1975a) indicated state 0 is the primitive

state.

CHARACTER 31, petrohyoideus anterior muscle:

state 0, muscle insertion on lateral edge of hyoid

plate; state 1, muscle insertion on ventral body

of hyoid.

Heyer (1975a) indicated state 0 is the primitive

state.

CHARACTER 32, petrohyoideus posterior muscles:

state 0, three slips present; state 1, two slips present.

Most frogs have state 0 (Trewavas, 1933; Liem,

1970); the state is assumed primitive (Criterion II

of Heyer, 1975a).

CHARACTER 33, dilatator laryngis muscles: state

0, one pair of slips present; state 1, some species

with one pair, others with two pairs of slips present; state 2, two pairs of slips present.

The usual state for frogs is state 0 (Trewavas,

1933); state 0 is assumed the primitive state (criterion II of Heyer, 1975a). The direction of change

of states is:

0 • l • 2

CHARACTER 34, pupil shape: state 0, pupil vertical; state 1, pupil horizontal.

Heyer (1975a) indicated state 0 is the primitive

state.

CHARACTER 35, abdominal skin texture: state 0,

skin of abdomen smooth; state 1, skin of abdomen

coarsely granular in some or all species.

The majority of frogs have state 0; it is assumed

primitive (criterion II of Heyer, 1975a).

CHARACTER 36, tongue: state 0, tongue free

posteriorly; state 1, tongue adherent to mouth

floor.

About 10 genera of frogs have state 1; the state

is considered derived (criterion II of Heyer, 1975a).
NUMBER 233

CHARACTER 37, toe webbing; state 0, toes

webbed; state 1, toes fringed in some or all species;

state 2, toes free in some or all species.

Heyer (1975a) indicated the direction of change

of character states to be:

CHARACTER 38, inner metatarsal tubercle: state

0, inner metatarsal tubercle not enlarged and

shovel-shaped; state 1, inner metatarsal tubercle

enlarged, shovel-shaped, compressed, and keeled.

State 0 is considered primitive (Heyer, 1975a).

CHARACTER 39, outer metatarsal tubercle: state

0, absent; state 1, absent in some species, present

in others; state 2, present, unmodified; state 3,

present, strongly compressed.

Heyer (1975a) indicated state 0 to be the primitive state. The direction of change of character

states is:

CHARACTER 40, intermandibularis muscle: state

0, no intermandibularis-submentalis connection;

state 1, intermandibularis-submentalis connection

present.

The generalized distribution of state 0 (Tyler,

1971) suggests this state to be primitive (criterion

II of Heyer, 1975a).

CHARACTER 41, egg mass: state 0, not foamy;

state 1, foamy.

Heyer (1975a) indicated state 1 to be derived.

CHARACTER 42, site of egg development: state

0, aquatic; state 1, non-aquatic, including terrestrial and parental brooding.

Heyer (1975a) indicated that state 0 is primitive.

The distribution of states among the taxa is

presented in Table 1. The recoding of states as

used in the next section is presented in Table 2.

Analysis of Relationships

The combinatorial method of Sharrock and

Felsenstein (1975) was used to organize the data.

The combinatorial method locates all monothetic

clusters in the data set and prints them out. Information from the character state trees was included;

the clusters were formed with derived states

only. The construction of the first phylogeny

(Figure 1) maximized the number of shared

derived states at each clustering point. Certain

conclusions can be drawn from the phylogeny of

Figure 1. There are three major groupings of

myobatrachid frogs: Rheobatrachus forms a group

by itself and apparently has little if any relationship to the remaining genera; the remaining genera

form two major clusters, corresponding with taxonomic proposals made by previous workers. The

cluster including Myobatrachus corresponds with

the subfamily Myobatrachinae; the cluster including Limnodynastes corresponds with the subfamily

Limnodynastinae (Liem, in press) (=Cycloraninae

of Lynch, 1971, and Parker, 1940, except for the

exclusion of Cyclorana). The basal clusters defining the two groups each contain unique state

appearances within the tree. As discussed later,

three of the muscle characters may be miscoded;

nevertheless, each cluster still has unique state

appearances within the tree. These two groups are

convincing sister groups in terms of Hennig's

(1966) terminology. There are no other meaningful arrangements of these two groups.

There are several rearrangements possible within

each group, however, that need detailed comment.

The cluster of Assa, Metacrinia, Pseudophryne,

and Pseudophryne occidentalis is the strongest

cluster in the data set, indicating that the closest

relationship of any of these taxa is with the other

three. Characters not used in the analysis provide

evidence to suggest a rearrangement within the

cluster. First, however, the cluster of Pseudophryne

occidentalis and Pseudophryne is substantiated: all

members lack tympani and columellae, have a

unique locomotion of crawling (except P. douglasi)

(Liem, in press), and have a uniform mating call

(Liem, in press). Pseudophryne does form a sister

group with P. occidentalis; all Pseudophryne (except P. occidentalis) have the derived state of the

depressor mandibulae muscle and have femoral

glands (a derived condition, Heyer, 1975a), while

P. occidentalis has narrow sacral diapophyses, a

compressed outer metatarsal tubercle, and two

slips of the dilatator laryngis muscles (Liem, in

press). These morphological differences indicate

different adaptive complexes in these two groups,

differences best reflected at the generic level. Liem

(in press) pointed out the distinctiveness of

P. occidentalis but preferred not to recognize it

at the generic level. Blake (1973) did not include

P. occidentalis in his analysis.

Blake (1973) proposed placing Metacrinia in the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 1.—Distribution of character states among genera as used for computer coding ("9"

indicates no information)

Genus

Adelotus

Assa

C. georgiana

C. haswelli

Geocrinia

Ranidella

Glauertia

Heleioporus

Kyarranus

Lechriodus

Limnodynastes

Metacrinia

Mixophyes

Myobatrachus

Neobatrachus

Notaden

Philoria

Platyplectron

Pseudophryne

P. occidentalis

Rheobatrachus

Taudactylus

Uperoleia

Genus

Adelotus

Assa

C. georgiana

C. haswelli

Geocrina

Ranidella

Glauertia

Heleioporus

Kyarranus

Lechriodus

Limnodynastes

Metacrinia

Mixophyes

Myobatrachus

Neobatrachus

Notaden

Philoria

Platyplectron

Pseudophryne

P. occidentalis

Rheobatrachus

Taudactylus

Uperoleia

1h-i

0

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26

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2

0

2

2

2

2

1

2

2

0

0

0

8

0

1

1

1

1

1h-i

0

0

u

0

1

1

0

0

0

0

1

1

0

1

1

29

0

1h-i

1

1

1

1

0

0

0

0

1

0

0

0

0

0

0

1h-1

0

1

1

9

1

0

0

0

0

0

0

1

1

1

1

0

n

u

0

1

1i-H 1

0

0

0

0

0

30

1

1

1

1h-i

1

1

0

1

0

0

1

0h-l

0

1

1

0

1

1

0

1h-1

Numbers

10

0

1h-i

1

1

1

0

0

0

u

0

1

u

0

0

0

0

0h-i h-l

0h-l h-1

31

0

1

1

1

1h1 1

0

0

0

0

1

0

1

0

0

0

0

1

1

0

1

1

11

0

0

0

0

0

0

0

0

0

1

0

0

0

0

0

0

0

0

0

0

0

0

32

0

1

0

0

0

0

0

0

0

0

0h-l

0

1

0

0

0

0

0

0

1

0

0

and

12

1

0

1h-i i—i 1

1

1

1

1

1

1

1

1

1

1

0

1

0

0

0

33

0

0

0

2

1

0Is)

0

0

0

0

0

0

2

0

0

0

0

0

0

0

1

0

States

13h-i

1

1h-i

1

1

1h-i

1

1rH r

1

1

M

1

1

1h-i

0

1

1

34

1

1

1i-t 1h-i

1

0

1

1H-I M

0

1

0

0

1

0

1

1

0h-l

0

14

1

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

35

0

0

1

0

0

1

1

0

0

0

0h-l

0

0

0

0

0

0

0

0

0

0h-l

15

1

1

0

0

0

0

1

1

1h-i h

1

1

1

1

1r-i h-i

1

1

0

1

36

1

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

1

0

0

16

0

0

0

0

0

0

1

1

0

1

0

0

1

1

1

1

0

0

0

0

0

37

0

2

2

1

2

2

0

1

2

2

2

2

0ts)

0

0

2

0

2ts)

0

1

1

17

0h-i

1h-i

1h-i

1

0

0

n

U

0

D h-i

u

1

0

0

0

0

1

1

0

1

1

38

0

0

0

0

0

0

0h-i

0

0

0

0

0

0

1

1

0

1

0

0

0

0

0

18

1

0

0

0

0

0

0

1

1

1

X

1

0

1

1

0

1

1h-l h1 0

0

0

0

0

39

2

2

2

2

0

2

3

0

0

0

1

2

0

0

0

1

0

0

2

3

2

2

3

19

1

1

0

0

0

0

1

0

1

T

J0

D h-i h-i C

0

1h-l

0

1h-l

1

1

1

40

1

0

0

0

0

0

0

1

1

1

1

0

1

0

1

1

1

1

0

0

0

0

0

20

0

9

0

0

0

1

M

0

0

nU

0

0

r\

U

0

0

0

2

0

2

0

0

0

2

41

1

0

0

0

0

0

0

1

1h-1 h-l

0

0

0

0

0

1

1

0

0

0

0

0

21

1

1

1

1

1

1

2

1

1

r>U

1

1

r\

U

1

1

1

1h-i

1

1

1

1

1

42

0

1

0

0

1

0

0

1

1

0

0

1

1

1

0

0

1

0h-l

1

1

0

0
NUMBER 233

TABLE 2.—Recoding key of character states from states as analyzed to states used in the

computer program

New state

Character

Old state

New state

Character

Old state

New state

Character

Old state

New state

Character

Old state

number

nunber

nurrber

nunber

nunber

nunber

nunber

nunber

nunber

nunber

nunber

nunber

1

1

1

15

10

j.

29

22

1

43

33

1

2

1

2

16

11

T

X

30

23

1

44

33

2

3

2

1

17

12

X

31

24

1

45

34

1

4

3

1

18

13

32

24

2

46

35

1

5

3

2

19

14

33

25

1

47

36

1

6

4

1

20

15

34

26

1

48

37

1

7

5

1

21

16

35

27

1

49

37

2

8

6

1

22

17

36

27

2

50

38

1

9

7

1

23

18

37

28

1

51

39

1

10

7

2

24

19

38

28

2

52

39

2

11

7

3

25

20

39

29

1

53

39

3

12

7

4

26

20

t

l

40

30

1

54

40

1

13

8

1

27

21

i

i

41

31

1

55

41

1

14

9

1

28

21

2

42

32

1

56

42

1

PSEUDOPHRYNE

25,26 .

P. OCCIDENTAL IS

,12,17,53

METACRINIA

y

\ 17,42,46

20,24,49,56

\ .

37,38

RANIDELLA

11,17,25,

46,45

15,27,29,39,

48,51,52^

\

GLAUERTIA

4,21,25,28,

29,39,46,51,

52,53 /

2,11,12,17,20,

24,37,38,43,44

MYOBATRACHUS

3,27,42,48,

«9,56

TAUDACTYLUS

30,35,43,45

6,15,24,27'

39,48,51,52.

UPEROLEIA

1,4,5,11,20,25,

26,29,31,46,53

/

9,10

31,«5

C. GECRGIANA

46,51,52

\ /

15,17,27,29,31,

39,45,48,49

(NO)

GEOCRINIA

9,37,«3,56

\

7,8,13,22,40,41,

PHILORIA KYARRANUS

3,21,25,26 8,29

4,5,24,40,

48,49,56

1«,27,36,37,'

38,45,55

PLATYPLECTRON

14,27,29,36,

37,50,55

4,5,21,32,34

HEIEIOPORUS

«»5,14,21,27,

34,48,50,55,56

NEOBATRACHUS NOTAOEN

30,33 2,7,8,24,

V 29,<°,51

4,14,21,27,

34,37,38,50

17,20,2a, 51

RHEOBATRACHUS

1,8,9,10,20,24,

27,29,42,47,51,

52,56

FIGURE 1.—Predicted phylogenetic relationships among the myobatrachid genera, ("(NO)" —

cluster is nonmonothetic; underlined states — unique appearances within phytogeny; see text.)
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

synonymy of Pseudophryne. This action is not

supported by the present study. Metacrinia and

Pseudophryne (including occidentalis) form robust

sister groups: Pseudophryne lacks a columella and

crawls (except P. douglasi); Metacrinia has the

derived state of the posterior petrohyoideus muscle, a granular belly, and terrestrial larvae. These

differences are best expressed at the generic level.

The relationships of Metacrinia appear to lie closest with Assa, in fact. Within the characters used

for analysis, all of the derived states of Assa are

shared with Metacrinia. Thus, within the cluster

of four, Assa and Metacrinia together share the

derived state of the posterior petrohyoideus muscle. In addition, both taxa have the derived condition of reduction of a phalanx in the first finger,

a state found additionally only in Uperoleia.

Metacrinia and Assa do form sister groups when

more life history information is added: Metacrinia

has narrow sacral diapophyses and a granular

belly; Assa has male brooding pouches. This distinctive life history pattern is best expressed at the

generic level as currently recognized.

In the phylogeny of Figure 1, Crinia georgiana

does not form a mbnothetic cluster with Geocrinia.

A better arrangement, in which all clusters are

monothetic, involves pairing C. georgiana with

Ranidella, and Geocrinia with C. haswelli. Interestingly, a combination of these four taxa does not

form a monothetic cluster. The combination of all

four with Assa, Metacrinia, and Pseudophryne

(including occidentalis) is a monothetic cluster, as

is a cluster of either C. georgiana-Ranidella or

Geocrinia~C. hasivelli with Assa, Metacrinia, and

Pseudophryne. There is no clear advantage to be

gained from choosing between these last alternatives in terms of numbers of shared states within

clusters, or number of convergent states. Thus,

while all of the above taxa seem to be most closely

related among themselves, the combination of

C georgiana, C. haswelli, Geocrinia, and Ranidella

is a poor cluster since it is not monothetic.

Placement of C. haswelli with Geocrinia agrees

with Liem (in press), but disagrees with Blake

(1973), who placed C. haswelli together with

C. georgiana. Crinia georgiana and C. hasivelli do

not form a monothetic cluster in the present data

set, indicating that they are not most closely

related. Geocrinia and C. haswelli appear to be a

robust sister group (Figure 2) because C. haswelli

has 5 unique states and Geocrinia 2. However,

many of these states are the uniform expression

of a variable state. For example, the omohyoideus

is absent in haswelli, but it is also absent in some

Geocrinia. Thus these kinds of states can not be

used to define clear cut sister groups. Each taxon

has one unique state: C. haswelli has an outer

metatarsal tubercle; Geocrinia has terrestrial egg

deposition. A further complication in attempting

to deduce relationships is occasioned by the fact

that C. haswelli actually shares two more derived

states with the cluster of Assa, Metacrinia, and

Pseudophryne (including P. occidentalis) than with

Geocrinia. The taxonomic decision of whether to

recognize C. haswelli as a distinct genus has a

parallel, in part, with the Ranidella complex; the

decision involving C. haswelli will be deferred

until the situation with Ranidella is described.

On the basis of the characters analyzed for this

study, all of the derived states of Crinia georgiana

are found within Ranidella, suggesting that these

two taxa are most closely related to each other.

Once the individual characters are examined in

detail, it becomes apparent that R. riparia is distinctive within Ranidella, and for purposes of

present discussion should be separated. The best

placement of Ranidella riparia is with C. georgiana

and the remaining Ranidella Liem (in press)

examined (Figure 2). States 9, 10, and 11 separate

C. georgiana from Ranidella (restricted sense). All

these states involve reduction of the vomer and

vomerine teeth. Vomerine teeth are absent in

Ranidella (restricted sense), the vomer is present

or absent. Since there is a trend within Ranidella

(restricted sense), it is not unreasonable to carry

the trend back one step further to include

C. georgiana within the same genus. The larval

evidence (Watson and Martin, 1973) does not support inclusion of C. georgiana in with Ranidella

(restricted), because C. georgiana has a stream

adapted tadpole (as do R. riparia and R. tasmaniensis). Of the species examined in this study,

R. riparia is distinctive from a cluster of C. georgiana and Ranidella (restricted sense) in mandibular musculature, terrestrial egg placement, and

possibly loss of the columella (Blake, 1973, so

indicates; Liem, in press, does not). The mating

call of C. georgiana is a unique quacking call,

distinctive within the Ranidella complex (Main,

1968).
NUMBER 233

PSEUDOPHRYNE P. OCCIDENTALIS METACRINIA

25,26. 12,17,53

2,11

RIPARIA RANIDELLA

25,26,56 11

5,10

C. GEORGIANA

25,31,

2,17,46

- I -

ASSA C. HASWELLI

42 10,38,44,

KYARRANUS

8,25

GEOGRINIA

45,56

/

17,43

TAUOACTYLUS UPEROLEIA

30,35,43,45 1,4,5,11,20,25

Y 26,25,31,46,53

6,9,10,24,51,52

PHILORIA

3,21,25,

26

4,5,24,40,

48,45,56

38,*5

AOELOTUS LIMNOOYNASTES

1,6,15,24, 21,48,45

34,40,47,

29,32,51

/

/

PLATYPLECTRON

4,5,21,25,32,

34,50

27,35,36,37

\

GLAUERTIA MYOBATRACHUS - \ ./-

4,21,25,28, 3,27,42,48,

29,39,46, 45,56 15,27,35,48

HELEIOPORUS LECHRIODUS

27,50,56 6,16,19,24,29,

32,35,45,49

,21,34,48

NEOBATRACHUS NOTADEN

30,33 2,7,8,24,

C 29,40,51

4,21,27,34,

37,38,50

MIXOPHYES

3,4,5,6,16,19,21,

30,32,33.34,35,56

RHEOBATRACHUS

1,8,9,10,20,24,27,

29,42,47,51,52,56

FIGURE 2.—Alternate phylogenetic relationships among the myobatrachid genera. (Underlined

states = unique appearances within phylogeny; dashed lines = lineages are not sister groups;

see text.)

In attempting to determine the relationships

among Crinia georgiana, C. haswelli, Ranidella

riparia, Geocrinia, and Ranidella, certain characters are keys to understanding the relationships:

vomer and vomerine teeth, omohyoideus muscle,

outer metatarsal tubercle, belly texture, and egg

placement. There is no way to group the myobatrachine taxa so that two of the derived states of

these characters define the same assemblage.

Rather, a grouping which results in a cluster having all the taxa with the same derived state of one

character leads to convergence of states in the

other characters. The most reasonable assumption

is that the common ancestor of the group had the

genetic potential to produce the states in question,

but, in part, the results were a mosaic pattern.

Based on this assumption, the most reasonable

taxonomic criteria to describe the situation is the

recognition of the following as distinct genera,

indicating separate evolutionary lineages: (1)

Geocrinia, (2) Crinia haswelli, (3) Crinia georgiana,

(4) Ranidella, and (5) R. riparia and R. tasmaniensis (latter was not included in Liem's, in press,

study sample). This arrangement is consistent with

the available larval evidence (Watson and Martin,

1973) and has the advantage that each unit is

homogeneous with respect to life history mode.

Separation of tasmaniensis from Crinia, sensu lato,

has been previously suggested (Littlejohn, 1970).

Of the remaining myobatrachine taxa, Taudactylus and Uperoleia are most closely related, and

when joined with the taxa mentioned above, form

a monothetic cluster with a unique state appearance (Figure 2). Glauertia and Myobatrachus are

closely related and can also form a monothetic

cluster at the same level as Taudactylus and

Uperoleia of Figure 2, but the cluster does not

have a unique state appearance.

The proposed relationships among the limnodynastines of Figure 1 contain one non-monothetic
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cluster. If clusters are chosen to maximize the

number of appearances of unique states within

the total phylogeny, one major difference in placement occurs: the location of Mixophyes. With

Mixophyes removed basally, the remaining limnodynastines have fused cervical and second vertebrae. Mixophyes is unique in that the nervus

mandibularis lies between the levator mandibulae

externus and subexternus muscles, it is the only

myobatrachid genus with axillary amplexus (DSL),

the genus is unique in egg placement (Liem, in

press) and tadpole morphology (Watson and Martin, 1973), further supporting the basal dichotomy

as represented in the phylogeny of Figure 2.

Liem (in press) thought Heleioporus and Neobatrachus to be closely related. This hypothesis is

not substantiated by this study; in fact, the two

taxa do not share a monothetic cluster. Assuming

that Neobatrachus has closest relationships with

Notaden, then the remaining limnodynastines are

characterized by eggs laid in foam nests.

Liem (in press) indicated a close relationship

between Adelotus and Lechriodus. Apparently this

was occasioned by shared primitive states, since

analysis of the derived states indicates the closest

relationships of Adelotus and Lechriodus lie elsewhere rather than with each other (Figure 2).

Liem (in press), following the lead of Littlejohn

(1963), proposed placing Kyarranus in the synonymy of Philoria. These two taxa share the most

derived states among the limnodynastines. Further,

the two taxa uniquely share the derived life history

pattern of nonaquatic, nonfeeding larvae among

the limnodynastines. Within this study, Kyarranus

and Philoria form sister groups, Kyarranus having

derived states of the squamosal and the supracoracoideus muscle, Philoria having derived states of

the nasal and frontoparietal relationship, prehallux, and depressor mandibulae muscle. Philoria

has undergone a separate evolutionary history

from Kyarranus although both had a common

ancestor. It seems rather a point of personal preference whether the emphasis is on the common

ancestry, resulting in recognition of a single genus,

or on the evolutionary divergence, suggesting two

genera. In fact, we disagree on this one point: the

first author prefers to recognize two genera, the

second author prefers to recognize one. For purposes of this study, Kyarranus and Philoria are

considered as separate generic units in order to

present more information.

SISTER GROUPS WITHIN THE PHYLOGENY

Within the phylogeny of Figure 2, those dichotomies not representing sister groups (Hennig,

1966) are indicated by dashed lines. In two cases,

additional information is needed to recognize

sister groups. In the case of Mixophyes and the

remaining limnodynastines, the life history information has been presented previously. In the case

of Rheobatrachus compared with the rest of the

genera, Rheobatrachus has a unique sacro-coccygeal

articulation (Liem, 1973) and a unique developmental pattern of gastric larval incubation (Corben, Ingram, Tyler, 1974). As argued elsewhere

(Heyer, 1975a), the weakest areas in the phylogenies

occur where sister groups are not present. It seems

reasonable to assume that higher taxa recognized

in an analysis of this sort should be sister groups.

COMPARISON WITH PREVIOUS SCHEMES

Lynch (1971) summarized the classificatory

schemes of the myobatrachids, with the exception

of Rheobatrachus, which was unknown to science

at that time. Parker (1940) was the first to recognize the distinctiveness of the myobatrachines and

limnodynastines. Lynch's (1971) study, as well as

the present study, supports Parker's conclusions.

Lynch (1971) further recognized two tribes in the

limnodynastines. The present study does not support that conclusion, since Lynch included

Heleioporus, Mixophyes, Neobatrachus, and Notaden in a single tribe. The results of this study

point to the distinctiveness of Mixophyes within

the limnodynastines.

Liem (in press) included Rheobatrachus in the

subfamily Myobatrachinae. In contrast to the present study, Liem (in press) also drew some different

systematic conclusions at the generic level, as did

Blake (1973) in his study of the Myobatrachinae.

Differences between this study and Liem's and

Blake's studies are due to the different analytic

procedures used. Both Liem (in press) and Blake

(1973) used a phenetic approach in which primitive and derived states were used to produce phylogenetic hypotheses. In this study, only derived state

information was used to produce phylogenetic
NUMBER 233 11

hypotheses. We think that clusters based on shared

primitive states are weak, since there is no information with respect to change from an ancestral

condition. All taxa can be systematically thought

of as collections of primitive and derived character

states. The primitive states indicate no change in

that character from the ancestral condition. The

derived states do indicate a change from an ancestral condition. If groups of taxa share the same

derived states, they may form a common evolutionary lineage. From a philosophical point of

view, we think the approach used in this study is

superior for deducing relationships than is a phenetic analysis, which indiscriminately uses primitive

and derived character state information. The

approximation of reality in the kind of analysis

presented herein depends on the correctness of

determination of primitive and derived states.

Systematic Conclusions

The relationships, as presented in the phylogenies (Figures 1, 2), are best represented by recognizing three subfamilies: one for Rheobatrachus,

one for the myobatrachines, and a third for the

limnodynastines. The limits of this study preclude

an evaluation of the familial status of these three

units. It may be that all three units are actually

familial units. Some authors (Liem, in press; Watson and Martin, 1973) suggest that the myobatrachines and limnodynastines are familial units; this

study demonstrates that Rheobatrachus is very

distinctive from the myobatrachines and limnodynastines. Recognizing each as a subfamily has

the advantage of pointing out the major differences within the total study group, while not adding taxonomic confusion to any future situation

(a suprageneric taxon is available for a family

name).

The only possible formal classification below the

subfamily level that could be proposed confidently

would be to recognize a tribe for Mixophyes. There

seems, however, to be no advantage to emphasizing

the distinctiveness of Mixophyes in a formal action.

The new taxonomic categories or combinations

arising from this study are presented formally.

Diagnostic definitions only are provided since the

full character complements of the taxa are presented elsewhere within the paper (Table 1).

The genera of each of the subfamilies as recognized in this study are:

NEW SUBFAMILY

Rheobatrachus

MYOBATRACHINAE

Assa

Crinia

Geocrinia

Glauertia

Metacrinia

Myobatrachus

Pseudophryne

Taudactylus

Uperoleia

New genus for Pseudophryne occidentalis

New genus for Crinia haswelli

New genus for Ranidella

riparia and tasmaniensis

LlMNODYNASTINAE

Adelotus

Heleioporus

Kyarranus

Lechriodus

Limnodynastes

Mixophyes

Neobatrachus

Notaden

Philoria

Platyplectron

RHEOBATRACHINAE, new subfamily

TYPE-GENUS—Rheobatrachus Liem, 1973.

DIAGNOSIS—Maxillary teeth fanglike; fontanelle

not roofed over by frontoparietals; columella

present; cervical cotyles narrowly separated; sacral

diapophyses broad; sacro-coccygeal articulation

monocondylar; coccygeal prezygapophyses and

transverse processes present; cricoid a complete

ring; alary process of hyoid broad and winglike;

pupil vertical; tongue adherent to mouth floor;

outer metatarsal tubercle present; female gastric

brooding of young.

CONTENT.—Monotypic, Rheobatrachus silus.

MYOBATRACHINAE Schlegel, 1850

Crinia Tschudi, 1838

TYPE-SPECIES.—Crinia georgiana Tschudi, 1838.

DIAGNOSIS.—A myobatrachine genus; cervical

cotyles widely separated; vomer and vomerine

teeth present; narrow sacral diapophyses; depressor

mandibulae muscle with slip from dorsal fascia;

omohyoideus muscle present; tympanum present;

belly smooth; toes without fringe or web; outer

metatarsal tubercle present; eggs placed in water;

stream larvae.

CONTENT.—Monotypic, Crinia georgiana.

Geocrinia Blake, 1973

TYPE-SPECIES.—Pterophrynus laevis Gunther,

1864.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DIAGNOSIS.—A myobatrachine genus; cervical

cotyles widely separated; vomer present, vomerine

teeth present or absent; narrow sacral diapophyses;

depressor mandibulae muscle with slip from dorsal

fascia; omohyoideus muscle present or absent; tympanum present; belly smooth; toes lacking fringe

or web; outer metatarsal tubercle absent; eggs laid

in concealed sites on land; pond larvae or terrestrial, non-feeding larvae.

CONTENT.—Geocrinia laevis, leai, lutea, rosea,

victoriana.

Pseudophryne Fitzinger, 1843

TYPE-SPECIES.—Phryniscus australis (in part)

Dume'ril and Bibron, 1841 (—Pseudophryne semimar morata Lucas, 1892).

DIAGNOSIS.—A myobatrachine genus; cervical

cotyles widely separated; vomer present or absent,

vomerine teeth absent; broad sacral diapophyses;

depressor mandibulae muscle lacking a slip from

dorsal fascia; omohyoideus muscle absent; columella absent; belly smooth; toes free, no fringe

or web; outer metatarsal tubercle present; eggs

placed on land (except P. douglasi, which places

eggs in water); pond larvae.

CONTENT.—Pseudophryne australis, bibroni, coriacea, corroboree, dendyi, douglasi, guentheri, major, semimarmorata.

Ranidella Girard, 1853

TYPE-SPECIES.—Ranidella signifera Girard, 1853.

DIAGNOSIS.—A myobatrachine genus; cervical

cotyles widely separated; vomer present or absent,

vomerine teeth absent; narrow sacral diapophyses;

depressor mandibulae muscle with slip from dorsal

fascia; omohyoideus muscle present; tympanum

present; belly granular; toes fringed or free; outer

metatarsal tubercle present; eggs placed in water;

pond larvae.

CONTENT.—Ranidella glauerti, insignifera, parinsignifera, pseudinsignifera, remota, signifera, sloanei, subinsignifera, tinnula.

Paracrinia, new genus

TYPE-SPECIES.—Crinia haswelli Fletcher, 1894.

DIAGNOSIS.—A myobatrachine genus; cervical

cotyles widely separated; vomer present, vomerine

teeth absent; narrow sacral diapophyses; depressor

mandibulae muscle with slip from dorsal fascia;

omohyoideus muscle absent; tympanum present;

belly smooth; toes fringed; outer metatarsal tubercle present; eggs placed in water; pond larvae.

CONTENT.—Monotypic, Paracrinia haswelli.

ETYMOLOGY.—From the Greek para ( = beside),

indicating the fact that the relationships of this

genus fall within the Crinia complex. The genus is

feminine in gender.

Australocrinia, new genus

TYPE-SPECIES.—Pterophrynus tasmaniensis Giinther, 1864.

DIAGNOSIS.—A myobatrachine genus; cervical

cotyles widely separated: vomer present, vomerine

teeth present or absent; narrow sacral diapophyses;

depressor mandibulae muscle lacking slip from

dorsal fascia; omohyoideus muscle present; tympanum present; belly granular; toes fringed or

free; outer metatarsal tubercle present; eggs placed

on land or water; stream or pond larvae.

CONTENT.—Australocrinia riparia, tasmaniensis.

ETYMOLOGY.—From the Latin australis ( = southern), indicating the southern geographic distribution of this member of the Crinia complex. The

genus is feminine in gender.

Kankanophryne, new genus

TYPE-SPECIES.—Pseudophryne occidentalis Parker,

1940.

DIAGNOSIS.—A myobatrachine genus; cervical

cotyles widely separated; vomer and vomerine teeth

absent; narrow sacral diapophyses; depressor mandibulae muscle with slip from dorsal fascia;

omohyoideus muscle absent; columella absent;

belly smooth; toes free of fringe or web; outer

metatarsal tubercle present; eggs placed on land;

pond larvae.

CONTENT.—Monotypic, Kankanophryne occidentalis.

ETYMOLOGY.—From the Greek kankanos (= dry),

in reference to the arid distribution of this genus.

The genus is feminine in gender.

Phyletic Character Analysis

The analysis of characters into primitive and
NUMBER 23S 13

derived states seems overall to be intuitively sound,

since the recognition of the myobatrachines and

limnodynastines as distinct units is in agreement

with every other systematist that has worked with

the myobatrachids in any detail.

As discussed in detail elsewhere (Heyer, 1975a),

an analysis of the phylogenetic usefulness of each

character state can be made after the fact. This

is done by using the information from the better

phylogenetic tree to determine the number of

appearances of each state (as printed) on the tree

divided by the total number of taxa that actually

have that state. The numeric values for each state

are then ranked and grouped into categories. A

value of 1 indicates no phylogenetic information;

the smaller the number, the better the state links

the taxa as found in the preferred phylogeny.

The phylogeny of Figure 2 was used for analysis,

except that riparia and Ranidella were considered

as a single unit to be consistent with the original

data. For multistate characters, averages are used.

Best characters have values of 0.04 to 0.20, good

characters have values of 0.25 to 0.42, average characters have values of 0.50 to 0.60, poor characters

have values of 0.67 to 1.00.

The best characters with a unique appearance

in the phylogenetic tree are: intervertebral disc,

cervical vertebral fusion, cervical cotyles, coccyx,

cricoid, alary process of the hyoid, anterior petrohyoideus muscle, intermandibularis muscle, egg

mass type. The best characters with two or more

appearances in the tree are: sphenethmoid and

geniohyoideus muscle.

Good characters are: squamosal, sacral diapophyses, carpals, adductor longus muscle, adductor magnus muscle, omohyoideus, sternohyoideus,

pupil shape, and toe webbing.

Average characters are vomer and vomerine

teeth, anterior process of the hyale, supra coracoideus muscle, semitendinosus muscle.

Poor characters are: maxillary teeth, nasal contact with frontoparietial, carotid canal, frontoparietal fontanelle, transverse process width, dorsal crest

of ilium, prehallux, depressor mandibulae muscle,

adductor mandibulae muscle, extensor cruralis

brevis muscle, petrohyoideus posterior muscles,

dilatator laryngis muscle, belly texture, tongue,

inner metatarsal tubercle, outer metatarsal tubercle, site of egg development.

There are seven multistate characters in which

the states of a given character differ with respect

to clustering behavior. State 1 of the vomervomerine teeth complex (character 7) has good

clustering behavior, states 2 and 3 are average,

state 4 is poor. State 1 of the adductor mandibulae

muscle (character 21) has good clustering behavior,

state 2 is poor. State 1 of the semitendinosus is

good (character 24), state 2 is poor. State 1 of the

abductor brevis plantaris hallucis muscle (character

27) is average, state 2 is best. State 1 of the dilatator laryngis muscle (character 33) is average, state

2 is poor. State 1 of toe webbing (character 37) is

good, state 2 is average. States 1 and 2 of the outer

metatarsal tubercle are average, state 3 is poor.

Examination of the characters indicates a discrepancy with respect to the hyoid complex. As

noted in the character analysis, the broad, winglike state of the alary process is thought to be

the primitive state, in contrast to previous coding

(Heyer, 1975a). The associated muscles, corresponding with broad winglike alary process, however,

were coded as derived states, as in the previous

study (Heyer, 1975a). The distribution of states

indicates that in order to be consistent, the primitive and derived states should be reversed for the

following characters: geniohyoideus medialis muscle (character 29), sternohyoideus muscle (character

30), and anterior petrohyoideus muscle (character

31).

The purpose of this entire analysis is to produce

a likely phylogenetic hypothesis for the myobatrachids. The analysis thus far concludes that there

are several changes that could be made in the

original data set that would improve the phylogenetic hypothesis. The changes are: (1) deletion

of Rheobatrachus, since its evolutionary history

appears to be distinct from the evolutionary history

of the other two subfamilies; (2) addition of Anstralocrinia; (3) reversal of direction of character

states for character numbers 29, 30, and 31; (4) deletion of characters in the poor category (as analyzed above). The purpose of making the changes

in the data set is not to draw new taxonomic conclusions. Taxonomic conclusions were drawn on

the basis of the entire original data set. Rather,

the purpose of making the changes is to determine

whether the changes allow new insights into the

relationships among the taxa.

The best arrangement of the taxa in terms of

maximization of character states at each cluster,
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

numbers of monothetic clusters, and unique state

appearances is presented in Figure 3. All clusters

are monothetic.

Within the myobatrachines, sister groups are

represented at the basal branchings involving

Taudactylus, Glauertia, and Myobatrachus. None

of the branchpoints from Geocrinia onward are

sister groups. This pattern contrasts with the pattern seen in Figure 2, where the distal branchings

all represented sister groups, but the basal branchings did not represent sister groups. As discussed

previously, it appears that evolution in the Criniacomplex involved a mosaic pattern of characters.

This hypothesis is consistent with a phylogeny in

which no sister groups are present. A mosaic pattern of evolution in the Crin ja-complex is best depicted in the phylogeny of Figure 3.

Among the limnodynastines, there are two alternate sequences at the basal branching. The

position of Notaden and Mixophyes can be interchanged. In the interchanged arrangement, character state 14, involving fusion of the cervical

vertebrae, is not a uniquely derived state, but the

state of the adductor magnus muscle (state 33) is.

With the data in the revised set, Notaden forms

a sister group with all other limnodynastines

whereas Mixophyes does not. As presented earlier,

data not originally coded will allow Mixophyes to

form a sister group with all other limnodynastines.

We prefer the arrangement depicted in Figure 3.

The cluster involving Adelotus, Limnodynastes, and

Platyplectron in Figure 3 represents the taxonomic

thinking of most previous workers, as Platyplectron

has usually been included in Limnodynastes. Adelotus does form a sister group with Limnodynastes

and Platyplectron. As in the phylogeny of Figure

2, the relationships proposed in Figure 3 involve

the unique development of the foam nest. Heleioporus does not have expanded fingers in the female as do all the other genera with foam nests.

The female finger flanges are used to beat up the

foam. The method of foam nest construction in

Heleioporus is unknown (Martin, 1972). The lack

of finger flanges in Heleioporus suggests that

Heleioporus represents a divergent evolutionary

lineage within the foam nest builders. This divergent pattern is best expressed in Figure 3.

The revised data set does allow construction of

PSEUOOPHRYNE OCCIDENTAL IS

\ 12,17,53

ASSA METACRINIA HASUELLI RANIOELLA RIPARIA UPEROLEIA

\ \ 17 17,37,38, 11,17,45,49 17,45,49 11,20,24,

-\—-\—I - 45 \ / ^ ^5 3

20,24,37,38,45,49^^=_^_^ \ /

GEORGIAN]! 9,10

17,45,49 /

GEOCRINIA 51,52

9,17,37, /

MYOBATRACHUS 15,29

9,10,11,12,17

20,24,37,39.

-3,-.45,4,N /

27,48

\ /

3 1 TAUOACTYLUS

\ 9.10,15.24,27.30

\ 35,45,48,51,52

7,8,13,22

GLAUERTIA

9,10,11,12,17,20,24,28,

29,37,38,45,51,52,53

KYARRANUS

8,29 /

\ /

24,27,36,37,

38,45,48,49

LIMNOOYNASTES PLATYPtECTRON

38,45,48,49,51 /

40" ADELOTUS

PHILORIA \ 24,38,45.51,52

\ /

27,36,37 LECHRIOOUS

w 24,n40",45,48,49

29,32

30,33

\

NOTAOEN

7,8,24,27,29,37,38,51

i l

MIXOPHYES

\ 30,32,33,35,"40"

17,20,20, "39","4i",51

FIGURE 3.—Preferred phylogenetic relationships among the myobatrachid genera. (Underlined

states - unique appearances within phylogeny; dashed lines = lineages are not sister groups;

quotation marks - direction of change of state has been reversed; see text.)
NUMBER 235 15

a phylogenetic tree that differs from the trees produced by the original data set. We think the

phylogeny based on the revised data set is a more

plausible phylogenetic hypothesis, representing the

best fit of the data with the probable phylogeny of

the group.

Evolutionary Zoogeography

GENERIC DISTRIBUTION PATTERNS

The known geographic distributions of the

genera, summarized in map form, are essential data

for the determination of distribution patterns.

This information for all genera is not available in

a single source. Since the discussion that follows is

based upon the basic distribution data, those data

are presented here (Figures 5-28). In grouping the

individual distribution maps into similar clusters

and attempting to associate those patterns with

features of the Australian climate, landscape, and/

or vegetation, certain facts become apparent. No

genus of myobatrachid frog is primarily adapted

to the desert habitat (compare Figure 20, Notaden,

the genus with the most extensive desert distribution, with Figure 4). In associating the distribution

patterns with other features, the distributions can

be understood best in terms of the vegetation types

found in the region. The geographic distributions

of the genera (Figures 5-28) show four major patterns associated with broad vegetational categories:

1. Distributions primarily restricted to the antarctic vegetation unit (temperate rainforest and

subtropical rainforest), > 1520 mm rainfall isohyet.

Assa, Kyarranus, Lechriodus, Rheobatrachus, and

Taudactylus share this pattern.

2. Distributions limited to wet forest vegetation

types (wet sclerophyll forest), ± 1000 mm isohyet.

The genera are either found only in the eastern

wet forest (Adelotus, Mixophyes, Paracrinia, Philoria), the southwestern wet forests (Crinia, Metacrinia), or both eastern and southwestern wet

forests (Geocrinia).

3. Distributions in both wet forest and subhumid woodland, about 500 mm to 1000 mm isohyts. The distributions are either primarily eastern

(Australocrinia, Limnodynastes), western (Myobatrachus), or eastern and western (Heleioporus,

Platyplectron, Pseudophryne, Ranidella, Uperoleia).

4. Open woodland and shrubland distributions,

neither wet forest nor primarily desert, < 500 mm

isohyet. Glauertia, Kankanophryne, Neobatrachus,

and Notaden share this pattern. Of these genera,

Notaden has the most arid distribution pattern,

being found in shrubland, spinifex plains, and sand

dunes in the rainfall distribution region < 360

mm isohyet. For zoogeographic purposes, the arid

distribution pattern of Notaden is not recognized

as a distinct pattern because there is only one

genus demonstrating the pattern.

The historical record of vegetation types in the

Australian region allows some generalization on

evolutionary trends and sequences within the Myobatrachidae. Details of past vegetation type distribution and paleoclimates in the Australian

region are not well known. The following summary, which appears to be sufficient to explain

major evolutionary and zoogeographic trends in

myobatrachids, seems reasonably substantiated by

the available data. The antarctic forest type (Notho-

(Continued on p. 22)

FIGURE 4.—Broad vegetation zones critical to analysis of

myobatrachid distribution patterns. (Solid areas = antarctic

vegetation unit; hatched areas = wet forests; stippled areas

= sandy deserts.)
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 5.—Geographic distribution of the myobatrachid FIGURE 6.—Geographic distribution of the myobatrachid

genus Adelotus. genus Assa.

FIGURE 7.—Geographic distribution of the myobatrachid

genus Australocrina.

FIGURE 8.—Geographic distribution of the myobatrachid

genus Crinia.
NUMBER 233 17

FIGURE 9.—Geographic distribution of the myobatrachid

genus Geocrinia.

FIGURE 10.—Geographic distribution of the myobatrachid

genus Glauertia.

FIGURE 11.—Geographic distribution of the myobatrachid

genus Heleioporus.

FIGURE 12.—Geographic distribution of the myobatrachid

genus Kankanophryne.
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 13.—Geographic distribution of the myobatrachid FIGURE 14.—Geographic distribution of the myobatrachid

genus Kyarranus. genus Lechriodus.

FIGURE 15.—Geographic distribution of the myobatrachid FIGURE 16.—Geographic distribution of the myobatrachid

genus Limnodynastes. genus Metacrinia.
NUMBER 233 19

FIGURE 17.—Geographic distribution of the myobatrachid

genus Mixophyes.

FIGURE 18.—Geographic distribution of the myobatrachid

genus Myobatrachus.

V

FIGURE 19.—Geographic distribution of the myobatrachid

genus Neobatrachus.

FIGURE 20.—Geographic distribution of the myobatrachid

genus Notaden.
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 21.—Geographic distribution of the myobatrachid FIGURE 22.—Geographic distribution of the myobatrachid

genus Paracrinia. genus Philoria.

FIGURE 23.—Geographic distribution of the myobatrachid FIGURE 24.—Geographic distribution of trie myobatrachid

genus Platyplectron. genus Pseudophryne.
NUMBER 233 21

FICURE 25.—Geographic distribution of the myobatrachid

genus Ranidella.

FICURE 26.—Geographic distribution of the myobatrachid

genus Rheobatrachvu.

V

FIGURE 27.—Geographic distribution of the myobatrachid

genus Taudactylus.

FIGURE 28.—Geographic distribution of the myobatrachid

genus Uperolcia.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fagus, podocarps) has had a long history in the

Australian region and during the early Tertiary,

at least, had a broader distribution than present.

Cookson (1953), Cookson and Pike, (1953, 1954),

and Darlington (1965) have shown that fossil

remains of the antarctic vegetation have been

found at several localities across southern Australia,

suggesting a former continuous forest belt. It is

probable that the extent of this forest unit has

been affected by world wide climatic changes,

especially glaciation cycles. Arid environments

have long been a feature of the Australian continent. The Great Dividing Range was formed at

the end of the Mesozoic, and arid environments

have been present in Australia throughout the

Tertiary to the present.

One of the most striking features in the evolution of myobatrachid frogs has been the lack of

radiation in the strict desert environment. The

desert habitat has been present in Australia for

a long enough time to allow a radiation. The

Australian deserts are apparently too harsh an

environment for frogs; some particular species can

tolerate desert conditions, but none are primarily

adapted for desert life. A similar situation occurs

with the Namib desert anuran fauna in South

Africa (Channing, 1974).

The antarctic forest vegetation appears to be

the vegetation type associated with the ancestral

myobatrachid stock. This assumption is supported

by the historical presence of this vegetation type

and by the distributions of several genera limited

strictly to this vegetation type. The genera presently restricted to the antarctic vegetation type

represent the descendants of the original stock

which was associated with the temperate antarctic

forests. If a generalized life history pattern is assumed for the ancestral stock, then the following

major evolutionary trends have occurred within

the beech and podocarp forests: trends toward

nonaquatic larvae; either direct development, inguinal pouch brooding, or gastric brooding; and a

stream adapted life history.

A second stage in the historical zoogeography

of the myobatrachids was the association with temperate wet forest (including Eucalyptus) types. The

same adaptive trends are seen in this assemblage

of genera as in the previous assemblage: trends toward direct development and stream adaptations.

A third stage was the evolution of forest forms

that could also live in more open formations. Adaptive trends in this group of genera include stream

adaptations, burrowing, and possibly direct development.

A fourth stage in the evolutionary history of the

myobatrachids involved adaptations leading to a

primary radiation in open, rather arid vegetation

types. The genera with this pattern either show a

distribution associated with water (oasis effect)

within a more arid region (Glauertia, Kankanophryne), or burrowers that can utilize temporary

bodies of water (Neobatrachus, Notaden).

To summarize, the major adaptations associated

with the four generic distribution patterns are as

follows:

Pattern 1 (Temperate/subtropical rainforest)

Gastric brooding: Rheobatrachus

Inguinal pouch brooding: Assa

Terrestrial egg

Nonfeeding larvae: Kyarranus

Steam adapted larvae: Taudactylus

Montane stream adaptations: Rheobatrachus, Taudactylus

Pattern 2 (Wet sclerophyll forest)

Terrestrial egg

Stream adapted larvae: Crinia, Mixophyes

Nonfeeding larvae: Philoria

Trend toward direct development: Geocrinia, part (leai,

laevis)

Direct development: Geocrinia, part (lutea, rosea), Metacrinia

Pattern 3 (Wet forest and subhumid woodland)

Rapid egg development: Platyplectron, part (ornatus,

spenceri)

Burrowing adaptations: Heleioporus, Platyplectron

Terrestrial egg

Steam adapted larvae: Australocrinia

Advanced larval development: Pseudophryne (except

douglasi)

Foam nest in burrow: Heleioporus

Possible direct development: Myobatrachus

Pattern 4 (Open woodland and shrubland)

Short larval life: Kankanophryne, Neobatrachus, Notaden

Burrowing adaptations: Neobatrachus, Notaden

Opportunistic breeding: Neobatrachus (part), Notaden

All four stages occurred roughly contemporaneously, early in the Tertiary at the latest. If the

four stages occurred in sequence, there should be

an overall reduction in the average number of

primitive states per genus from the first to the

fourth stage. This is not the case. The first three

groups all have an average of 55% primitive states

per genus, the fourth group has an average of 54%.

A rough indication of recency of evolutionary
NUMBER 233

activity can be gained by examining the average

number of species per genus. The assumption is

that lower number of species per genus represents

an older radiation, a higher number of species

per genus represents more recent speciation. The

first distribution pattern group has an average

number of species per genus of 2.6, the second

group an average of 2.0, the third group an average

of 5.0, and the fourth group an average of 3.25. If

the difference between the first and second group

is real, it could be interpreted in the following

way. The general wet forest associated group has

had a more continuous habitat available since the

Tertiary, while the antarctic forest associated group

has had a more fragmentary distribution, with populations of frogs becoming isolated in forest fragments sometime in the Tertiary. The isolation of

these latter populations has led to relatively more

speciation in this group than in the more general

wet forest associated group. This hypothesis is supported by the fact that most of the antarctic forest

associated genera that are not monotypic typically

have allopatrically distributed species. The fourth

group, primarily arid adapted, has an intermediate

amount of relatively recent speciation, while the

forest-open formation group has the greatest

amount of recent speciation. Apparently, in terms

of fluctuating environments there has historically

been greater fragmentation and coalescing of open

formation habitats. This has allowed speciation to

continue throughout the Tertiary to the present.

The phenomenon of more recent speciation in

open eucalypt habitat compared to rainforest habitat has also been observed in Australian birds

(Schodde and Calaby, 1972).

In summary, it appears that the major expansion

from the ancestral temperate antarctic forest habitat into other habitat types occurred more or less

synchronously; that the radiations within the major habitats are old, stemming from the early

Tertiary; and that the most recent evolutionary

activity is associated with open formation habitats,

intermediate between forest and desert environments.

Another overall trend is that direct development

is approached or attained only in wet forest genera,

as pointed out previously by Main, Littlejohn, and

Lee (1959). The only possible exception is Myobatrachus, which has huge ovarian eggs, suggesting

direct development.

EVOLUTIONARY TRENDS WITHIN THE SUBFAMILIES

When the historical zoogeography of the myobatrachids is viewed in terms of evolutionary relationships, additional trends can be postulated.

SUBFAMILY RHEOBATRACHINAE.—The sole member of this subfamily, Rheobatrachus silus, represents an early stream adaptational complex from

the ancestral myobatrachid stock. Rheobatrachus

has remained in the temperate antarctic forest

vegetation and represents a combination of a large

number of primitive states (67%) together with

some of the most bizarre derived states known

involving life history (Corben, Ingram, and Tyler,

1974).

SUBFAMILY MYOBATRACHINAE.—Members of the

Myobatrachinae demonstrate all four geographic

patterns; however, there are definite emphases

within the patterns, most of the genera being associated with wet forests. Taudactylus represents,

within the antarctic forest type, an early specialization to stream adaptations, with respect to the

larvae of all known species and all adults (Liem

and Hosmer, 1973). The Crmia-complex, represented by Assa, Australocrinia, Crinia, Geocrinia,

Paracrinia, Ranidella, all appear to represent more

or less independent specializations from the ancestral wet forest myobatrachine stock. Each generic

unit likely represents a previous isolation in a wet

forested area with concomitant adaptational responses to that given environment. The result of

several independent adaptational responses in a

generalized myobatrachine is a mosaic pattern of

derived states. The generalized myobatrachine

morphology has been retained, so the similarity

among these taxa is based on a suite of primitive,

rather than derived, states. Ranidella had the

genetic plasticity to adapt to environments bordering the wet forest habitat. The genera demonstrating the arid adaptation geographic pattern,

Glauertia and Kankanophryne, are exploiting local

mesic habitats within an arid landscape. The divergence of Glauertia into this specialized habitat

type probably occurred earlier than for Kankanophryne. Pseudophryne, Ranidella, and Uperoleia

have adapted in part to open formation vegetation

types. The large terrestrial egg of Pseudophryne

with initial development out of water has adaptive

value in areas of unreliable rainfall. There are no

particular obvious adaptations in the life history
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

patterns of Ranidella and Uperoleia to predict a

habitat shift to open formation vegetation. Myobatrachus is a very specialized form, a subterranean

termite eater. It is the only burrowing myobatrachid that can burrow head first and is restricted

to sandy areas in southwest Australia. Within the

Myobatrachine, it appears that Taudactylus,

Glauertia, and Myobatrachus represent the earliest

specializations and divergences from the common

myobatrachine ancestral stock. The remainder of

the myobatrachine genera appear to have had a

common ancestor that was widespread throughout

the late Mesozoic-early Tertiary wet temperate

forests, and that fragmentation of this common

stock associated with fragmentation of the temperate forests, with evolution proceeding independently in each unit, gave rise to the mosiac complex of genera present today.

SUBFAMILY LIMNODYNASTINAE.—Mixophyes appears to be an early specialization of the limnodynastine ancestral stock to the stream environment.

Neobatrachus and Notaden represent early specializations to the arid environment involving fossoriality in the adult stage. Notaden and Neobatrachus

are the most arid adapted myobatrachid genera

and, interestingly, have almost completely allopatric distributions. The evolutionary history of

the remaining limnodynastines is associated with

the foam nest. Four of the genera having a foam

nest are wet forest genera, the other three are

forest and open formation forms. Either the foam

nest was initially evolved in response to selective

pressures associated with wet forest environments

or open formation environments. The evidence

points to the original adaptive significance of the

foam nest as having been associated with wet

forests. None of the genera that are primarily arid

adapted have a foam nest, while several of the

genera with foam nests are found only in wet forest

habitats. The wet forest foam-nesting genera have

an average of 2.5 species per genus, contrasted

with a 5.7 average for the open formation foamnesting genera, suggesting the wet forest radiation

is older than the open formation radiation. The

foam nest, then, served as a preadaption for exploitation of a more seasonal environment associated with open formation vegetations. Lechriodus

is noteworthy among the myobatrachids in that

it is the only genus in which more species are

found outside of Australia than within Australia.

Lechriodus seems to be associated with the podocarp vegetation unit of the temperate antarctic

vegetation and has undergone a minor radiation

associated with, in part, the podocarp forests of

New Guinea. There is no evidence, however, to

suggest that the genus arose in New Guinea; in

fact, the Australian species is the most primitive in

the genus (Zweifel, 1972).

MAJOR TRENDS AMONG SUBFAMILIES

Even though the adaptive trends within the

Myobatrachinae and Limnodynastinae are similar

in that they include adaptational complexes associated with open formations and arid environments, direct development and stream habitats,

the adaptations associated with these shifts are

different. The myobatrachines have no truly arid

adapted genera; the limnodynastines have two.

Direct development involves a foam nest in the

limnodynastines; a foam nest is not involved in

direct development in the myobatrachines. The

foam nest is the key preadaptation to open formation utilization by some of the limnodynastines;

the foam nest is not a feature of the open formation myobatrachines.

Within all three subfamilies, the ancestral pattern appears to be an association with temperate

antarctic vegetation and a generalized life history

pattern involving a free living larva. The most

parsimonious explanation of the early evolution of

the myobatrachids is that the present myobatrachids represent three very early divergences in

the myobatrachid ancestral stock. One lineage

became adapted as adults to the stream habitats

with an extremely specialized life history pattern

of gastric brooding. The degree of specialization

apparently led to an evolutionary dead end represented in Rheobatrachus silus. The early divergence of the myobatrachine and limnodynastine

stocks involved morphological features which at

present are difficult to associate with adaptive

trends. Both groups were associated with the temperate antarctic vegetation, and they likely had a

generalized life history pattern. The basic morphological differences between the two stocks allowed

their evolution to act in independent ways in relation to the same environmental changes of increasing aridity in the late Mesozoic to present.
NUMBER 233 25

COMPARISONS WITH PREVIOUS ZOOGEOGRAPHIC

SCHEMES

Previous workers in discussing the zoogeography

of the Australian anuran fauna have proposed

three major ideas. The first is that a segment of

the anuran fauna is old, stemming at least from

the early Tertiary. The second is that the anuran

fauna can be explained best by using a modified

Spencerian (1896) scheme of four faunal subregions

in Australia. The third is the proposal of a mode

of speciation in some genera involving evolution

of species in southeast Australia with dispersal to

southwest Australia.

This study confirms the importance of recognizing the antiquity of the myobatrachid frog

fauna. The myobatrachids were certainly present

in the Australian region by late Mesozoic and

much of the radiation of the group may well have

occurred in early Tertiary.

The faunal subregions used by most workers to

explain the geographic distribution are as follows

(from Littlejohn, 1967:152):

(1) Bassian—coastal, cool temperate, relatively uniform rainfall; east and south-east of the Eastern Highlands and south

of the Clarence River; (2) Torresian—northern and eastern

tropical area north of the Clarence River with a summer

maximum rainfall; (3) Eyrean—all the drier inland portions

of the continent, including the north-west coast; (4) Southwest—the humid south-western corner of Western Australia

which receives a reliable winter maximum rainfall.

Moore (1961) has been the only recent worker

that did not use the above scheme. Moore's scheme,

in part, recognized two additional regions and he

emphasized that only one of his regions was truly

distinctive: that corresponding with the South-west

pattern described above.

None of the myobatrachids have a strictly Torresian distribution. Several of the genera have

distributions combining Bassian and Torresian

distributions. Most of the genera with distributions

in the Eyrean region also are found in the Bassian,

the Torresian, and/or the South-west regions.

Neither the modified Spencerian nor the Moore

(1961) schemes adequately describe the distribution

patterns of the myobatrachids. We think the key

to understanding the zoogeography of the myobatrachids is the recognition of the temperate antarctic vegetation associated myobatrachid fauna as a

distinctive unit.

The Spencerian scheme appears to account much

better for reptile distributions than for myobatrachid distributions (e.g. Keast, 1959; Kluge, 1967).

We think that the Spencerian subregion scheme

associates best with those faunal units having a

tropical origin. For those faunal units having a

temperate origin, such as the myobatrachids, the

scheme proposed herein apparently is a more

realistic zoogeographic hypothesis.

Lee (1967:428-429) summarized the speciation

scheme proposed for certain genera:

Faced with the incongruous situation of a multiplicity of

species within Heleioporus and two other genera of frogs

(Crinia and Neobatrachus) in the topographically featureless

south-western pocket. Main, Lee, and Littlejohn (1958)

suggested the following explanation for their origin:

(1) Within each genus two elements or species groups were

recognized and identified as bassian and eyrean elements

(after the terminology of Serventy and Whittell (1951).

The breeding biology of the bassian frogs was adapted

to uniform rainfall or continuously moist situations, and

that of the eyrean species to seasonal rainfall and seasonally arid conditions. Designation of the eastern species

based on these adaptations presented no problems, but

with the specialization of the western fauna to the pronounced seasonal conditions of the area, use had to be

made of similar morphology to provide clues of relationship. Judged by their morphological dissimilarity, this

division into two elements was a pre-Pleistocene event.

(2) South-eastern rather than south-western Australia was

favoured as the centre of origin of the genera. The southeast is more diverse topographically and ecologically,

and despite the greater number of species in these genera

in south-west, there are more species and genera of frogs

in the south-east.

(3) The multiplicity of species within each genus in the

south-west resulted from a series of invasions of the

eastern representatives of each element into the southwestern pocket where some persisted. The invasions were

separated by periods of isolation of sufficient duration for

the populations to retain separate identity when brought

together again.

(4) The invasions were considered to be coincident with the

pluvial or glacial phases, and isolation with the arid interglacial phases of the Pleistocene. The Pleistocene

events of the northern hemisphere were probably paralleled in the southern hemisphere, but only with minor

continental glaciation. Browne (1957) and Browne and

Vallance (1963) have been able to recognize three phases

of glaciation in southeastern Australia, and this corresponds to the number of western eyrean representatives

in each genus, but not to the number of bassian species,

of which there are either one or two. Main, Lee, and

Littlejohn (1958) suggested that early bassian invaders

may have succumbed to the marked arid conditions of

later interglacial phases.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(5) Finally, the disproportionate number of species in eastern and western Australia was attributed to the failure

of the western representatives to reinvade the east after

specialization during isolation.

The speciation scheme includes recognizing

species pairs in southeast and southwest Australia.

The kinds of information used to determine relationships have been morphological similarity, mating call similarity, and hybridization results. The

authors who support this mode of speciation do

not distinguish primitive and derived states; thus

the morphological and mating call similarities

could be based on primitive states and contain no

information regarding relationships. Hybridization

data as an index to relationships among anurans

must be used with great caution. There are many

examples of viable intergeneric crosses. Postmating

isolating mechanisms involving hybrid inviability

may only be selected for in closely related forms

that have had historical overlap in geographic

ranges. We present these arguments not to invalidate the conclusions of the authors who have proposed this mode of speciation, but to indicate that

there is room for another hypothesis.

The hypothesis we favor is that for such genera

as Heleioporiis, during glacial periods, when the

mesic forests would have had an expanded distribution, local arid habitats may have persisted due

to such factors as drainage peculiarities of the soil.

These isolated arid habitats would serve as refugia

for the arid adapted fauna. Due to the isolated

nature of these arid refugia, evolution could operate independently on the faunas of each of them

allowing allopatric speciation to occur. The origin

of the species would be within the same general

region where they are found at present, not in

southeast Australia exclusively.

MYOHATRACHID-LEPTODACTYLID COMPARISONS

The leptodactylid radiation in the Neotropics

resulted in at least 47 genera and 500 species. The

vastness of this radiation appears to overwhelm

the myobatrachid radiation which resulted in 24

genera and about 80 species. The comparison of

the total leptodactylid and myobatrachid radiations

is in one sense unrealistic, since the myobatrachids

have not radiated into tropical environments in

the way that leptodactylids have. A more meaningful comparison is to compare that portion of the

leptodactylid radiation that corresponds with the

myobatrachid radiation. The myobatrachid history

includes original association with temperate forests

and evolution into open formations and arid environments. A comparable situation is found in

South America in the case of the anuran fauna of

Chile and adjacent Argentina, including the Gran

Chaco, but excluding Misiones and east coastal

Argentina. In this Neotropical region, the original

association of the leptodactylids was with the temperate forests and radiations have occurred into the

open formations and arid environments.

Although the fauna of southern South America

is reasonably well understood, the leptodactylids of

Chile and Argentina are not as well understood

systematically as are the myobatrachids of Australia.

There are 15 genera and 50 species of leptodactylids found in the area of comparison in Chile

and Argentina. This larger number of taxa indicates that the myobatrachid radiation has been

larger than the leptodactylid radiation in similar habitats. Interestingly, both the southern

leptodactylids and myobatrachids have the same

average number of species per genus, 3 1/3, suggesting that the radiations are of equivalent age.

A feature of the myobatrachid radiation is that

there are relatively few species per genus, Ranidella

and Pseudophryne being high with 8 and 9 known

species. This compares poorly with the 300 plus

Neotropical species of Eleutherodactylus or even

the 20-40 species in such Neotropical genera as

Leptodactylus and Physalaemus. When one examines the leptodactylid fauna of Chile and Argentina, the situation is exactly that found in the

myobatrachids. Leptodactylus, Pleurodema, and

Telmatobius are all high with a total of 8 known

species.

The more extensive radiation of the myobatrachids than the southern leptodactylids appears

to be valid. The only other group of frogs with

extensive distributions in Australia is the tree frog

family Pelodryadidae. The tree frog family Hylidae

is well represented in southern South America. In

addition, there are two bufonid genera, two

microhylid genera, and one pseudid genus with

extensive distributions in Chile and Argentina.

This last set of genera appears to have ecological

equivalents among the myobatrachids. The most

striking example is the fossorial termite eating
NUMBER 2S3 27

myobatrachid Myobatrachus and the fossorial termite eating microhylid Dermatonotus mulleri.

There is, incidently, only one such species in each

region. If this last set of genera is added to the

leptodactylids, there is a total of 20 genera and

64 species. This figure is rather close to the

myobatrachid figure, taking into account the fact

that the southern South American anuran fauna

is not as well known as the Australian anuran

fauna and the total area of comparison is smaller

for southern South America. It appears as though

the total anuran frog faunas of these two regions

are equivalent and that the myobatrachid radiation

has been more extensive than the leptodactylid.

Convergence of adaptive complexes between the

myobatrachids and southern leptodactylids is striking. In both groups there are riparian forms,

fossorial forms, and trends toward and including

direct development. Certain of these are worth

more detailed comment.

Parental brooding of the larvae is found in both

the myobatrachids and southern leptodactylids.

The evolution of this pattern, however, has clearly

occurred three different times. The myobatrachid

Assa has male inguinal brooding pouches, the

myobatrachid Rheobatrachus has a female gastric

brooding pouch, and the leptodactyloid

Rhinoderma has a male vocal sac brooding pouch.

The evolution of the foam nest has had a similar, but independent, evolutionary history in both

the myobatrachids and leptodactylids. In both

groups, the original adaptive significance of the

foam nest was in response to selective pressures

associated with wet forest environments. The presence of the foam nest then served as a preadaptation allowing radiation into open formation

habitats. The known methods of foam nest formation are very different, as pointed out by Martin

(1970). In the myobatrachids, the fingers of the

female are modified to whip the jelly into foam,

while in the leptodactylids the hind legs of the

male whip the jelly into foam.

Recently, a complex association of specialized

larval microhabitats and morphology has been

described for Leptodactylus pentadactylus (Heyer,

McDiarmid, and Weigmann, 1975). Briefly, L.

pentadactylus, a foam nest breeder, utilizes small

aquarium sized puddles as larvae. These puddles,

occurring in wet environments, are replenished by

almost daily rainfall, so they do not dry up. The

larvae are facultatively carnivorous, a rare condition among tadpoles. It was postulated that the

carnivorous tadpole mode is possible in these particular microhabitats because there are no fish in

these small puddles (see Heyer, McDiarmid, and

Weigmann, 1975, for a fuller explanation).

Lechriodus fletcheri, a wet forest associated, foam

nest breeding myobatrachid, also utilizes small

aquarium sized puddles and has facultatively carnivorous larvae.

The myobatrachids and leptodactylids most

likely had a common ancestor that was widespread

throughout the Antarcto-Tertiary Geoflora of

Gondwanaland. Both the myobatrachids and

leptodactylids have members that are still associated with this vegetation. Arid centers developed

independently in Australia and South America.

The convergence of life histories and morphologies

of the myobatrachids and leptodactylids with derivative temperate forests, open formation vegetations,

and arid environments is nothing short of spectacular.
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