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I

Evolutionary Relationships,
Osteology, and Zoogeography of
Leptodactyloid Frogs

By

John D. Lynch

UNIVERSITY OF KANSAS

LAWRENCE 1971 June 30, 1971

UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY

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Kansas 66044.

University of Kansas
Museum of Natural History

Miscellaneous Publication No. 53
June 30, 1971

Evolutionary Relationships, Osteology, and

Zoogeography o£

Leptodactyloid Frogs

By

John D. Lynch

Department of Zoology

University of Nebraska

Lincoln, Nebraska 68508

University of Kansas

Lawrence

1971

1
University of Kansas Publications, Museum of Natural History

Editors: Craig C. Black, William E. Duellman,
Philip S. Humphrey

Miscellaneous Publication No. 53

pp. 1-238; 131 figures

Pubhshed June 30, 1971

Museum of Natural History

University of Kansas

Lawrence, Kansas 66044

U.S.A.

Printed by

University of Kansas Printing Service

Lawrence, Kansas

CONTENTS

INTRODUCTION .. 7

Acknowledgments 8

Nomenclatoiial Resume 8

Materials and Methods 17

Generic Concept 18

ANALYSIS OF CHARACTERS 19

Non-Osteological Characters 21

Amplectic Position 22

Breeding Behavior and Development 24

Tadpole Morphology 25

Secondary Sex CJiaracters 28

Pupil Shape 29

Hands and Feet 30

Body Glands 32

Myology 33

Hyolaryngeal Apparatus 34

Chromosome Numbers 36

Osteological Characters 37

Maxillary Arch 38

Lower ]aw 42

Roof of the Skull 43

Temporal Architecture 47

Palate 48

Otoccipital and Columella 52

Vertebral Column 53

Pectoral Girdle 58

Pelvic Girdle 60

Limb Elements 64

SYSTEMATIC ACCOUNTS 67

Genera Excluded from the Leptodactylidae 68

Geobatrachus Ruthven, 1915 69

Hylopsis Werner, 1896 69

3

Rhinoderma Dumeril and Bibron, 1841 69

Keys to the Genera of Leptodactylid Frogs 71

Key to Africa7i and Australo-Papiian Genera 71

Key to African and Neotropical Genera 72

LEPTODACTYLIDAE Berg, 1896 ( 1838) 75

CYCLORANINAE Parker, 1940 75

Cycloranini Parker, 1940 76

Cyclorana Steindachner, 1867 76

Heleioporus Gray, 1841 78

Mixophyes Giinther, 1864 79

Neobatrachiis Peters, 1863 81

Notaden Giinther, 1873 82

Limnodynastini New Tribe 83

Adelotus Ogilby, 1907 84

Lechriodns Boulenger, 1882 85

Limnodytuistes Fitzinger, 1843 86

Kyarranus Moore, 1958 -— 88

Philoria Spencer, 1901 89

MYOBATRACHINAE Schlegel, 1850 90

Crinia Tschudi, 1838 91

Pseudophryne Fitzinger, 1843 94

Taudactylus Straughan and Lee, 1966 95

Glauertia Loveridge, 1933 97

Uperoleia Gray, 1841 99

Metacrinia Parker, 1940 100

Myohatrachus Schlegel, 1850 101

ilndohatrachus Noble, 1930 103

HELEOPHRYNINAE Noble, 1931 103

Heleophryne Sclater, 1898 104

CERATOPHRYINAE Tschudi, 1838 105

Ceratophrys Wied, 1824 107

Lepidobatrachus Budgett, 1899 110

fWawelia Casamiquela, 1963 112

TELMATOBIINAE Fitzinger, 1843 112

4

Telmatobiini Fitzinger, 1843 113

Caudiverbera Laurenti, 1768 114

Telmatohufo Schmidt, 1952 116

Telmatohius Wiegmann, 1835 118

■\Neoprocoela Schaeffer, 1949 120

Batrachophrynus Peters, 1873 122

Alsodini Mivart, 1869 : 123

Eupsophus Fitzinger, 1843 124

Hylorina Bell, 1843 126

Batrachyla Bell, 1843 128

Thoropa Cope, 1865 129

Odontophrynini New Tribe 130

Odontophrynus Reinhardt and Liitken, 1862 131

Proceratophrys Miranda-Ribeiro, 1920 133

Grypiscini Mivart, 1869 135

Crossodactylodes Cochran, 1938 136

Cycloramphus Tschudi, 1838 137

Zachaenus Cope, 1866 139

Eleutherodactylini Lutz, 1954 142

Amblyphrynus Cochran and Coin, 1961 143

Eleutherodactylus Dumeril and Bibron, 1841 144

Euparkerella Griffiths, 1959 150

Holoaden Miranda-Ribeiro, 1920 152

Hylactophryiie Lynch, 1968 153

Ischnocnerna Reinhardt and Liitken, 1862 155

Niceforonia Coin and Cochran, 1963 156

Sminthillus Barbour and Noble, 1920 158

SyrrJwphtis Cope, 1878 159

Tomodactylus Giinther, 1900 160

Tribe incertae sedis 162

Scythrophrys new genus 162

ELOSIINAE Miranda-Ribeiro, 1926 163

Crossodactylus Dumeril and Bibron, 1841 164

Hylodes Fitzinger, 1826 166

5

Megaelosia Miranda-Ribeiro, 1923 168

LEPTODACTYLINAE Berg, 1896 (1838) 169

Pleurodeniui Tschudi, 1838 172

Limnomedusa Fitzinger, 1843 175

Hydrolaetare Gallardo, 1963 176

Edalorhina Jimenez de la Espada, 1870 177

Litlxodijtes Fiizingex, 1843 179

Phijsalaemus Fitzinger, 1826 180

Paratelmatohius Lutz and Carvalho, 1958 182

Pseudopahidicola Miranda-Ribeiro, 1926 184

Leptodactijhis Fitzinger, 1826 186

Banjcholos Heyer, 1969 190

PHYLOGENY AND RELATIONSHIPS 192

The Fossil Record 192

Pelobatid-Leptodactylid Relationships 198

Intrafamilial Relationships of the Leptodactylidae 204

Extrafamilial Relationships of the Leptodactylidae 209

ZOOGEOGRAPHY 211

SUMMARY AND CONCLUSIONS 221

APPENDIX 225

LITERATURE CITED 229

6

INTRODUCTION

Frogs of the family Leptodactylidae
(auctomm) are distributed throughout
the Neotropics north to Texas and Flor-
ida, in southern Africa, and over most of
the Australo-Papuan region (Fig. 1).
The family is one of the largest amphib-
ian families and was known to the early
describers of the world's fauna. Linne
(1758) knew two species of the family,
Rana ocellata (^=Leptodactylus ocella-
tus) and Rana cormita (^Ceratophnjs
cornuta), and the tadpole of a third, La-
certa caudiverhera (=Caudwerbera cau-
diverbera), all from South America. The
first representative of the Australian
fauna was not described until Shaw's
(1795) work naming Rana australiaca
(=:Heleioporus australiacus). The fam-
ily, as presently conceived, contains ap-
proximately 650 species in 57 living gen-
era arranged in seven subfamilies. Two
of these subfamilies are restricted to
Australia, New Guinea, and Tasmania,

one to the Cape region of southern
Africa, and the other four to the Neo-
tropics. Seventeen genera are restricted
to the Australo-Papuan realm, one to
south Africa, and 39 to the Neotropics.

Three factors have contributed to our
current deplorable lack of general knowl-
edge concerning this large family: 1) the
most recent synopsis of a revisionary na-
ture is that of Boulenger, 1882; 2) the
bewildering number of generic names
and the indiscriminant application of
family and subfamily names of numer-
ous authors; and 3) the problem of
dealing with Eleutlierodactyliis, one of
the largest vertebrate genera known. No
part of this genus (either systematic or
geographic) has been monographed.

Initially, this study was restricted to
a survey of the skeletal morphology of
frogs of the genus EletitJierodactylus;
however, this proved impractical, be-
cause of the lack of understanding of

Figure 1. Distributions of the family Leptodactylidae (stipple) and the pelobatid subfamily Mego-

phryinae ( hatching ) .

8

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

intrageneric relationships. Therefore, I
decided to re-evaluate the genera of the
family and to attempt to point out supra-
generic relationships. Detailed study and
analysis of infrageneric relationships
have, of necessity, been postponed, pend-
ing a more complete and realistic under-
standing of the genera involved.

ACKNOWLEDGMENTS

This study could not have been com-
pleted were it not for the generosity of
curators of numerous institutional and
private collections. For loans and/ or
gifts of specimens I am grateful to Wer-
ner C. A. Bokermann, Sao Paulo; the late
Doris M. Cochran and James A. Peters,
United States National Museum; Alan
Leviton, California Academy of Sciences
( CAS ) ; Alice G. Grandison, British Mu-
seum ( Natural History ) ; Robert F. Inger
and Karel Liem, Field Museum of Nat-
ural Histoiy; Celia Limeses and Maria
Tia, Laboratorio de Investigaciones
Herpetologicas, Buenos Aires; A. R.
Main, Department of Zoology, Univer-
sity of Western Australia, Nedlands;
Albert Schwartz, Miami; Hobart M.
Smith, University of Illinois Museum of
Natural History; Glen M. Storr, West-
ern Australian Museum, Perth; I. R.
Straughan, Department of Zoology, Uni-
versity College of Townsville, Australia;
Paulo Vanzolini, Museu de Zoologia,
Universidade de Sao Paulo; C. F. Walk-
er, University of Michigan Museum of
Zoology; E. E. Williams, Museum of
Comparative Zoology; and R. G. Zweifel,
American Museum of Natural History.
The majority of the material used in the
study is deposited in the University of
Kansas Museum of Natural History col-
lection or in my own skeleton collection.
Special thanks are extended to Werner
C. A. Bokermann for always responding
to requests for yet another specimen of
an uncommon Brasilian frog and to
James A. Peters and Richard G. Zweifel
for pennitting preparation of skeletons
of paratypes of Crossodactylodes pintoi
and Noblella peruviana, respectively.

Werner C. A. Bokermann, Jose Cei,
M. J. Littlejolm, and J. A. Moore were
especially helpful in providing informa-
tion concerning reproduction. William
E. Duellman, Arthur C. Echternacht, Jay
M. Sa\age, Hobart M. Smith, Gerald R.
Smith, Linda Trueb, Edward H. Taylor,
and Charles F. Walker offered construc-
tive critcism during the evolution of my
concept of the evolution of leptodac-
tyloid frogs. Special thanks are given to
William E. Duellman, under those guid-
ance this study was made; Theodore H.
Eaton, George W. Byers, and Gerald R.
Smith, read and criticized the manu-
script.

Field support was provided by grants
from the Committee on Systematics and
Evolutionary Biology, University of Kan-
sas; a University of Kansas Graduate
School Honors Fellowship; The Society
of the Sigma Xi; and the Watkins Mu-
seum of Natural History Grant in Aid of
Research, University of Kansas. For as-
sistance in the field I am grateful to
Carlos Arroyo, William E. Duellman,
Jerry James, Federico Medem, Juan
Leon, Marsha Lynch, and Jesus Perez.

NOMENCLATORIAL RESUME

More genera are contained in the
Leptodactylidae than in any other am-
phibian family. The most recent author
to monograph the Leptodactylidae was
Boulenger ( 1882 ) , whose analysis suf-
fered from a misunderstanding of what
characters were of use in assessing fami-
lies (Noble 1922, 1931). Boulenger
( 1882 ) recognized 179 species of lepto-
dactylids and placed them in 34 genera
in 7 famihes. No author subsequent to
Boulenger has treated any major section
of the Neotropical leptodactylid fauna,
other than Parker (1927), who treated
the frogs then placed in Paludicola
[^=PhysaIaenms, Pleiirodema, and Pseu-
dopaludicoki], although several works of
major importance have appeared treat-
ing faunal units and therefore parts of
leptodactylid genera (Cei, 1962a; Coch-
ran, 1955; and Cocliian and Coin, 1970).

LYNCH: LEPTODACTYLOID FROGS

9

Gallardo ( 1965 ) presented a "synopsis
of leptodactylids in South America" but
this work in my opinion falls far short of
its intended goal. Parker ( 1940 ) and
Moore ( 1961 ) pro\'ided a good under-
standing, albeit incomplete, of the Aus-
tralo-Papuan leptodactylids. The status
of the species of Heleophnjne in South
Africa is now relatively well known.

Twenty-three family group names
have been proposed for frogs based on
type-genera which are here included in
the Leptodactylidae (Table 1). A num-
ber of other family group names have
been applied to leptodactylid frogs but
do not have nomenclatural significance
in a classification of the leptodactylids.

Tschudi ( 1838 ) proposed two family
group names, Cystignathi and Cerato-

phrydes, based on Cystignathus Wagler,
1830, and Cemtophrys Wied, 1824, re-
spectively. Although the Cystignathi
was widely used ( Cystignathidae of
Boulenger, 1882; Nieden, 1923), few
authors (Cope, 1866 and Miranda-Ri-
beiro, 1926) used the Ceratophrydes (or
an emendation of it). The concept of
the group changed whenever it was
used. Parker (1935) pointed out that
Noble's ( 1931 ) Pseudinae was probably
an artificial assemblage and regardless
should take the oldest available name,
Ceratophryinae. Recently, several au-
thors have proposed using Ceratophryi-
dae (or an emendation of this name) as
a family containing three genera of Neo-
tropical frogs. With the discovery that
Cystignathtis was an obligate synonym

TABLE 1. Family group names proposed in the Leptodactylidae and the subfamilial

or tribal name used herein.

Name proposed Name used herein

Cystignathi Tschudi, 1838 Leptodactylinae

Ceratophrydes Tschudi, 1838 Ceratophryinae

Telmatobii Fitzinger, 1843 Telmatobiinae, Telmatobiini

Myobatrachidae Schlegel, 1850 Myobatrachinae

Uperoliidae Giinther, 1859 Myobatrachinae

Hylodidae Giinther, 1859 Eleutherodactylini

Criniae Cope, 1866 Myobatrachinae

Pleurodemae Cope, 1866 Leptodachtylinae

Alsodina Mivart, 1869 Alsodini

Cacotina Mivart, 1869 Alsodini

Paludicolina Mivart, 1869 Leptodactylinae

Plectromantidae Mivart, 1869 Leptodactylinae

Giypiscina Mivart, 1869 Grypiscini

Leptodactylidae Berg, 1896 (1838)^' Leptodactyhnae

Elosiidae Miranda-Ribeiro, 1926 Elosiinae

Heleophryninae Noble, 1931 Heleophryninae

Cycloraninae Parker, 1940 Cycloraninae, Cycloranini

Telmatobiinae Vellard, 1951 [Homonym]

Cyclorhamphinae Lutz, 1954 Grypiscini

Eleutherodactylinae Lutz, 1954 Eleutherodactylini

Calyptocephalellinae Reig, 1960 Telmatobiini

Batrachylinae Gallardo, 1965 Alsodini

Pseudopaludicolinae Gallardo, 1965 Leptodactylinae

Limnodynastini Proposed herein

Odontophrynini Proposed herein

" The family name Leptodactyhdae is a conserved name in the sense of Article 40 (International
Code of Zoological Nomenclature, 1961) and hence dates from 1838, not 1896, for purposes of priority.

10

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

of Leptodactyhis Fitzinger, 1843, the
family gioup name Cystignathidae be-
came unavailable according to the rules
then in effect, and Leptodactylidae was
proposed as a family replacement name
— this name has gained nearly universal
acceptance and is thus a conserved name
in the sense of the provisions of Article
40 of the International Code of Zoologi-
cal Nomenclature ( 1961 ) and takes pri-
ority over any other names proposed
between 1838 and 1896. Therefore the
oldest valid family group name for these
frogs is Berg's ( 1896 ) Leptodactylidae.
Tschudi ( 1838 ) knew eleven genera and
twelve species of frogs that are now
placed in the Leptodactylidae. He ar-
ranged them in five families — Bombina-
tores (Pletirodema and Telmatobius),
Ceratophiydes (Ceratophnjs and Phryn-
oceros), Cystignathi (Cystignotlnis and
Crinia), Hylae (Cornufer, Elosia, and
two species of Hylodes), and Ranae (Cy-
doramphus and Peltoceplialus).

Fitzinger, an active and often erratic
classifier, proposed ( 1843 ) placing the
25 genera known to him in 7 families —
Alytae (Calyptocephalus, Ceratophrys,
Crinia, Hammatodactyhis, Leiuperus,
Phrynoceros, Stomljiis, and Tomopterna),
Gastrophrynae (Pcdiidicola), Limnodytae
(Euhyas, Hylodes, and Lithodytes), Pe-
lobii (Enydrohius), Phyllobatae [an older
family name for Dendrobatidae (Crosso-
dactylus and Scinacodes)], Ranae (Cys-
tignatJnis, Cycloramphtis, Eupsophus,
Gnathophysa, Leptodactyhis, Limnody-
nastes, Limnomedusa, Physalaemus, and
Sihilatrix). Burger (1954) correctly
pointed out that the terminal endings of
family group names such as these pro-
posed by Fitzinger do not affect their
priority and that Vellard's (1951) Tel-
matobiinae is a homonym of Fitzinger's
Telmatobii.

Giinther ( 1859a ) attempted a world-
wide catalogue of the amphibians, and,
although less successful than Boulen-
ger's (1882) second edition of this im-
portant project of the British Museum, it
did present a synthesis of most of the

genera of frogs as then known. Giin-
ther's family concepts were very differ-
ent from those used during the following
century, which explains in part why he
placed the 23 leptodactylid genera
known to him in elcxen families — Alyti-
dae (Heleioporus), Brachycephalidae
(Pseudophryne), Bombinatoridae (Al-
sodes and Tehnatohiiis), Cystignathidae
(Cystignathus, Leiuperus, Limnodynas-
tes, and Pleurodema), Discoglossidae
(Chiroleptes), Engystomatidae (Chelydo-
batracJms), Hylodidae (Crossodactylus
and Hylodes), Myobatrachidae (Myo-
hatrachus), Polypedatidae (Cornufer and
Elosia), Ranidae (Calyptocephalus, Cera-
tophrys, Cyclorhamphus, Hylorhina,
Limnocharis, Pithecopsis, and Pyxiceph-
alus), and Uperoliidae (Uperoleia). Of
these eleven families, the Hylodidae and
Uperoliidae represented new groups
based on leptodactylid genera. The
Hylodidae was based on Hylodes of
Fitzinger (1843, not 1826), and thus ap-
plies to the group I designate as Eleu-
therodactylini; in addition, the type
genus, Hylodes Fitzinger (1843), is a
homonym. Myobatrachidae was first
used by Schlegel (1850). It perhaps re-
flects the difficulties of communication
that Giinther recognized both Chelydo-
hatrachus and Myobatrachus, which are
synonymous, and included them in dif-
ferent families, Engystomatidae and
Myobatrachidae.

Cope (1866) considerably altered the
previous classifications, and relied so
heavily on osteological characters that
most subsequent authors have over-
reacted (in my opinion) in arguing that
the osteocranium cannot reasonably
serve in anuran classification. Cope rec-
ognized 37 genera of leptodactylid frogs
and placed all but two of these in the
Cystignathidae. Cryptotis (^Adelotus)
was placed in the Asterophrydidae and
Thoropa in the Hyhdae. Cope placed
the other 35 genera in six subfamilies,
two of which (Criniae and Pleurodemae)
were new. Cope recognized the Cera-
tophrydes (Ceratophrys, Chiroleptes,

LYNCH: LEPTODACTYLOID FROGS

11

Limnomedusa, Stombus, Tomopterna,
and Zachaenus), Criniae (Abodes, Bor-
horocaetes, Crinia, Cycloramplms, Etiso-
plnis, Helioporus, Hyperoleio, Isleohatra-
chus, and Plotyplectnim), Cystignathi
(Cys-ti<i,natJius, Gnathophysa, Gompho-
bates, and Torsopterus), Hylodes (Enhy-
drobius, Epirhexis, Hylodes, Limno-
charis, Litlwdytes, Pliyllobates, and Stra-
bomantis), Pleurodemae (Hylorhma, Liu-
perus, and Pleuwdema) and Pseudes
(Mixophyes, Pitliecopsis, Lysapsus and
Pseudis). The genera Batrachyla, Nat-
tereria, Plectromantis, and Tehnatobius
were included in the Cystignathidae but
not assigned to any of the six subfami-
Hes. Noble (1931) used Cope's Criniae
(as Criniinae) for all Australo-Papuan
leptodactylids, and the classification he
put forth bears sufficient resemblance to
Cope's (1<S66) that it seems reasonable
to assume that Cope's classification
served as Noble's prototype.

Mivart (1869) attempted to integrate
the two classifications proposed by Giin-
ther (lS59a) and Cope (1866) but in
my opinion did not contribute to a bet-
ter understanding of anuran relation-
ships. He continued the use of two un-
reliable characters — reduction (or loss)
of the auditory apparatus and loss of
maxillary teeth. He placed 38 leptodac-
tylid genera in 9 families and proposed
several new (and unnecessary) subfam-
ily names — Pseudophryne was placed in
the Phryniscidae ( Phryniscina ) ; Adeno-
mera, Chelydobatrachtis, and Poludicola
in three subfamilies ( Engystomina, Bre-
vicipitina, and Paludicolina) of the En-
gystomidae; AJsodes and Tebnatobius in
the Alsodina and Cacotus in the Caco-
tina, both subfamilies of the Bombina-
toridae; Plectromantis was the only
member of the Plectromantidae; Hyper-
olius {^Uperoleia, nee Rapp, 1842),
Helioporus, and Nattereria in the sub-
family Uperoliina (Alytidae); Thorpa
I in the Hylidae; Leiyla in the Acridina
and Crossodoctylus, Elosia, Epirhexis,
Hylodes, and Strabomantis in the Hylo-
dina ( Polypedatidae ) ; Odontophrynus

was placed in the Ranina (Ranidae)
and Ceratophrys, Crinia, Cycloramphus,
Cystignathus, Eusophleus (=Eupso-
phus), Hylorhimu Leiiiperiis, Lim-
nocharis, Limnodynastes, Mixophyes,
Neobatrachus, Pithecopsis, Platyplec-
trtini, Pleurodema, and Zachaenus in the
Cystignathina (Ranidae); and four gen-
era were placed in the Discoglossidae,
Calyptocephalus and Chiroleptes (Chi-
roleptina), Cryptotis ( Asterophiydina )
and Gryplsciis ( Grypiscina ) . Of these
family group names, six were new and
based on leptodactylids — Alsodina, Ca-
cotina, Chiroleptina, Grypiscina, Paludi-
colina, and Plectromantidae. Parker
( 1940 ) pointed out that the Chiroleptina
cannot be used because the type genus is
a homonym; this was the only family
group name proposed for the group
Parker (1940) named the Cycloraninae.
Paludicolina and Plectromantidae are
synonymous with the Leptodactylinae.
Alsodina and Cacotina are based on type
genera which are synonymous (Lynch,
1968b) with Eupsophus but are other-
wise available and older names for Gal-
lardo's (1965) Batrachylinae. Grypis-
cina is based on Grypiscus, a synonym
of Cycloramphus, and thus is an older
name for Lutz's ( 1954 ) Cycloramphinae.
Boulenger (1882) recognized 34 gen-
era of leptodactylids and placed these in
seven famifies — Grypiscus ( Amphigna-
thodontidae ) , Engystomops, Myobatra-
chus, Notaden, and Pseudophryne (Bu-
fonidae), Batrachophrynus (Dendro-
phryniscidae ) , Adenomera (Engystomat-
idae), Thoropa (Hylidae), Batrachopsis
and Ranmter (Pelobatidae), and 28 gen-
era (two were members of other fami-
lies) in the Cystignathidae [Abodes,
Borborocoetes, Calyptocephalus, Cera-
tophrys, Chiroleptes, Crinia, Cryptotis,
Cyclorhamphus, Edalorhina, Elosia, He-
leioporus, Hylodes, Hylorhina, Hypero-
leia, Leptodactylus, Limnodynastes,
Limnomedusa, Mixophyes, Nattereria,
Paludicola, Pliyllobates (in the sense of
Syrrhophus), Plectromantis, Tehnato-
bius, and Zachaenus]. Many of the gen-

12

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

era proposed by previous authors were
combined with the genera he recog-
nized, the first major step toward a syn-
thetic classification. Boulenger recog-
nized no subfamiHes of Cystignathidae.
Noble's ( 1931 ) classification com-
bined two large families ( Bufonidae and
Leptodactylidae ) , and he recognized
five subfamilies for the leptodactylids —
Criniinae, Elosiinae, Heleophryninae,
Leptodactylinae, and Pseudinae. Only
SinintJuUus was erroneouslv associated
( Brachycephalidae ) , although Pseudis
was included in the Leptodactylidae.

Several additional works of limited
geographic coverage are of significance.
In his study of the Brasilian frogs, Mi-
randa-Ribeiro (1926) recognized six
families of leptodactylids — Ceratophry-
didae, Elosiidae (new), Hylodidae, Lep-
todactylidae, Paludicolidae, and Telma-
tobiidae. More significant, however, was
a paper by Parker (1940) in which he
analyzed the variation in the Australo-
Papuan leptodactylids (Criniinae of
Noble, 1931; Davis, 1936) and recog-
nized two subfamilies, the Cycloraninae
(new) and the Myobatrachinae. The
acceptance of these two well-defined
groups has been delayed because too
little has been known of the African and
American genera.

Lutz (1954) proposed two subfami-
lies, Cyclorhamphinae and Eleuthero-
dactylinae, and placed a single genus in
each, although in later papers she ex-
panded the groups. Gallardo (1965)
emended the Cyclorhamphinae to Cy-
cloramphiinae, but both authors were
apparently unaware of Mivart's (1869)
Grypiscina. Reig ( 1960a ) concluded
that Calyptocephalella ( ^Caudiverbera,
Myers, 1962) was not so closely related
as had been previously supposed to the
other telmatobiine frogs and named the
CalyptocephalelHnae.

Gallardo (1965) attempted to ar-
range all of the Neotropical leptodacty-
lids in subfamilies and proposed two
new subfamilies. Unfortunately he did
not assign all of the genera to subfami-

lies, and several of his associations are
highly heterogeneous and unnatural.
The subfamilies he recognized are: Bat-
rachylinae (new, but synonymous with
Mivart's Alsodina and Cacotina), Cera-
tophryinae, Cycloramphiinae ( synony-
mous with Mivart's Grypiscina), Eleu-
therodactylinae, Elosinae, Leptodactyli-
nae, Pseudopaludicolinae (new), and
Telmatobiinae.

Perhaps the most different classifica-
tion of the Leptodactylidae at the fam-
ily-level is that of Kuhn (1965). His
Leptodactylidae included six subfamilies
—Elosiinae, Ceratophryinae, Calypto-
cephalellinae, Telmatobiinae ( erroneous-
ly credited to Vellard), Cystignathinae,
and Criniinae. He recognized the fami-
lies Ceratophrynidae (although he also
listed it as a leptodactylid subfamily one
page earlier) and Heleophrynidae. He
considered Myohatrachus the sole mem-
ber of the Myobatrachinae which he
placed as a subfamily of the Bufonidae.

The most recent compilation of
names in the family (Gorham, 1966) is
especially important because it includes
all but one generic name (valid or not)
proposed for leptodactylid frogs as well
as about 98 percent of the specific names
proposed prior to 1963. Gorham recog-
nized 63 genera and 598 species in the
family; a few genera have since been
named. Many others are considered in-
valid by me; numerous additional species
have been named, and others have been
synonymized since Gorham ceased to
accumulate data ( 1963 ) .

One hundred and sixty-one generic
names have been proposed for the 57
Recent genera recognized here (Table
2). Of the recognized genera, 50 per
cent contain one or two species. The 57
genera considered by me to belong to
the family Leptodactylidae are distrib-
uted over much of the southern land
masses of the world (excepting Antarc-
tica). The seventeen Australo-Papuan
genera are here placed in two subfami-
Hes (those of Parker, 1940), one of
which is further subdivided into two

LYNCH: LEPTODACTYLOID FROGS 13

TABLE 2. Generic names of leptodactylid frogs and the corresponding generic names

used here.

Name proposed Name used herein

Adelottis Ogilby, 1907 Adelotus

Adenomera Steindachner, 1867 Leptodactylus

Abodes Bell, 1843 Eupsophus

Amblyphrymis Cochran & Goin, 1961 Amhlyphrymts

Barycholos Heyer, 1969 Barycholos

Basonitia Miranda-Ribeiro, 1923 Eleiitherodactylus

Batrachophrynus Peters, 1873 Batrachophrynus

Batrachopsis Boulenger, 1882 Lechriodus

Batrachyla Bell, 1843 Batrachijla

Borhorocaetes Cope, 1866 Eupsophus

Borborocoetea Strand, 1928 Eupsophus

Borborocoetes Bell, 1843 Eupsophus

Bubonias Cope, 1874 Edalorhina

Bufonella Girard, 1853 Pseudophryne

Cacotus Giinther, 1868 Eupsophus

Calamobates deWitte, 1930 Crossodactylus

Calyptocephala Nieden, 1923 Caudiverbera

Cahjptocephalella Strand, 1928 Caudiverbera

Calyptocephahis Dum. & Bib., 1841 Caudiverbera

Camariolius Peters, 1863 Crinia

Caudiverbera Laurenti, 1768 Caudiverbera

Ceratophrys Wied, 1824 Ceratophrys

Chacophrys Reig & Limeses, 1963 Ceratophrys

Cheiroleptes Spencer, 1901 Cyclorana

Chehjdobatrachus Giinther, 1859 Myobatrachus

Chiroleptes Giinther 1859 Cyclorana

Cophaeus Cope, 1889 Tehnatobius

Cornufer Tschudi, 1838 Eleutherodactyhis

Craspedoglossa Miiller, 1922 Zachaenus

Craugastor Cope, 1862 EJeutherodactylus

Crinia Tschudi, 1838 Crinia

Crossodactylodes Cochran, 1938 Crossodactylodes

Crossodactylus Dum. & Bib., 1841 Crossodactylus

Cryptotis Giinther, 1863 Adelotus

Ctenocranius MeHn, 1941 Eleutherodactylus

Cyclocephalus Jimenez de la Espada, 1875 Eleutherodactylus

Cycloramphus Tschudi, 1838 Cycloramphus

Cyclorana Steindachner, 1867 Cyclorana

Cijclorhamphus Agassiz, 1846 Cycloramphus

Cystignathus Wagler, 1830 Leptodactylus

Edalorhina Jimenez de la Espada, 1870 Edalorhina

Eleutherodactylus Dum. & Bib., 1841 Eleutherodactylus

Elosia Tschudi, 1838 Hylodes Fitz., 1826

Engystomops Jimenez de la Espada, 1872 Physalaemus

Entomoglossus Peters, 1870 Leptodactylus

Enydrobius Wagler, 1830 Hylodes Fitz., 1826

Epirhexis Cope, 1866 Syrrhophus

Euhyas Fitzinger, 1843 Eleutherodactyhis

14 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

TABLE 2. Generic names of leptodactylid frogs and the corresponding generic names

used here— Continued

Name proposed Name used herein

Euparkerella Griffiths, 1959 EuporkereUa

Ettpemphix Steindachner, 1863 Physakiemus

Eupsophiis Fitzinger, 1843 Eiipsophiis

Eusophus Gope, 1865 Eiipsophus

Geohatrachus Rnthven, 1915 MIGROHYLIDAE

Glauertia Loveridge, 1933 Glauertia

Gnathophysa Gope, 1865 Leptodactyhis

Gomphohaies Rein. & Liitk., 1862 PJujscdaemtts

Grypiscus Gope, 1867 Cycloramphiis

Hammatodactylus Fitzinger, 1843 Eiipsophus

Heleioponis Gray, 1841 Heleioporus

Heleophryne Sclater, 1898 Heleophryne

Helioporus Gray, 1841 Heleioporus

Heliorana Steindachner, 1867 Limnodynastes

Holoaden Miranda-Ribeiro, 1920 Holoaden

Hydrolaetcire Gallardo, 1963 HydroJaetare

Hylactophryne Lynch, 1968 Hylactophryne

Hylodes Fitzinger, 1826 Hylodes

Hylodes Fitzinger, 1843 (Auctorwn) Eleutherodactyhts

Hylopsis Werner, 1896 HYLIDAE (liyki)

Hylorhina Agassiz, 1846 Hylorina

Hylorina Bell, 1843 Hylorina

Hyperolia Gope, 1865 Uperoleia

Hyperoodon Philippi, 1902 Odontophrynus

Hypodictyon Gope, 1885 Eleutherodactyhts

IJioJ)ates Fitzinger, 1867 Physakiemus

Iliodisctis Miranda-Ribeiro, 1920 Cycloramphus

Ischnocnema Rein. & Liitk., 1862 Ischnocnema

Kyarranus Moore, 1958 Kyarranus

Lechriodus Bonlenger, 1882 Lechriodus

Leiuperus Dnm. & Bib., 1841 Pleurodema

Leiyla Keferstein, 1868 Eleutherodactyhts

Lepidohatrachus Budgett, 1899 Lepidobatrachus

Leptodactyhis Fitzinger, 1826 Leptodactyhis

Limnocharis Bell, 1843 Crossodactyhis

Limnodynastes Fitzinger, 1843 Limnodynastes

Limnomedusa Fitzinger, 1843 Limnomedusa

Limnophys Jimenez de la Espada, 1870 Eleutherodactyhts

Liohyla Cope, 1894 Eleutherodactyhts

Lithodytes Fitzinger, 1843 Lithodytes

Litopleura Jimenez de la Espada, 1875 Limnomedusa

Liuperus Gope, 1860 __._ Physalaemus

Liyla Gope, 1869 Eleutherodactyhts

Lystris Gope, 1868 Pleurodema

Macrogenioglottus Garvalho, 1946 Odontophrynus

Malachylodes Gope, 1879 Syrrhophus

Megaelosia Miranda-Ribeiro, 1923 Megaelosia

Metacrinia Parker, 1940 Metacrinia

LYNCH: LEPTODACTYLOID FROGS 15

TABLE 2. Generic names of leptodactylid frogs and the corresponding generic names

used here— Continued

Name proposed Name used herein

Microhatrachijhis Taylor, 1940 Eleutheroclacttjhis

MicropJiryne Peters, 1873 Phijsalaemus

Mitrohjsis Cope, 1889 Cyclorana

Mixophijes Giinther, 1864 Mixophyes

Myohatrachus Schlegel, 1850 .. Myolxit radius

Nattereria Steindachner, 1864 Phy.sakiemus

Neobatrachus Peters, 1863 Neobatrachus

Niceforonia Coin & Cochran, 1963 Niceforonia

Niedcnia Ahl, 1923 Cycloramphus

Noblella Barbour, 1930 Eleutherodactylus

Notaden Giinther, 1873 Notaden

Odontophrynus Rein. & Lijtk., 1862 Odontophrynus

Oocormus Boulenger, 1905 Zachaenus

Opisthodon Steindachner, 1867 Limnodynastes

Oreobates Jimenez de la Espada, 1872 Ischnocnema

Paludicola Wagler, 1830 Phy.salaemus

Paratehnatobhis Lutz & Carvalho, 1958 Paratehnatobius

Parvulus Lutz, 1930 Leptodactylus

Peltocephahis Bibron, 1838 Caudwerbera

Peralaimos Jimenez de la Espada, 1875 Physalaemus

PeriaUa Gray, 1845 Heleioporus

Phanerotis Boulenger, 1890 LecJiriodus

Philocryphus Fletcher, 1894 Heleioporus

Philoria Spencer, 1901 Philoria

Phractops Peters, 1867 Cyclorana

Phrynanodus Ahl, 1933 Eleutherodactylus

Phrynoceros Bibron, in Tschudi, 1838 Ceratophrys

Phrynopus Peters, 1873 Eupsophus

Physalaemus Fitzinger, 1826 Physalaemus

Physalaemus Fitzinger, 1843 Pleurodema

Pithecopsis Giinther, 1859 Cycloramphus

Platyplectron Peters, 1863 Limnodynastes

Platyplectrum Giinther, 1863 Limnodynastes

Plectomantis Cope, 1874 (emendation?) Leptodactylus

Plectromantis Peters, 1862 Leptodactylus

Pleurodema Tschudi, 1838 Pleurodema

Pristimantis Jimenez de la Espada, 1870 Eleutherodactylus

Proceratophrys Miranda-Ribeiro, 1920 Proceratophrys

Pseudobatrachus Peters, 1873 Telmatobius

Pseudohyla Andersson, 1945 Eleutherodactylus

Pseudopaludicola Miranda-Ribeiro, 1926 Pseudopaludicola

Pseudophryne Fitzinger, 1843 Pseudophryne

Pterophryne Giinther, 1867 Crinia

Pternophrynus Liitken, 1863 Crinia

Ranaster MaCleay, 1878 Limnodynastes

Ranidella Girard, 1853 Crinia

Rhinoderma Dum. & Bib., 1841 RHINODERMATIDAE

Scythrophrys new genus Scythrophrys

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

TABLE 2. Generic names of leptodactylid frogs and the corresponding generic names

used here.— Concluded

Name proposed Name used herein

Sihilatrix Fitzinger, 1843 Leptodacfylus

Sminthillus Barbom- & Noble, 1920 SminthiUus

Stombus Gravenhorst, 1825 Ceratophrys

Stombiis (Auctonim) Proceratophrys

Strahomantis Peters, 1863 Eleutherodactyhis

Syrrhaphus Giinther, 1900 Syrrhophiis

Syrrhophus Cope, 1878 Syrrhophiis

Syrrhopus Boulenger, 1888 Syrrhophus

Tarsopterus Rein. & Liitk., 1862 Crossodachjhis

Taudactyhis Straughan & Lee, 1966 Taudactyhis

Tehtrema Miranda-Ribeiro, 1937 Eleutherodactylus

Telmatohius Wiegmann, 1835 Telmatobius

Tehnatobufo Schmidt, 1952 Tehnatobufo

Thoropa Cope, 1865 Thoropa

Tomodactylus Giinther, 1901 Tomodactylus

Trachyphrynus Coin & Cochran, 1963 Eleutherodactylus

Trigonophrys Hallowell, 1856 Ceratophrys

Uperoleia Gray, 1841 Uperoleia

Wagleria Girard, 1853 Limnodynastes

Zachaenus Cope, 1866 Zachaenus

iEophractus Schaeffer, 1949 Caiidiverbera

iGigantobatrachus Casamiquela, 1959 Caiidiverbera

findobatrachus Noble, 1930 ^Indobatrachus

iNeoprocoeki Schaeffer, 1949 iNeoprocoela

fWawelia Casamiquela, 1963 iWawelia

tribes. These two subfamilies might well
represent, as apparently does the South
African Heleopliryninae, independently
derived stocks of megophiyine peloba-
tids. The Neotropical part of the family
is divided into four subfamilies, one of
which is further divided into five tribes.
The classification I propose is as follows:

Subfamily Ceratophryinae (Neotropical).
Included genera: Ceratophrys,
Lepidobatrachus, and the
Miocene Waicelia.

Subfamily Cycloraninae (Australo-
Papuan).
Tribe Cycloranini (included genera:
Cyclorarm, Heleioporus,
Mixophyes, Neobatrachus,
and Notaden).
Tribe Limnodynastini (included
genera: Adelotus, Kyarranus,

Lechriodus, Limnodynastes,
and Philoria).

Subfamily Elosiinae (Neotropical).
Included genera: Crossodactylus,
Hylodes, and Megaelosia.
Subfamily Heleophryninae (South
African).
Included genera: Heleophryne.

Subfamily Leptodactylinae (Neotropical).
Included genera: Barycholos,
Edalorhina, Hydrolaetare,
Leptodactyhis, Limnomedusa,
L ithodytes. Pa rate hnatobitis,
Physalaemus, Pleurodema,
and Pseudopaludicola.
Subfamily Myobatrachinae (Australo-
Papuan ) .
Included genera: Crinia, Glauertia,
the Eocene Indobatrachus
(peninsular India), Metacrinia,

LYNCH: LEPTODACTYLOID FROGS

17

Mijohatrachus, Pseudophnjne,
Taudactylus, and Uperoleia.
Subfamily Telmatobiinae (Neotropical).

Tribe Alsodini (included genera:
Batrachijla, Eupsophus, Hylorina,
and Thoropa).

Tribe Eleutherodactylini (included
genera: Amhlijphnjnus,
Eleutherodactijlus, Euparkerella,
Holoaden, Hylactoplnijne,
Ischnocnema, Nicefownia,
SmintliiUus, Syrrhophus, and
T omodactylus) .

Tribe Grypiscini (included genera:

Tribe Odontophrynini (included
Crossodactylodes, Cycloramphus,
and Zachaenus) .
genera: Odontophrynus and
Proceratophrys) .

Tribe Telmatobiini (included genera:
Batrachophrynus, Caudiverbera,
the Oligocene Neoprocoela,
Telmatobius, and Telmatohujo).

Tribe incertae sedis: Scythrophrys.

The justifications for the above men-
tioned taxonomic rearrangements and
the recognition of as many or as few
( depending on the reader's inclinations )
as 57 Recent genera are the substance of
the following account.

MATERIALS AND METHODS

In the course of this study 304 dried
skeletons, 343 cleared and stained speci-
mens, and innumerable preserved speci-
mens were examined. In addition, four
species were studied by means of stereo-
radiographs. The material used in for-
mulating the generic accounts is listed
in the Appendix.

Illustrated material is identified in
the legends to figures by museum and
catalogue number; the abbreviations cor-
respond to the following institutions and
collections:

AMNH American Museum of

Natural History, New York

AS Private collection of

Albert Schwartz, Miami

BMNH British Museum (Natural

History), London
EHT Private collection of

Edward H. Taylor,

Lawrence
FMNH Field Museum of Natural

History, Chicago
JDL Private skeleton collection

of John D. Lynch
KU University of Kansas

Museum of Natural History
LSUMZ Louisiana State University

Museum of Zoology
MCZ Museum of Comparative

Zoology, Harvard

University
UIMNH University of Illinois

Museum of Natural History
UMMZ University of Michigan

Museum of Zoology
USNM United States National

Museum

Observations on the skeleton, which
form the greater part of this study, were
made by the following techniques: 1)
examination of dried, dermestid-cleaned
skeletons or of macerated, disarticulated
skeletons; the latter were of special im-
portance in the study of smaller species;
2) examination of cleared and stained
specimens prepared by the technique
described by Davis and Gore (1936);
this technique was used for study of rela-
tively small preserved specimens (less
than 40 mm. snout- vent length); 3)
stereo-radiographs prepared in the man-
ner described by Smith and Smith
( 1967 ) for study of genera known only
from type-specimens and/ or very few
specimens; 4) limited dissection used on
rare species and to check relatively su-
perficial characters in large series of
specimens; this technique usually in-
volved careful peeling back of the derm
of the head and then careful separation
or removal of muscle blocks; the tech-
nique was used effectively to study the
arrangement and relationships of the
roofing bones, squamosal architecture,
and to check for the presence or absence
of a frontoparietal fontanelle; and 5)

18

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

microscopic study of serial sections of
the head in the manner described by
Trueb ( 1968 ) ; this technique was used
for comparisons with the works of du
Toit, de VilHers, and Baldauf.

The hyolarynx, thigh and jaw mus-
culature, and the pectoral anatomy were
studied by dissection of preserved ma-
terial or study of cleared and stained
material.

All drawings used in the following
account were prepared by me using a
Wild binocular dissecting microscope
with an attached drawing tube. No ef-
fort was made to correct for distortion or
to idealize or improve the drawings over
the condition of the specimen, inasmuch
as such corrections might confound more
than clarify. The species (skeletal ma-
terial only) used in formulating the gen-
eric accounts are listed alphabetically in
the appendix.

GENERIC CONCEPT

Early in my study, I decided that the
basic unit would be the genus. Because
all categories above and below the
species are arbitrary, some comment on
my generic concept is in order.

There are two approaches towards
genera and their definition. These have
been termed "typological" and "non-
typological" by Inger ( 1958 ) . Likewise,
they might be termed "morphological"
and "adaptive" approaches. There are
advantages and disadvantages to each
approach. The morphological approach
has been, in practice, the more widely
used, but as we learn more about the
biology of the organisms with which we
work, we are finding that most, if not
all, genera have an ecological niche of
sorts. This generic niche is an integral
part of the adaptive approach to genera
as discussed by Inger ( 1958 ) .

In terms of morphology, there is a
tacit philosophy that states that external
characters are used in species and
species-group classification, whereas
osteological characters are those of gen-
era. Although there is some measure of

fact in this philosophy, it cannot be re-
garded as an iron-fast rule as pointed out
by Inger. The single-character classifi-
cation, so prevalent in many groups, is a
prominent liability at the generic level
in leptodactylid classification. It is most
often applied to superficially similar, yet
not closely related, genera such as Cera-
tophnjs, Odontophnjmis, Proceratophnjs,
and Zachaenus. This wide usage of one
or two characters to define any leptodac-
tylid genus has resulted in almost all
leptodactylid genera being very poorly
defined. Most authors have concerned
themselves with the genera and species
of a rather limited geographic area, such
as southeastern Brasil or Mexico, and
have emphasized rather trivial charac-
teristics as being important in generic
diagnoses. Myers (1962) justifiably criti-
cized these workers for their use of such
characters as the presence or absence of
prevomerine teeth or some of the body
glands.

Within the order Salientia, generic
concepts differ from group to group.
Amalops and Rana, both Malaysian
frogs, cannot be satisfactorily separated
as adults, but they are very distinctive in
tadpole morphology and, to a lesser ex-
tent, in ecology. This case is certainly
not equivalent to the case of two other
Malaysian genera, Bufo and Pseudobufo,
which differ in morphology and ecology
of both the adults and tadpoles. In
Inger's ( 1958 ) "adaptive approach to
genera" he used the Malaysian Bufoni-
dae as his example. Within this group,
and for the Bufonidae in general, Inger's
generalizations are true. However, un-
like the Leptodactylidae, small divergent
groups of species recognized as mono-
typic or small genera are rare in the
Bufonidae. Inger (1958) discussed many
of the theoretical and practical difficul-
ties of the generic concept. In some
groups of animals, monotypic genera will
be an absolute necessity even after the
evolutionary trends of that group be-
come apparent and a more synthetic ap-
proach to their classification is possible.

LYNCH: LEPTODACTYLOID FROGS

19

The Leptodactylidae were in the stage
of poorly known evolutionary patterns
until a few years ago. We now have a
reasonably complete picture of the gen-
eral evolutionary trends within the fam-
ily and some, if not all, of the adaptive
patterns. Therefore, one might argue,
there should be little need for, or sense
to, the recognition of many monotypic
genera. Nevertheless, after considerable
study, I still advocate the recognition of
18 monotypic genera. Of course, it is
unknown how many of these will con-
tain as yet undescribed species, but this
difficulty need not concern us.

Mayr, Linsley, and Usinger (1953:
48) required that genera be units that
were ". . . separated from other genera
by a decided gap." Difficulties in this
essentially resemblance-difference system
stem from the definition of a "decided
gap." These authors further suggested
that the size of the gap be in inverse pro-
portion to the size of the genera. Thus
the difference between monotypic gen-
era should be proportionately greater
than the difference between medium-
sized or large genera.

Genera are radiations in some cases
(medium-sized and large genera) but

monotypic genera, in the absence of a
fossil record, cannot be referred to as
radiations. Lcptodactylid genera are
monotypic, small (2-4 species), medium-
sized (5-10 species), large (14-35 spe-
cies), or very large. The last category
includes only one genus, Eleutherodac-
tyliis, with more than 350 species. All
genera, if they are to be realistic, must
be morphologically distinctive as adults.
I would therefore not separate Amalops
from Rana, although it is desirable that
genera likewise be distinctive as larvae.
All of the genera recognized here are
morphologically discrete units charac-
terized by a relatively high degree of
homogeneity, in terms of both morphol-
ogy and ecology. Available data, albeit
incomplete, suggest that each of the
genera exhibits distinctive ecological
parameters. Some of these are the same
among members of unrelated phyletic
lines, but this does not render the gen-
eralization any less important. For exam-
ple, the presence of a foam nest in the
breeding biology of both Pleurodema
and Limnodijnastes does not mean that
they are not generically distinct eco-
logically.

ANALYSIS OF CHARACTERS

Many biologists have gone before me
in the quest of biological and morpho-
logical bases for a classification of the
advanced, though generalized frogs of
the family Leptodactylidae. To these
pioneers I owe special thanks for their
published accounts, which have served
as my sources for much of the data on
behavior and larval morphology. In the
following discussion I have, except where
circumstance dictates that use of a gen-
eric synonym is more explanatory, used
the generic names that I recognize as
valid in the systematic summary.

The higher classification of frogs is a
source of much argument and debate.
Much literature has accumulated on the
subject, in general attacking proposed
classifications. The most recent proposal

and attack respectively are the papers by
GriflBths (1963) and Inger (1967). The
two divergent sets of criteria proposed
by anuran systematists are 1) a system
based primarily upon the nature of the
pectoral girdle; the proponents have
been Cope, Boulenger, and Griffiths;
and 2) a system based primarily upon
the nature of the vertebral centrum and
the thigh musculature; the principle pro-
ponent was Noble. Neither system has a
profound effect on the Leptodactylidae,
although the former would in some cases
align some leptodactylids with ranids.
The latter system would not permit asso-
ciation of genera with paedomorphic
vertebral centra with the families with
which they belong. Furthermore, the
second system does not take into ac-

20

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

count the variation in the disposition of
the distal tendons of the thigh muscles.

Davis (1936), among others, used
the term Bufonoidea to embrace those
groups of frogs which tend to be more
like the bufonids than the ranids. The
recognition of the Bufonoidea and Ra-
noidea is a means of subdividing the
more advanced frogs. This was the ob-
ject of Noble's Procoela and Diplasio-
coela, and Boulenger's Arcifera and
Firmisterna, which approximate the Bu-
fonoidea and Ranoidea as here used.
The Bufonoidea can be defined as fol-
lows: ribs absent in both larvae and
adults; m. sartorhis present; sternum
present (except in Brachycephahis);
scapula not overlain anteriorly by lateral
development of clavicle; pectoral girdle
arciferal with free epicoracoidal horns,
or if firmisternal, epicoracoidal horns
present but fused medially or epicora-
coidal horns absent (Dendrobatidae);
prezonal and/ or post-zonal elements
lacking bony styH; vertebral centra pae-
domorphic or procoelous; larvae type 4
of Orton, 1953, (spiracle sinistral, beak
present, labial papillae and teeth pres-
ent); parahyoid bone absent; arytenoid
cartilage lacking apical cartilages; glan-
dular pharyngeal folds lacking.

The Bufonoidea includes the follow-
ing famihes: Bufonidae (including the
Atelopodidae auctorum), Centrolenidae,
Dendrobatidae, Hylidae, Leptodactyli-
dae, and Pseudidae. The Leptodactyli-
dae are regarded as the basal stock from
which all other families of Bufonoidea
have evolved. The Atelopodid-Bufonid
line differs by the presence in the adult
(or in the young) of an organ of Bidder.
The families differ from one another in
only the mesial fusion of the epicoracoi-
dal halves of the pectoral girdle. Dr.
Roy McDiarmid has recently completed
an osteological study of the family Atelo-
podidae and considers it inseparable
from the Bufonidae — a position with
which I concur on the basis of my own
observations. The Hylidae, Centroleni-
dae, and Pseudidae differ from the other

three families in the possession of an
intercalary phalangeal element. It is
elongate and bony in the pseudids and
short and cartilaginous in the centro-
lenid-hylid complex. The occipital con-
dyles of the pseudids, hylids, and cen-
trolenids are widely separated and
stalked. The centrolenids are the only
bufonoid frogs exhibiting fusion of the
astragalus and calcaneum and uniform
T-shaped terminal phalanges throughout
the family. T-shaped terminal phalanges
occur in several bufonids and leptodac-
tylids, and in the dendrobatids. The
omosternum is lost in all centrolenids
but is retained by at least some members
of all other families. The dendrobatids
are the only bufonids with a truly firmi-
sternal pectoral girdle. They further dif-
fer in the loss of both the prevomers and
the palatines. The Leptodactylidae are
defined as the remaining bufonoid frogs.
All lack an organ of Bidder, intercalary
phalangeal element ( cartilages or bone ) ,
and have an arciferal or modified arcif-
eral pectoral girdle (the epicoracoidal
horns are visible near the posterior edge
of the sternum ) .

Few characteristics are demonstrably
primitive in the Anura. Frogs are un-
questionably amphibians but the lack of
a fossil record makes it difficult or im-
possible to relate them to any fossil
group. Parsons and Williams (1962,
1963) presented convincing evidence
supporting the idea that the three living
amphibian orders are more closely re-
lated to one another than any one is
related to some extinct amphibian order.
Thus characteristics which are shared by
a few anuran groups and most other
Lissamphibia are more likely to be prim-
itive than a character unique to the
order Anura. The presence of free ribs
(in the adult or larva) is therefore a
primitive character in frogs as are the
following: high number of presacral
vertebrae, imbricate neural arches, arcif-
eral pectoral girdles, fully aquatic lar-
vae, and the retention of tail muscles.
These characteristics (here termed first

LYNCH: LEPTODACTYLOID FROGS

21

degree) occur in some or all of the
archaic frog families (Ascaphidae, Dis-
coglossidae, Pipidae, and Rhinophryni-
dae ) as well as in a few of the advanced
families.

Another set of characteristics (sec-
ond degree) are usually not evident (or
difficult to homologize) in non-anurans
and are largely restricted to the archaic
frog families, but occur in some ad-
vanced families (principally the Pelo-
batidae but sometimes in leptodacty-
lids). Because this set of characteristics
is associated (correlated) with the dis-
tribution of first degree characteristics,
the second degree characteristics may be
reasonably asserted to be primitive. In-
guinal amplexus, the presence of a para-
hyoid, and the absence of a discrete m.
sartorius are second degree characteris-
tics.

First and second degree character-
istics are therefore considered primitive
throughout this study. Following the
reasoning of Sporne ( 1954 ) , I also recog-
nize a third set of characteristics as
primitive because the distribution of
these characteristics among frogs is cor-
related with the distribution of the first
and second degree characteristics. Third
degree characteristics are usually ap-
parent in some of the archaic frog fami-
lies and are always apparent in the
Pelobatidae. The following character-
istics are third degree characteristics:

1) occipital condyles large and closely
approximated medially, and cervical
cotyles closely approximated medially;

2) transverse processes of anterior pre-
sacral vertebrae elongate and transverse
processes of posterior presacral vertebrae
shortened; 3) sacral diapophyses broadly
dilated; 4) sacral vertebrae with post-
zygapophyses and coccyx with prezyga-
pophyses; 5) intervertebral discs free;
6) pupil vertical; 7) outer metatarsal
tubercle absent; 8 ) tadpole with median
vent; and 9) tadpole dental formula in-
cluding at least three rows anterior to
and three rows posterior to the beak.

These nine chai'acter states are consid-
ered primitive.

NON-OSTEOLOGICAL
CHARACTERS

A number of morphological and non-
moi-phological character complexes or
traits are considered here. Some of these
have not previously been given serious
consideration, whereas others have been
afforded considerable weight in the con-
struction of classifications.

Early classifications utilized obvious,
ecologically adaptive characters, such as
are used by the layman today. The
"frog" versus "toad" classification is still
prevalent and is (and was) based largely
on the presence or absence, respectively,
of webbing on the digits of the hind
limb. Within the leptodactylids exam-
ples of the "frog" would be Cijcloram-
phtis, Paratelmatohius, or Telnmtohius
in the Neotropics and Cyclorana and
Mixophijes in the Australo-Papuan re-
gion. The "frog-toad" classification also
used pustularity of the skin, the presence
of parotoid glands, and environmental
preference (field or pond). Abundant
evidence has been presented refuting
such a classification. Nonetheless, a num-
ber of behavioral and externally appar-
ent morphological features can be inves-
tigated for possible use in constructing
phylogenies and classifications. Among
such features discussed below are the
following: amplectic position, larval
characteristcis, developmental patterns,
breeding sites and nests, secondary sex-
ual features of the male (vocal appara-
tus, nuptial callosities, excresences,
spines), pupil shape, tongue morphol-
ogy, foot morphology (webbing, meta-
tarsal arrangement, tarsal folds), and
glands on the body.

Recently, serious consideration has
been revived in myological variation and
taxonomic usefulness. Three major areas
of concern here involve the myology of
the jaw, the myology of the thigh, and
the hyoid musculature. Noble (1922),
Parker (1940), and Limeses (1963,

22

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

1964) contributed significantly to the
study of thigh musculature. Noble used
variation in the thigh musculature (es-
pecially distal thigh) to support his
classification of the frogs (1922). Star-
rett (1968) reclassified the Anura on the
basis of the jaw musculature of adults
and tadpoles.

Several trends are evident in these
non-osteological features, some of which
are suggested below to be of phyloge-
netic significance. Most, however, are
extremely labile features appearing in-
dependently in many lines of evolution
in response to similar ecological condi-
tions.

Amplectic Position

It is generally conceded (Tihen,

1965) that "primitive frogs" amplex in a
pelvic position and that "advanced
frogs" amplex in an axillary position.
Few authors (Thomas, 1854, Bruch,
1863) have seriously suggested utilizing
amplectic position as a major taxonomic
criterion. Indeed, most authors (e.g.,
Boulenger, 1912, Jameson, 1955, Grif-
fiths, 1963) stressed the variability of
amplectic position and ruled the char-
acter useless in ascertaining relation-
ships. For example, in breeding con-
gresses of toads (Bufo), males can be
found amplexing females anterior to,
above, or a variable distance behind the
arms; all of these variations are minor,
temporary, or fortuitous deviations from
the axillary amplectic pattern character-
istic of the Bufonidae. Tihen (1965) sug-
gested that the transition from a pelvic
to an axillary amplectic position was an
evolutionary trend but noted that in-
dividual genera may deviate to a highly
specialized degree. The genus most
often cited is Alytes. Alytes, unlike other
discoglossid frogs (Barhomla, Bombina,
and Disco glossus), is usually character-
ized as having cephalic amplexus instead
of the pelvic amplexus seen in the other
genera. The cephalic amplexus of Alytes
is a post-fertilization specialization on
the part of the male for entwining the

egg strings about his hind legs where the
early development ensues. Just prior to
and often during fertilization, Alytes,
Hke other discoglossids, engages in pel-
vic amplexus (Boulenger, 1897).

Since Jameson's (1955) review of
breeding behavior in the Anura, amplec-
tic pattern data have been gathered for
examples of all but one family ( Sooglos-
sidae) and for many more genera of
frogs. We find the following distribu-
tion of pre-fertilization amplectic pat-
terns (pelvic or axillary) in fourteen
families of frogs (not all genera or
species have been observed in all fami-
lies, but based on available observations,
these families can be characterized as
either pelvic or axillary ) :

Pelvic amplexus:

Ascaphidae

Discoglossidae

Pipidae

Rhinophrynidae

Pelobatidae

Leptodactylidae ( part )
Axillary amplexus:

Bufonidae

Centrolenidae

Hylidae

Pseudidae

Leptodactylidae (part)

Microhylidae

Ranidae

Hyperoliidae

The evolutionaiy trend suggested by
Tihen (1965) — pelvic to axillary amplex-
us— is reasonable. Fewer sperm are
probably required when the male and
female gametes are released in close
proximity. Some pelvic amplexing frogs
have specialized their breeding behavior
to compensate for their less efficient am-
plectic type. Ascaphus has an intromit-
tent organ, the "tail"; pipids have a "flip-
over" fertilization cycle (Rabb and Rabb,
1960, 1963a, 1963b). Most of the slight
variation in clasping position probably
reflects modifications brought about by
selection for greater fertilization econ-
omy. It is inconceivable that there is any

LYNCH: LEPTODACTYLOID FROGS

23

selective advantage in shifting from axil-
lary to pelvic amplexus. Were there any
advantage, it would probably be evident
by the occurrence of some bufonids, hy-
lids, or ranids exhibiting pelvic am-
plexus.

As is evident by examination of the
above table, only the Leptodactylidae
exhibit both axillary and pelvic amplexus
(Fig. 2). All other families are uniform-
ly axillary or pelvic amplexing frogs.

Figure 2. Amplexing pairs of Pseiidophnjne
australis (top) and Phtjsaloemtis pustulostis
(bottom); the latter also illustrates a foam nest.
Photo of Pseudophryne courtesy of John A.
Moore; that of Physalaemus courtesy of William
E. Duellman.

Amplexus has not been observed (or
reported) in the following leptodacty-
lid genera: Amblyphrynus, Banjcholos,
Crossodactylodes, Edalorhina, Heleo-
phryne, Hydrolaetare, Ischnocnema,
LWwdytes, Megaelosia, Myohatrachus,
Niceforonia, Notaden, Scythrophrys,
Taudactylus, Telmatobufo, and Thoropa.

The following genera have been ob-

served to engage in pelvic amplexus:
Adelotus, Batrachyla, Crinia, Cyclorana,
Glauertia, Heleioporus, Kyarranus, Lech-
riodus, Limnodynastes, Metacrinia, Mix-
ophyes, Neobatrachus, Philoria, Pseudo-
phryne, and Uperoleia. The only varia-
tions known within these genera are the
report of Fletcher (1889) that Mixo-
phyes engages in axillary amplexus (M.
J. Littlejohn and A. A. Martin observed
only pelvic amplexus in this genus), and
a photograph of an axillary amplexing
pair of Batrachyla taeniata sent to me by
Jose Cei (Barrio, 1967a, observed only
pelvic amplexus in the other two species
of the genus ).^

The following genera have been ob-
served to engage in only axillary am-
plexus: Batrachophryniis, Catidiverbera,
Ceratophrys, Crossodactyhis, Cycloram-
phus, Eletitherodactyhis, Euparkerella,
Eupsophus, Holoaden, Hylactophryne,
Hylodes, Hylorina, Lepidobatrachus,
Leptodactyhis, Limnomediisa, Odonto-
phrynus, Paratehnatobms, Physalaemus,
Pleurodema, Proceratoplirys, Pseudo-
paludicola, Syrrhophus, Tehnatobius,
Tomodactylus, and Zachaenus.

Examination of these lists immedi-
ately points out that no Australo-Papuan
genus exhibits axillary amplexus (if we
dismiss Fletcher's, 1889, report) and
only one Neotropical genus exhibits pel-
vic amplexus. The Australo-Papuan lep-
todactylids are judged to be closely re-
lated to megophryine pelobatids, based
on morphology, and the primitive am-
plectic pattern corresponds with this
close relationship. In view of the wide
distribution of pelvic amplexus in the
Australo-Papuan leptodactylids and the
probable irreversibility of this evolution-
ary trend, I consider the Australo-Papu-
an leptodactylids to have primitively
exhibited pelvic amplexus.

All but one (Batrachyla) of the 26

' Dr. Ian Straughan (in litt.) suggested that all
cycloranines employ axillary amplexus. He has
observed axillary amplexus in several taxa. His
observations are contrary to observations pub-
lished by Littlejohn (1963), Martin (1968),
and Moore (1961).

24

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Neotropical leptodact)4id genera for
which pre-fertilization amplectic posi-
tions are known exhibit axillary am-
plexus. The large number of genera
with axillary amplexus and their diver-
sity suggests that the shift to axillary
amplexus was made early in the evolu-
tion of the Neotropical leptodactylid
stock, because each time a shift occurs
independently, it is less likely that the
frogs of a given amplectic type are re-
lated. For example, if of 20 genera (all
judged closely related), one exhibited
pelvic amplexus and the other 19 axillary
amplexus, it is more likely that the shift
occurred only once (prior to the evolu-
tion of the 19 axillary amplexing genera)
and not that it may have occured inde-
pendently two, 10, or 19 times. The oc-
currence of a single pelvic amplexing
genus of Neotropical leptodactylids is
witness either to the primitive stock ex-
hibiting pelvic amplexus, or the invasion
of two stocks, one having pelvic am-
plexus and the other axillaiy amplexus.
It is not unlikely that isolated phy-
letic lines could have achieved the axil-
lary grade independently. Such units as
Heleophnjne, Rhinoderjua or the soo-
glossid ranoids may have independently
evolved axillary amplexus. Data are not
available for Heleophryne or the sooglos-
sid ranoids, but each group exhibits
many primitive characteristics and could
have either pelvic or axillary amplexus.

Breeding Behavior and Development

Many authors have reported on the
direct development of frogs of the genus
Eleutherodactylus. Most other frogs
typically lay the eggs in water and have
a free-swimming larval stage. Of the
"typical frogs" some construct a foam or
froth nest in which the earliest develop-
ment of the tadpole takes place. Lepto-
dactylid frogs can be divided into three
classes: those laying the eggs directly in
water, those depositing the eggs in a
foam nest, and those with direct devel-
opment. The first group can be sub-
divided into those laying their eggs in

ponds and slow streams or rivers (most
species) and those laying their eggs in
close association with torrential moun-
tain streams (e.g., Heleophryne and
Thoropa). Within the group that builds
foam nests some species deposit the eggs
in a nest built in moist terrestrial situa-
tions which are later flooded (e.g.,
Heleioporus and members of the Lepto-
dactyhis fuscus group); others deposit
the eggs in a foam nest ( Fig. 2 ) floating
on water (e.g., most limnodynastine
Cycloraninae, many Leptodactylus, Phy-
salaemus, and Vleurodema) .

No data on breeding biology or de-
velopment are known for the following
genera: Amhlyphrynus, Glauertia, Hy-
drolaetare, Ischnocnema, Limnomedusa,
Lithodytes, Notaden, Paratehnatobius,
Taiidactylus, and Tehimtobufo. Only in-
complete data are available for several
other genera.

Batrachophrymis, Caudiverbera, Cer-
atoplirys, Crinia (part), Crossed actyhis,
Cyclorana, Eupsophus, Hylodes, Hylo-
rina, Lepidobatrachus, Megaeloski, Mix-
ophyes, Neobatraclnis, Odontophrymis,
Proceratophrys, Pseudopaludicola, Pseti-
dopliryne (part), Tehnatobius, and Up-
eroleia lay their eggs in water without
building a foam nest. Glauertia and
Notaden probably also lay their eggs in
water (Main and Storr, 1966; Slater and
Main, 1963). Batrachyla, Tl^oropa, and
some Crinia and Pseudophryne lay their
eggs in moist situations and, upon hatch-
ing, the tadpoles enter water. The same
pattern is inferred for Heleophryne, but
field observations are not available.

Adelotus, Lechriodus, Leptodactylus
(fuscus, ocellatus, and pentadactylus
groups), Limnodynastes, Physalaemus
(including Engystomops and Eupem-
phix), and Pleurodema lay their eggs in
foam nests floating on the surface of
water. Heleioporus, Kyarranus, mem-
bers of the Leptodactylus marmoratus
group, and Philoria lay their eggs in
foam nests in moist, terrestrial situations;
development is direct in Kyarranus and
the L. marmoratus group and may be in

LYNCH: LEPTODACTYLOID FROGS

25

PJuloria but is not in Heleioporiis. In
Heleioporus, as in the Leptodactylus
fuscus group, the eggs are laid in a foam
nest in a terrestrial situation, but prior to
hatching the nest is flooded and the tad-
poles emerge in an aquatic environment.
In the foam-nest builders, the females of
Adelotus, Lechrioclus, Limnodyna.stes, as
well as Kyarranus and Thiloria (Moore,
1958, 1961; Littlejohn, 1963), have broad,
fleshy flanges on the inner fingers (cf.
Fig. 48). Flanges are lacking in the other
foam-nest builders. In those genera with
flanged fingers that lay the eggs in a
nest floating on water, the female churns
a mucous in water with her hands and
traps air bubbles in the froth. In Lepto-
dactylus, Physalaemus, and Pleurodema
the male kicks in a mucous and water
mixture to trap air bubbles. The mode
of building the nest in Heleioporus is
unknown; in Kyarranus and Pluloria,
nest-building is accomplished by the
female using the hands but not in water.
Members of the Leptodactylus marmora-
tus group lay the eggs in a froth nest in
a cavity (usually a hole dug in moist
soil), and the embryos develop without
a free-swimming tadpole stage (Heyer,
1969a).

Cycloramplms and Zachaenus de-
posit their large eggs amidst wet leaves
(terrestrial situation), and development
proceeds within the egg. In contrast to
eleutherodactyline frogs, the egg hatches
as an advanced tadpole, and the tadpole
lives in the wet leaves for a short period
and then metamorphoses (Lutz, 1944).
Crossodactylodes lay their large eggs in
the axillae of terrestrial bromeliads. The
tadpoles complete development in the
moist axillae.

Direct development (without a foam
nest) occurs in Eleutherodactylus, Eu-
parkereUa, Holoaden, Hylactophryne,
Sminthillus, Syrrhophus, and Tomodac-
tylus, and probably also occurs in Nice-
foronia (Goin and Cochran, 1963) and
Metacrinia and Myohatrachus (Parker,
1940). In this group development pro-
ceeds within the egg membrane, and on

hatching, a miniature froglet emerges
(Goin, 1946, Griffiths, 1959, Lynn, 1942,
Lynn and Lutz, 1946a, 1946b, Lutz,
1958, Noble, 1926a, and Valett and Jame-
son, 1961). The oviducal eggs in every
case are large and few in number.

The habit of building a foam nest
probably has developed independently
on different continents. In those Aus-
tralo-Papuan genera which build foam
nests and the females of which have
spatulate fingers, the palatal osteology
suggests a close relationship. The other
Australian genus with a foam nest is
closely allied to genera which do not
build foam nests. The Neotropical foam-
nest builders all possess an osseous ster-
nal style and are unquestionably related;
these genera also are distinctive in the
mode of nest building in that the male
kicks air into the nest (Bokermann,
1962).

Direct development has indepen-
dently evolved in limnodynastine Cyclo-
raninae (Kyarranus and Philoria), the
Myobatrachinae (Metacrinia and Myo-
hatrachus), the Leptodactylinae (Lepto-
dactylus marmoratus group), and Telma-
tobiinae (the genera here placed in the
tribe Eleutherodactylini).

Tadpole Morphology

The use of tadpole morphology in
generic diagnoses has not gained wide
acceptance chiefly because of the gen-
eral lack of material of most species in
collections. When tadpoles have been
available, they have often provided addi-
tional evidence relative to the advisabil-
ity of generic recognition (Inger, 1954,
1966). Examination of available speci-
mens and utilization of reliable reports
in the literature revealed some intriguing
variation in the tadpoles of leptodacty-
lids, especially in fight of Inger 's (1967)
recent comment that some tadpole char-
acters are invariate below the family
level (e.g., the position of the anal
opening ) .

The following characters were ob-
served for each available genus : 1 ) posi-

26

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

sion of the anal opening; 2) tooth rows;
and 3) distribution of labial papillae. In
addition, some other characters were
used when deemed of sufficient depar-
ture from the generalized body form to
be characteristic at the generic level —
for example, body form in the highly
stream-adapted tadpoles of TJwwpa
(Fig. 3).

Figure 3. Tadpoles of (A) Thoropa petropoli-
tana (KU 112394, X 4.5), (B) Crossodacttjliis
dispar (after Bokemiann, 1963, X 1-8), and
(C) Hylodes aspera (KU 112387, X 1.8). Mouth-
parts of (D) a myobatrachine ( Crinia victoriana,
after Martin, 1965 ) and ( E ) a cycloranine
( Neobatrachiis pictus, after Martin, 1965).

Among the Neotropical genera, data
are not available for Amhhjphnjnus,
Barijcholos, Hydrolaetare, Ischnocnema,
Limnomedusa, Lithodytes, Scythrophnjs,
or Telmatohufor Crossodactylodes has
bromeliad dwelling tadpoles (W. C. A.
Bokermann, in litt.) and Edalorhirm has
aquatic tadpoles ( R. Etheridge, in litt. ) ;
I have no other data for these two gen-
era. Tadpole data are not available for
the following genera, all of which have
direct development: Cycloramphus,
EUutherodactylus, Euparkerella, Holoa-
den, Hylactophryne, Niceforpnia, Para-
telmatobius, SminthiUus, SyrrJwphus,
Tomodactylus, and Zachaenus. Data are
available (at least in part) for each of

- Tadpoles are now known for Telmatobufo.
The mouth is ventral, the tooth row formula
2/3, and the papillae form a complete series
around the mouth. The tadpoles are swift-
stream adapted.

the 18 remaining Neotropical genera
that I recognize.

Tadpoles of both ceratophryine gen-
era are known; the larvae of both genera
have a median vent, highly modified
tooth row formula, and relatively few,
unifomily spaced papillae across the an-
terior edge of the mouth. Ceratophrys
has 6/7 tooth rows and large horny beaks
whereas Lepidobatraclius lacks larval
teeth and has large but weakly devel-
oped beaks.

Data are available for four of the
nine leptodactyline genera. Leptodac-
tylus and Pleurodema have median
vents, and Physalaemus and Pseudopalu-
dicola have dextral vents. The tooth row
formula for all four genera is 2/3, and
the labial papillae are interrupted an-
teriorly.

Data are available for all three gen-
era of the Elosiinae. Crossodactylus has
a median vent, and Hylvdes and Megae-
losia have dextral vents. All thi-ee have
2/3 tooth rows and an anterior inter-
ruption in the labial papillae.

Tadpole morphology is not known
for the majority of the Telmatobiinae;
many of these genera exhibit direct de-
velopment. Two genera depart from the
typical bufonoid 2/3 tooth row formula:
Caiidiverbera has 3/3 tooth rows, and
Hylorina has 2/2. Only one genus dif-
fers in the pattern of the labial papillae;
Batrachyla has a complete row of labial
papillae across the anterior edge of the
mouth. All other genera for which data
are available have an interruption an-
teriorly. None has a posterior inter-
ruption.

Batrachyla, Eupsophus, Hylorina,
Proceratophrys, and Thoropa have me-
dian vents, whereas dextral vents occur
in Batrachophrynus, Caudiverbera,
Odontophrymis, and Telmatobius.

Cycloramphus and Zachaenus have
an alDbreviated larval life but do not
exhibit the direct developmental pattern
such as is seen in Eleutherodactylus.
During development in the former two
genera, the vent is medial and the tooth

LYNCH: LEPTODACTYLOID FROGS

27

row formula is 1/1. The upper row of
teeth is interrupted medially as in many
other leptodactylids.

Tadpoles of the Australo-Papuan
genera are relatively well known
through the efforts of Moore ( 1961 ) and
Parker (1940). Data are not available
for the myobatrachine genera GImiertia,
Metocrinia, Myobatrachus, or Taudacty-
his. At least two of these (Metacrinia
and Myobatrachus) probably lack a free-
living tadpole (Parker, 1940).

Two of the cycloranine genera ex-
hibit modified developmental patterns —
Kyanamis and Vhiloria. Both build a
foam nest and lay large eggs in a ter-
restrial situation (Moore, 1958, 1961;
Littlejohn, 1963). These genera have
median anal openings as tadpoles. Re-
ports in the literature concerning the
other cycloranines are contradictory.
Martin (1965), in the most recent studies
on these frogs, reported median vents in
Neobatrachiis pictus and three species of
Limnodynastes (dorsalis, peronii, and
ta.smmiien.sis). Moore (1961) also re-
ported median vents in L. dorsalis and
L. peronii but characterized Neobatra-
chiis pictus as having a dextral vent.
Parker ( 1940 ) , in contrast to reports by
Martin and Moore, reported a dextral
vent in a Limnodynastes (L. spenceri).
Parker (1940) and Moore (1961) pro-
vided descriptions of the larvae of
AdeJotus brevis, Cyclorana phtycepha-
lus, Heleioporus eyeri, Lechriodus fietch-
eri, and Mixophyes fa.sciolatus, all of
which were characterized by having a
dextral vent. Lee (1966) reported in-
traspecific variation ( median or dextral )
in one species of Heleioporus; most
species of the genus have dextral vents.
The cycloranines have three rows of
teeth behind the beak. Adelotus, Cy-
clorana, and Lechriodus have 2/3 tooth
rows insofar as they are known but
Heleioporus, Mixophyes, Limnodynastes,
Neobatrachiis, and Notaden have 3/3 to
6/3 tooth rows. The labial papillae are
not interrupted anteriorly in Mixophyes

and are only narrowly interrupted in
Adelotus, whereas there is a broad in-
terruption in all other genera and species

Data are available for three myoba-
known.

trachines — Crinia, Pseudophryne, and
Uperoleia. These tliree genera are dis-
tinctive among leptodactylids ( and most
bufonoids) in having the labial papillae
interrupted both anteriorly and posteri-
orly (Fig. 3). Crinia and Pseudophryne
have 2/3 tooth rows, whereas Uperoleia
has 1/3.

Intrafamilial varition in the position
of the vent was denied by Inger ( 1967 ) ;
however, the above data clearly indicate
that variation occurs within subfamilies
and even within genera and species.
Starrett (1967) reported that some spe-
cies of Bufo have a dextral vent, whereas
most have a median vent, as do members
of other genera in the family (Ansonia,
Atelopus, and Melanophryniscus). Den-
drophryniscus brevipollicatus also has a
dextral vent; data are not known for the
other three species of that genus, some
of which are only tentatively retained in
the genus.

The tadpole of the ancestral lepto-
dactylid probably exhibited the follow-
ing characteristics: 1) median vent; 2)
more than 2/3 tooth rows; 3) labial pa-
pillae not or only narrowly, interrupted
anteriorly, but not posteriorly; and 4)
relatively deep body with high fins on
the tail (deduced because the primitive
leptodactylid frog was a pond breeder ) .
Based on my analysis of the relationships
of the genera, I consider the dextral vent
to have appeared no fewer than five
times in the course of the evolution of
the American genera and no less than
twice in the evolution of the Australo-
Papuan genera.

The tadpole of Heleophryne is very
specialized. The beak has been lost as in
Lepidobatrachus, the vent is median,
and the tooth rows vary from 4/11 to
4/17. The labial papillae are complete
anteriorly.

28

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Secondary Sex Characters

The development or absence of some
of the secondary sex characteristics has
been used in defining genera. Liu ( 1936 )
and Noble ( 1931 ) published reviews of
the secondary sex characters in frogs,
and Parker ( 1940 ) made some interest-
ing speculations on the variation in nup-
tial excrescences. Of the numerous sec-
ondary sex characteristics that could be
discussed here, the vocal pouches or sacs
and the development of nuptial ex-
crescences of the male are the most im-
portant.

The nuptial excrescences on the
thumb (and sometimes second finger)
vary from roughened, sandpaper-like
patches, through small, numerous, close-
set spines, to a few large, horny spines.
Large heavy spines are uncommon in
leptodactylids — Heleioporus and some
species groups of Leptodactyhis (mela-
nonotus, ocellattis, and pentadactylus)
have thumb spines. I have observed
relatively little intrageneric variation in
the presence or absence of nuptial asper-
ities in leptodactylids, although this
variation does occur. Parker ( 1940 )
pointed out that apparently those frogs
which clasp on land and not in water
lack nuptial excrescences, whereas those
that clasp in water have them. This
hypothesis was put forth following a
study of the Australo-Papuan leptodac-
tylids, but he did not say whether the
hypothesis was formulated only on the
basis of Australo-Papuan leptodactylids,
or whether it was influenced by his wide
experience with frogs from all over the
world. Nonetheless, those genera of
Neotropical leptodactylids lacking nup-
tial excrescences all clasp on land; most
also exhibit direct development. Kyar-
ranus, an Australian genus exhibiting di-
rect development, does not clasp in
water and has nuptial cxcrespences
(Moore, 1958, 1961). A Neotropical
genus, Batrachyla, has nuptial excres-
cences but clasps on land ( Barrio, 1967a;
Cei, 1962a ) . There seems to be evidence
for and against Parker's suggestion. In

some cases the secondary sex characters
seem to have significance in suggesting
relationships ( e.g., within the Eleuthero-
dactylini), but in others the characters
reflect breeding behavior. This is espe-
cially apparent in Leptodactyhis, in
which the species of the two groups that
clasp on land (fiiscus and marmoratiis)
lack spines, and the three that clasp in
water (melanonotus, oceUatus, and pen-
todoctyhis) have spines. The species of
the melanonotus, ocellattis, and penta-
dactylus groups are closely related. The
species of the ftiscus group are more
closely related to the groups with thumb
spines than to the other group (mar-
moratiis) lacking spines (Heyer, 1969a).
This example illustrates the inadvisabil-
ity of applying extra weight to the taxo-
nomic value of variations in the sec-
ondary sex characteristics when con-
structing frog phylogenies at or above
the generic level.

Secondary sex characters have been
used to characterize some genera. For
example, Heleioporus usually has one to
several large spines on the thumb of the
male ( one species lacks spines ) , whereas
Neohatraclms always has a diffuse pad
on the thumb of the male. In external
appearance the frogs of these two gen-
era are otherwise difficult to separate.
Nuptial excrescences are lacking in male
Niceforonia but present in male Eupso-
phus. These genera had previously been
confused and can be separated osteo-
logically, but only with difficulty ex-
ternally.

Males of the ocellatiis and pentadac-
tylus groups of Leptodactyhis have
greatly enlarged muscles in the arm;
this enlargement is osteologically sup-
ported by the development of large
ffanges on the humerus (Fig. 41).

A curious secondary sex characteris-
tic is exhibited by males in some species
of Eiipsophus, Leptodactyhis, and Tel-
matohius, in which clusters of horny
spines develop on the chest. These
spines are seasonal in appearance at

LYNCH: LEPTODACTYLOID FROGS

29

least in Eiipsophtis and may be in all of
the species which have them.

Few secondaiv sex characteristics are
developed in female frogs. The females
of those frogs here placed in the Limno-
dynastini develop spatulate fringes on
the inner digits of the hand during the
breeding season (Fig. 48). The fringe
enables the female more easilv to make
the foam nest in which the eggs are
deposited. Of the five genera in this
tribe, males of all species have nuptial
pads. Two of the genera (Kyarranus
and Pliiloria) exhibit direct development
and do not lay their eggs in water.

Male leptodactylid frogs exhibit a
wide range of variation in the condition
and development of vocal pouches. Sev-
eral genera lack vocal sacs (e.g., Megae-
Josia), and this character varies intra-
generically (e.g., Eleiitlwrodactijhis).
Most leptodactylids have an internal or
external subgular vocal sac, but the sac
is paired and ventrolateral in the Lep-
todactylus fuscus group. In Hylodes the
vocal sacs are membranous and lateral.

Pupil Shape

A widely used "generic" character in
many anuran families is the nature of
the pupil — horizontally or vertically el-
liptical (Fig. 4). I am aware of no
author who has afforded the shape of
the pupil more than generic value or
who has suggested that it is anything
more than a handy "key character."

Several pupil shapes have been de-
scribed for the leptodactylid genera
(horizontal, round, horizontal with ven-
tral angle, rhomboidal, and vertical). In
most cases the specimens were pre-
served. Apparently pupil shape is al-
tered by death, inasmuch as species re-
ported as having a horizontal pupil with
a ventral angle by Parker ( 1940 ) have
either horizontal or vertical pupils in
life (Moore, 1961, Lee, 1966). Uperoleia
has a rhomboidal pupil when preserved
but the pupil is a vertical slit in life
(Main and Storr, 1966, Moore, 1961).
The pupil shapes in leptodactylids are

variations of two types — vertical or hori-
zontal ellipse. The only leptodactylid
genera with vertical pupils are Caiidi-
verbera, Heleioporus, Heleophryne, Hy-
drolaetare, Hylorina, Lepidobatrachus,
Limnomedusa, Mixophyes, Neobatra-
chus, and Tehnatobufo.

All primitive families (Ascaphidae,
Discoglossidae, Pipidae, Rhinophryni-
dae, and Pelobatidae) have vertical pu-
pils, with the possible exception of Neso-
bia, a rare, megophryine pelobatid. In
contrast, vertical pupils are uncommon
in higher frog families. No genus of
bufonids, centrolenids, dendrobatids,
pseudids, rhinodermatids, or sooglossids
has vertical pupils. Among the Hylidae,
vertical pupils occur in only the four
genera of the Phyllomedusinae (Aga-
lychnis, Nyctimistes, PacJiymedusa, and
PJiylloniedusa), although several other

Figure 4. Pupil shape in (A) Heleophryne
natalensis, KU 105923, X 2; (B) Neobatrachiis
pictus, KU 69276, X 2; (C) Hylodes aspera,
KU 92871, X 1.5; and (D) Eleuthewdactijhis
silvicola, LSUMZ 7557, X 2.5.

30

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

genera have been erroneously reported
to have vertical or rhomboidal pupils.

The only microhylid genus definitely
known to have vertical pupils is Dijsco-
phus, although CtenopJiryne may have
vertical pupils (Parker, 1934).

Other than the Leptodactylidae, the
Ranidae (sensu lato) are the only ad-
\'anced family ha\'ing several genera
with vertical pupils. Ranids are diverse
in terms of generic radiation except in
the Americas; vertical pupils occur only
in the African genera, whereas none of
the several genera of ranids in the Indo-
Australian Archipelago, Malaysia, and
China has vertical pupils. The only
ranid genera with vertical pupils belong
to the group recognized as the Hypero-
liidae by Laurent (1951), but several of
the genera in that family group have
horizontal pupils.

Hands and Feet

The degree of webbing, digital pad
expansion or development, variations in
the tarsal folds and tubercles, and sizes
of the subarticular, supernumerary, and
metatarsal tubercles have been used as
generic characters in the Leptodactyli-
dae. Within the large genera most of
these characters exhibit a continuum.
However, they have been used, and
probably will continue to be used, to
distinguish the smaller genera from the
larger ones (Eleutlwroductylus and Lep-
todactylus).

Leptodactylids are generally thought
of as free-toed frogs, whereas other
groups (e.g., Hyhdae and Ranidae) are
characteristically web-footed. The hands
of all leptodactylids lack webbing; some
authors have reported basal webbing of
the hands, but this apparently always
has been based on a poorly preserved
and/ or desiccated specimen. Lateral
fringes are present on the digits of some
species, but these cannot be interpreted
as webbing. Batrachoplirynus, Catidi-
verbera, Ceratophrys, Cyclorana, Glaii-
ertia, Hydrolaetare, Limnomedusa, Mix-
ophyes, Neobatrachus, Notaden, Para-

tehnafobius, Proceratophrys, Telmato-
hins, and Tehnatobufo have webbing
between the toes; the webbing is not
extensive in Ceratophrys, Limnomedusa,
or Proceratophrys. Several other genera
have basal webbing. Especially among
the Australo-Papuan leptodactylids, the
distinction between "free toes" and
"webbed toes" is tenuous. The full range
of webbing conditions occurs in Cyclo-
ramphus, Eletitherodactylus, and Limno-
dynastes.

Expanded pads occur on the tips of
the digits of many leptodactylids and are
apparently correlated with arboreal
habits. Frogs of the genus Eleuthero-
dactyhis exhibit a full range of variation
from terrestrial species with no pads to
fully arboreal species with large, emar-
ginate pads. The finger pads are usually
larger than those on the toes but in some
species the reverse is true. I previously
pointed out the value and significance
of the presence or absence of a terminal
ti-ansverse groove on the digital pads
(Lynch, 1968a). The grooves are best
developed on the outer digital pads and
usually better developed on the toes
than on the fingers.

Boulenger (1882) used the separa-
tion or union of the outer metatarsals as
a generic character. All leptodactylids
are closer to the united condition than to
the separated condition exemplified by
Xenopns. Separation of the metatarsals
is best developed in, but not exclusive
to, those species with webbed feet. I
consider the character to be of but
limited importance because I find no
myological basis for it in leptodactylids
and find a continuum of character states
relative to the separation. The separa-
tion is complete in Mixophyes, less so in
the Cycloramphus fulginosus group (in
which the fourth and fifth metatarsals
are separated by webbing for about half
of their length), and united (Fig. 5) in
the web-less species of Cycloramphus
(for example, C. eJeutherodactyhis) .

The tarsus is bedecked by inner
and/or outer tubercles or folds in many

LYNCH: LEPTODACTYLOID FROGS

31

leptodactylids. Usually the folds are
small, often ridge-like; these folds are
variable in genera and species groups
and the development of the folds is
sometimes sexually variable (EleutJiero-
dactylus rugulosiis). However, the elo-
siine leptodactylids have large, Hap-like
tarsal folds (Fig. 5) in both sexes.

The subarticular tubercles vary in
size and shape. Usually they are
rounded, light colored, and moderate
sized, but in several species they are
elongate, conical, or absent. I have not
observed bifid subarticular tubercles in
leptodactylids. Supernumerary plantar
tubercles occur in relatively few lepto-
dactylids and are useful in species-group
and generic classification. These tuber-
cles are arranged in rows along the meta-
tarsals in some species, but in most they

Figure 5. Plantar views of feet of leptodactylid
frogs. (A) Hylodes aspera, KU 92871; (B)
Cijcloramphus eleutherodactylus, KU 92781;
and (C) C. dubitis, KU 74196, illustrating lat-
eral fringes on the toes, tarsal fold, and presence
and absence of digital webbing. All X 1-5.

are irregularly scattered over the sole.
The relative sizes of the metatarsal
tubercles are of use in defining small
genera and species groups in moderate
sized and large genera. In a few semi-
fossorial genera, the inner metatarsal
tubercle is enlarged and spade-like
(compressed and keeled).

Figure 6. Plantar views of feet of (A) Heleo-
phrtjiie natalensis ( KU 105923 ) and ( B ) Eleu-
therodactylus chloronotus ( KU 117519) illus-
trating presence and absence of outer metatarsal
tubercles. The foot of Heleophryne also illus-
trates completely separated outer metatarsals
whereas those of Eleutherodactylus are united.

X2.

Primitive anuran families and primi-
tive genera of the Leptodactylidae lack
an outer metatarsal tubercle (Fig. 6).
Many hylids and some centrolenids and
ranids also lack an outer metatarsal
tubercle. Within the leptodactylids I re-
gard the development of an outer meta-
tarsal tubercle as a derived state. Most
genera with vertical pupils lack an outer
metatarsal tubercle; Uperoleia is an ex-
ception. The development of the tuber-
cle may be a manifestation of adaptation
to terrestrial modes of life, but the gen-
eralization runs contrary to the absence
of the tubercle in the fossorial pelobatid
genera and Rhinophrynus. The presence
or absence of an outer metatarsal tu-

32

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

bercle is \ariable in some leptodactylid
genera (Crinia, Cydorana, and Limno-
dijna.stes). The following genera lack an
outer metatarsal tubercle: Caudwerhera,
Crinia (except georgiana, glmiertia, tas-
manienms, and the signifera superspe-
cies), Cydorana (except inermis), He-
Jeioporus, Hdeophryne, Kyarranus, Lep-
idohatradnis, Limnodynastes (except
salmini), Mixophyes, Neobatradnis, No-
taden, Fhihria, Taudactylus, and Telma-
iohiifo.

Body Glands

The presence or absence of a variety
of glands on the body has been widely
used in the generic classification of lep-
todactylids. Those glands in question are
1) paratoid, 2) lumbar, 3) inguinal, 4)
tibial, and 5) flank glands. Some gen-
era, such as Holoaden, are heavily glan-
dular, whereas others lack all of the

above-mentioned glands. The presence
of parotoid glands is usually readily
demonstrated by external examination.
The same is true of the inguinal glands
of some Fhysalaemus and the lumbar
glands of Pleurodema and Tomodacty-
his. Inguinal glands usually are not
clearly discernible from external exam-
ination, and their presence can be veri-
fied only by examining the inner surface
of the skin. The inguinal glands vary
from well defined (as in Cydoramphus)
to loosely organized (many Eleuthero-
dactylus). Schematic drawings (Fig. 7)
of Cydoramphus, Odontophrynus, Pliil-
oria, Pleurodema, Telmatobufo, and Up-
eroleia show the most extensive develop-
ment of each kind of gland.

Parker (1940) used the number of
ducts of the intermaxillary glands to
separate the Australo-Papuan subfami-
lies Cycloraninae and Myobatrachinae.

Figure 7. Body glands in leptodactylid frogs. (A) Telmatobufo hullocki, FMNH 23842, X .5, paro-
toid glands; (B) Pleurodema cinera, KU 80829, X L lumbar gland; (C) Philoria frosti, KU 110344,
X 1, parotoid gland; (D) Odontophrynus cultripes, KU 92974, X -7, parotoid and tibial glands; (E)
Uperoleia rugosa, KU 93589, X L2, parotoid and flank glands; and (F) Cycloramphus asper, KU

84715, X 1, inguinal gland.

LYNCH: LEPTODACTYLOID FROGS

33

I have not examined the Neotropical
genera for these structures, except in the
cases of Crossodactyhis and Pljymlae-
mus; Baldauf and Tanzer (1965) pro-
vided data for Syrrlioplnis. All three
genera exhibit the intermaxillary duct
pattern seen in cycloranine frogs. Until
study of all of the leptodactylid genera
is completed, further comment on in-
ternal glands and their significance, if
any, must be deferred.

Myology

Thigh musculature, jaw musculature,
and the musculature of the hyolaryngeal
complex have been important characters
in the development of our current con-
cepts of the subfamilies and families of
leptodactylid and bufonid frogs. Noble
( 1922 ) divided the higher frogs into
two suborders (Procoela and Diplasio-
coela) based on the shape of the centra
in the first presacral and sacral vertebrae
and on the nature of the distal tendons
of the thigh musculature. Although nu-
merous authors pointed out that, at least
as far as the muscles were concerned,
the division was artificial, the most con-
clusive evidence against Noble's scheme
was reported by Parker ( 1940 ) , who
based his arguments on Australo-Papuan
leptodactylids. He demonstrated a pro-
coelan to diplasiocoelan transition in the
distal tendons of the thigh musculature;
the procoelan condition ( ventral position
of the distal tendon of the m. semiten-
dinosus relative to the m. gracilis) was
regarded as more generalized by Parker
and found to be restricted to the Cyclo-
raninae, whereas the advanced ( diplasio-
coelan) condition was found only in
the Myobatrachinae. Further evidence
that Noble's arrangement is artificial is
found in the elosiine leptodactylids.
Crossodactyhis has a ranoid pattern of
distal thigh tendons, whereas the closely
allied genera, Hylodes and Megaelosia,
have the bufonoid pattern. The signifi-
cance of this variation is discussed in
the systematic summary (Elosiinae, p.
164).

Limeses ( 1964 ) defined a bufonid and
leptodact>'lid pattern of thigh muscula-
ture based on the patterns in Bufo and
Leptodactylus respectively. She demon-
stiated a continuum of muscle patterns
within the Neotropical leptodactylids
from the bufonid condition (Cerato-
phrys, Lepidobatrachus, Odontophrynus,
and Froceraiophrys cristiceps) to an "in-
termediate" condition ( Macro genioglot-
tiis and Froceraiophrys, except P. cristi-
ceps) to the leptodactylid condition. She
regarded the leptodactylid condition as
grading from a cycloramphine pattern
(most like the bufonid type) through a
telmatobiine pattern to the leptodactylid
pattern. Limeses' study demonstrated
the inadvisability of using the thigh
musculature in classifications above the
generic level; she felt that the data sup-
ported the contention of Reig and his
students that the ceratophryines are fa-
milially distinct and more closely allied
to bufonids.

Limeses (1964) concluded that the
thigh muscle complexes of Noble were
"progressive" characters and therefore
should not be given much weight in de-
fining families or suborders. Griffiths
( 1959, 1963 ) pointed out variability in
the distal tendons in the dendrobatids ( a
variability which overlaps the spectrum
seen in myobatrachines ) , but he still at-
tached considerable importance to the
character complex because the muscula-
ture supported his unique contention
that dendrobatids are a subfamily of
ranids.

The conclusion of Tihen (1965) and
Inger (1967) that in the course of evo-
lution an increase in the complexity of
the musculature is to be expected car-
ries much weight in support of the argu-
ment that the diplasiocoelan condition
is labile as evidenced by its appearance
in the Myobatrachinae, Dendrobatidae,
and in Crossodactyhis. I am not of the
opinion, in view of the other data avail-
able, that the diplasiocoelan condition in
the Myobatrachinae suggests ranoid af-
finities, although it is not inconceivable

34

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

that the origin of the arthroleptine raiiids
might be the Australo-Papuan Myobatra-
chinae. At present, the arrangements of
the thigh muscles in the cycloranine and
myobatrachine leptodactyhds are re-
garded as only one additional character
in support of Parker's ( 1940 ) subfamilial
classification.

Griffiths (1959, 1963) used the mus-
culature of the temporal arch as one of
the two characters separating leptodac-
tylids and hylids from bufonids. In the
former group the m. depressor mandihu-
lae is divided into two slips (pars tym-
panicus and pars scapularis), whereas in
the bufonids the pars scapidaris is ab-
sent. Limeses ( 1965 ) investigated the
m. depressor mandibiilae in several of
the "ceratophrydids" and compared her
data with that for a number of "normal"
leptodactylids. I dissected at least one
specimen for each leptodactylid genus
except Amhlijphnjnus and Hijlorina in
order to examine the state of the m. de-
pressor mandihidae. The only taxa with
the bufonid type [pars tympanicus only)
are Crinia, Eupsophus juninensis, Lepi-
dobatrachiis, Philoria, and Pseudo-
phryne. A massive pars tympanicus and
minute pars scapularis are exhibited in
several Eleiitherodactylus (biporcatus,
bufoniformis, cornutus, galdi, stdcatus),
Proceratophrys (except in P. cristiceps),
and Tebnatobufo; the condition seen in
Ceratophrys and Macrogenioglottus is
slightly more leptodactylid, but the pars
scapularis is still small. In Odonto-
phrynus and Proceratophrys cristiceps
both slips are large. In the remaining
leptodactylids both muscle slips are pres-
ent, and the pars scapularis is usually
larger.

Starrett (1968) studied the jaw mus-
culature in many anurans and distin-
guished several insertional patterns of
the m. depressor mandibulae. She con-
sidered the condition of this muscle too
variable to be used in defining families
but useful at lower taxonomic levels
when used in combination with the ad-
ductor muscles. Based on my study of

most of the leptodactylid genera, I con-
sider the m. depressor mandibulae to be
of questionable value in leptodactylid
frog classification. I did not investigate
the condition of the adductor muscles in
leptodactylids. The reader is referred to
Starrett (1968) for a detailed review of
anuran jaw musculature.

An associated character is the path-
way of the mandibular ramus of the tri-
geminal nerve through the muscle
blocks. Limeses (1965) investigated this
character for several of the broad-
headed South American leptodactylids
and considered it useful in defining her
"Ceratophrydidae" — which included only
parts of some genera (for example, Pro-
ceratophrys).

The musculature of the hyolarynx has
been successfully used by several authors
in systematic studies of frogs. The mus-
culature is discussed with the hyoid ( see
below).

Hyolaryngeal Apparatus

Trewavas ( 1933 ) investigated the
hyolarynx in a wide variety of frogs.
Among the sixty species he treated were
nine leptodactylids. Parker (1940) used
the shape of the hyoid plate, condition
of the cricoid cartilage ventrally, and
the patterns of insertion of the deeper
throat muscles on the hyoid plate in
separating the two subfamilies of the
Australo-Papuan leptodactylids. Grif-
fiths (1959) described the hyolarynx of
Sminthillus. Among the potentially use-
ful characters in the hyolarynx of lepto-
dactylids are 1) the extent and shape of
the alary process of the hyoid plate, 2)
branching of the hyale, 3) variation in
the postero-lateral process of the hyoid
plate, 4) division (ventrally) of the
cricoid cartilage, and 5) attachment of
the m. petrohyoideus anterior and m.
sternohyoideus on the hyoid plate.

The cricoid is divided ventrally in all
genera of the Myobatrachinae but is
complete in all other genera of the fam-
ily. Griffiths (1963) reported the cri-
coid to be divided ventrally in Lepto-

LYNCH: LEPTODACTYLOID FROGS

35

dachjlus prognathus, but I have been
unable to verify this observation in my
specimens. The esophageal process of
the cricoid has been used as a systematic
tool, but it is apparently of such vari-
ability (poorly to well developed) that
its use at the generic level is not wise.
In ranoids the esophageal process is
elongate, as it is in Heleophryne, but in
most leptodactylids the process is either
reduced in size or only of moderate size.

In the Myobatrachinae, the alary
processes of the hyoid are broad and
wing-like, and the m. petrolnjoideus an-
terior inserts on the alary process. The
elongate insertion of the m. sternoJujoid-
eus begins anteriorly on the hyoid plate
medial to the alaiy process and pos-
teriorly moves nearer the midline of the
hyoid plate (Fig. 8). In contrast to the
condition seen in the myobatrachines,
the cycloranines have a narrow stalked
alary process and the m. petrohijoideus
anterior inserts on the posterolateral
edge of the hyoid plate. The m. sterno-
Injoideus inserts medially on, but at the
edge of, the hyoid plate. Anteriorly, its
insertion is like that seen in the myo-
batrachines, but posteriorly the muscles
are disposed more laterad. The muscle
disposition in Heleophryne is exactly
like that seen in the Cycloraninae; the
only differences in the hyolarynx be-
tween the Cycloraninae and Heleo-
phryne are the presence of an anterior
process on the cornu and the elongate
esophageal process in Heleophryne.

Among Neotropical leptodactyhds,
only the Leptodactylus marmoratus
group, Physahemus (including Engy-
stomops and Eupemphix), and Pseudo-
paludicola have the myobatrachine pat-
tern of muscle insertions on the hyoid
plate. These genera also have the ex-
panded and wing-like alary processes of
the hyoid plate as does Hydrolaetare.
The alary processes are absent in Cera-
tophrys, Euparkerella, Holoaden, Lepi-
dobatrachus, Limnomedusa, and Smin-
thillus. Holoaden and Pseudopaludicola
lack the posterolateral processes of the

hyoid plate. The posterolateral proc-
esses of the hyoid plate are large in the
Ceratophryinae and Limnomedusa (Fig.
9). This enlargement may be a struc-
tural compensation for the loss of the
alary processes in these genera. An-
terior processes of the cornua are lacking
in the Australo-Papuan genera and in
about one-half of the Neotropical gen-
era. I found anterior processes of the
cornua in Edalorhina, Eleutlierodacty-
lus, Euparkerella, Eupsophus, Holoaden,
Heleophryne, Hylactophryne, Ischno-
cnema, Lithodytes, Megaelosia, Nice-
foronia, Odontophrynus, Proceratophrys,
Physalaemus, Sminthillus, Syrrhophus,
and Tomodactylus. Specimens were not
available of Amhlyphrymis, Batrachyla,
Caudiverbera, Crossodactylodes, Hylo-

FiGURE 8. Hyoid plates of Heleophryne natalen-
sis (A, KU 105925, X 5) and Physalaemus
pusiulatus (B, UIMNH 80710, X 10) illus-
trating the two muscle insertional patterns. The
insertion site for the m. sternohyoideus is stip-
pled; the other muscle is the ni. petrohijoideus
anterior.

36

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

rina, and Telmatohufo. The cerato-
phiyines have broad cornua in compari-
son to other leptodactylids. The entire
hyoid plate is elongate in EuparhereUa,
Holoaden, and Smintliilhis in contrast to
all other leptodactylids (Fig. 9).

Figure 9. Hyoid plates of (A) Limnomediisa
macwglossa (female, KU 74327, X 2.5), (B)
Lepidohatrachus asper (male, KU 86782, x
4.5), (C) Holoden hradei (female, KU 107086,
X 4.5 ) and ( D ) Stninthilhis limbatiis ( after
Griffiths, 1959).

Variations in the hyolarynx appear
useful in comparing and contrasting lep-
todactylid genera. In some cases, the
hyolaryngeal data support the classifica-
tion at the tribe or subfamily level — the
Australo-Papuan groups and the Cerato-
phiyinae. The variation in the shape of
the hyoid plate among the Neotropical
leptodactylids is impressive in view of
the uniformity of the structure within
the two Australo-Papuan subfamilies.

Chiomosome Numbers'*

In recent years, increasing attention
has been given chromosome numbers in

' The chromosome number account presented
here includes only 15 species of leptodactylids.
The efforts in this area have greatly increased
the number of species (78) for which counts
are available. The pertinent literature includes
Barrio (1970), Barrio and Rinaldi de Chieri
(1970a, 1970b), Be(,ak (1968), Be(,ak, Denaro,
and Begak (1970), Bogart (1970), Brum-

Zorrilla and Saez (1968), Morescalchi and
Gargiulo (1968), and Morescalchi, Gargiulo,
and Olmo (1968). Rabello (1970) summarized
much of the data but was unaware of other
observations; she published counts contrary to
those reported in some of the original work. The
somatic counts range from 18 to 104, although
Becak, Denaro, and Becak (1970) stated that a
\'alue of 16 is the lowest known. The highest
counts are for polyploid populations. Values of
2N =8X^104 are known for Ceratophnjs dor-
sata and C. ornata ( Barrio and Rinaldi de
Chieri, 1970b). Tetraploid counts (2N=4X=
44) are known for Odontophnjnus americanus
(Bogart, 1967) and Pleurodema bibronii and P.
kricgi ( Barrio and Rinaldi de Chieri, 1970a).
The published counts for 67 other Neotropical
and six Australian leptodactylids range from 18
to 36. The six Australian leptodactylids (Crinia,
Limnodijnastes, Pseudophryne, and Uperoleia)
have 2N=24 (Morescalchi, Gargiulo, and Olmo,
1968).

The diploid counts for all leptodactylid gen-
era are siimmerized below. The data are pre-
sented as follows: genus (number of species
for which counts are known), 2N value(s), and
where germane, citations of intrageneric (and
intraspecific ) variations. Polyploid counts are
not repeated here. The names used in the orig-
inal accounts have been brought into line with
the generic classification employed in this work.
Batrachyla (2), 2N=26; Caudiverbera (1), 2N=
26; Ccmtophnjs (3), 2N=:26; Crmia (2), 2N=
24; Cro.ssodactyhis (2), 2N=26, however,
Brum-Zorrilla and Saez ( 1968) reported 2N=22
for C. gaudichaudii, whereas Bogart (1970)
obtained 2N=26 for the same species; Cyclo-
ramphus (3), 2N=26, however, Brum-Zorrilla
and Saez ( 1968) reported 2N=:22 for C. /t///g/-
nosiis (sic), whereas Bogart (1970) obtained
2N=26 for the same species; Eleuthewdactyhis
(11), 2N=18(2), 20(2), 22(4), 34(2), and
36(1), see Bogart (1970); Eupsophus (4),
2N=22(1), 26(1), 28(1), and 30(1), see Bo-
gart (1970) for a detailed account; Hylacto-
phryne (1), 2N=22; Hylodes (2), 2N=26;
Insuctoi)luynus (1), 2N=26; Lcpidobat radius
(3), 2N=26; Leptodactylus (8), 2N=22;
Limnodynastcs (2), 2N=i 24; Lithodytcs (1),
2Niizl8; OdontopJirymis (4), 2N=22; Physa-
lacnnis (6), 2N=22; Pleurodema (6), 2N=22;
Proccratophrys (2), 2N=22; Pseudopahidicola
(2), 2N = 18, 20, and 22, P. ameghini is 2N=20
and P. falcipes has 2N values of 18, 20, and 22
(Brum-Zorrilla and Saez, 1968, and Batistic,
Beyak, and Vizotto, 1969, discussed the situ-
ation in this genus); Pseudophryne (1), 2N=24;
Syrrhophus (1), 2N=30; Tehnatobius (2),
2N=22 and 26, see Barbieri (1954) and Brum-
Zorrilla (1968); Thoropa (1), 2N=26; Tomo-
dactylus (1), 2N=22; Uperoleia (1), 2N=24;
and Zachacnus ( 1 ), 2N=26.

James P. Bogart has a considerable amount

LYNCH: LEPTODACTYLOID FROGS

37

systematic studies. Data are only now
being accumulated for the leptodacty-
lids, and to date no counts are available
for the Australo-Papuan or African gen-
era. Likewise, no data have accumlated
for the Neotropical subfamily Elosiinae

Bogart (1967), Moreschalchi (1967),
and Saez and Brum ( 1959 ) reported
counts for the ceratophryine genera;
Ceratophn/s calcarata, Chacophnjs pier-
otti, and Lepidohatrachus Uanensis have
2n of 26, whereas Ceratophrys ornata
has been reported to have 2n of 88, 92,
98, 104, and 108. Bogart (1967) pre-
sented convincing arguments that this
species is octaploid (n = 13).

Of the subfamily Leptodactylinae,
counts of 2n of 22 are available for five
species of Leptodactylus and Pseudopal-
udicola falcipes (Barbieri, 1950, Bogart,
1967, and Saez and Brum, 1960).

Counts for the Telmatobiinae range
from 2n of 22 to 50. The major variation
occurs in the genus Odontophrynus — O.
americanus is tetraploid (2n of 44)
[Becak, et al., 1966, Bogart, 1967]. Saez
and Brum (1966) reported 2n = 42, 44,
and 50 for various populations of this

of unpublished observations on leptodactylid
chromosome numbers. He has counts for three
genera not included abo\'e — Edalorhina (1),
2N=22; Hcleophnjnc (1), 2N=26; and Isch-
nocncma (1), 2N=22 — and counts for addi-
tional species in eight genera. The additions are
Cycloratuphus (three more species, 2N=26),
Eleuthewdactijlus ( 19 more species, 2N=22 in
five, 26 in nine, 30 in three, and 34 in two);
Hylodes (one more species, 2N=26); Lepto-
dacttjlus (eight more with 2N=22, and two
species of the mannoratus group, subgenus
Adenomcra, with 2N=26); Physalaemus (six
more species, 2N=22); Syrrhophus (one more
species, 2N=26); Telmatobius (three more spe-
cies, 2N=26; one of these is T. montanus which
Gallardo, 1970, placed in Abodes); and Thowpa
(one more species, 2N=26). These 47 species
increase the total known to 126 representing 29
genera and all seven subfamilies. The sub-
families have counts of: Cycloraninae, 2N=24,
Myobatrachinae, 2N=24, Heleophryninae, 2N=
26, Ceratophryinae, 2N=26 (and 8X=104),
Elosiinae, 2N=26, Leptodactylinae, 2N=18, 20,
22, and 26 (and 4X=44), and Telmatobiinae,
2N = 18, 20, 22, 26, 28, 30, 34, and 36 (and
4X=44).

species. The other two species of the
genus for which counts are available
have 2n of 22 (Saez and Brum, 1966).
Tehnatohius schreiteri has a 2n =: 26
(Barbieri, 1954), whereas the other spe-
cies of the subfamily (except the tetra-
ploid Odontophrynus americanus) have
2n =r 22 (HylactopJiryne aufi^iisti, Tomo-
dactylus nitidus; Duellman, 1967).

It is generally conceded that the
more primitive members of a group have
higher chromosome numbers than do the
more advanced members of a given
group. Although this generalization is
supported by too few data to be clearly
stated, it is significant that except for i^he
cases of tetraploidy and octaploidy in the
leptodactylids, the more primitive Neo-
tropical groups ( Ceratophryinae and the
Telmatobiini ) have diploid numbers of
26, as opposed to 22 in the advanced
lines (as detennined on other grounds).
I anticipate the Cycloraninae and Heleo-
phryninae also to have diploid numbers
of 26, as do the pelobatids (Duellman,
1967, Moreschalchi, 1967). The Myo-
batrachinae might have fewer than 26
since they are morphologically diver-
gent from the Cycloraninae.

OSTEOLOGICAL CHARACTERS

With the notable exceptions of the
vertebral centra, sacral diapophyses,
terminal phalanges, and the gross pec-
toral structure, few osteological charac-
ters have been used in generic and su-
prageneric classifications of bufonoid
frogs. Cope (1865, 1866, 1889) placed
undue weight on minor variations of the
skull bones. Subsequent authors, in not-
ing the obvious errors in Cope's scheme,
tended to regard most osteological varia-
tion as trivial and therefore of no value
in a classification at or above the generic
level. The most recent author to ex-
pound this point was Griffiths (1963).
The major efforts in anuran phylogeny
have been based on the pectoral archi-
tecture ( Cope, Boulenger, and Griffiths )
and on the vertebral column and thigh
musculature (Nicholls and Noble).

38

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

I am convinced that at least within
the Bufonoidea, the osteocranium can
serve as a primary source of characters
for classification. I have completed a
generic-level analysis of the skeletons of
the frog family Leptodactylidae and
have briefly checked certain points
among the genera of the Bufonidae,
Centrolenidae, Dendrobatidae, Hylidae,
and Pseudidae. Additional data are
available in the pectoral and pelvic
girdles and in the vertebral column. I
consider the appendicular skeleton of
relatively little use, although the shape
of the terminal phalanges and the fusion
of the tarsal bones are useful sources of
characters.

Maxillary Arch

The premaxillae, maxillae, septomax-
illae, and quadratojugals make up the
maxillary arch. The quadratojugals are
the only bones that are variable in occur-
rence. In no case are the premaxillae
fused. Loss of the quadratojugals occurs
sporadically throughout frog families
from the most primitive (Leiopelma,
Pipa) to the advanced (some Hyla). The
quadratojugals have been lost in six lep-
todactylid genera — Batrachyla, Crosso-
dactylus, Hylorina, Notaden, Pleurode-
ma, and Pseud opaludicoh. The maxil-
lary arch is incomplete (Fig. 10) but
the quadratojugal is present in several
leptodactylids. The degree of separation
of the maxilla and quadratojugal is vari-
able. Most leptodactylids have the quad-
ratojugals in articular contact with the
maxillae, although the contact is tenuous
in some genera (Hylodes).

Fusion of the elements of the maxil-
lary arch (maxilla and quadratojugal) is
seen only in the genera of the subfamily
Ceratophryinae (Ceratophrys and Lepi-
dohatrachus). These frogs and the
Chilean Caudiverhera caudiverhera have
bony, stegocephalian-like skulls with a
complete and small orbit. In contrast to
the latter genus, the ceratophryines are
characterized by the fusion of all of the
skull bones (except the columellae, sep-

tomaxillae, and premaxillae) into an
akinetic unit.

Loss of the maxillary and premaxil-
lary teeth was once considered a family
character in frogs, and very often its use
resulted in the erroneous association of
several leptodactylids with the Bufoni-
dae. In his later works, George A.
Boulenger laid an ever decreasing em-
phasis on the lack of teeth in the maxil-
lary arch. Noble (1922) completely dis-
missed the character and combined the
Bufonidae and Leptodactylidae. Sub-
sequent authors have usually considered
loss of the teeth of the maxillary arch of
at least generic value; even Noble
( 1925 ) later decided that the presence
or absence of maxillaiy teeth was a ge-
neric character.

If Moore (1961) is correct in recog-
nizing a single species of Uperoleia, then
at least one species of frog has intra-
specific variation in the presence of teeth
on the maxillary arch. Most leptodacty-
lids have teeth on the maxillary arch;

Figure 10. Lateral views of the skulls of (A)
Baimrhyh Irptopiis ( UMMZ S-2246, X 6.5),
(B) Etii)soi)]iiis lusciis (AMNH 22104, X 4.2)
and (C) Cyclommphus eleutherodactyhis (KU
92785, X 3.3) illustratinjf variation in the extent
of the quadratojugal bone.

LYNCH: LEPTODACTYLOID FROGS

39

Batrachophnjmis, Glauertia, Metacrinia,
Myohatrachus, Notaden, some species of
PJiysolaenuis (maculwentris, nanus, nat-
tereri, ohtectus, petersi, pnstulosus, and
signifenis), Pseudophnjne, Sminthillus,
some species of Telmafobius, and some
Uperoleia lack teeth. Fhnjnanodus na-
nus Ahl was long thought to be tooth-
less, but examination of the unique holo-
type revealed a full complement of teeth
and satisfied me that the monotypic
genus was identical to Eleutherodactylus
parvus (Lynch, 1968c).

Loss of teeth is either uniform or
\'ariable in ten leptodactylid genera;
these genera belong to four clearly de-
finable subfamilies, none of which is de-
fined by tooth loss. Notaden is the only
cycloranine which has no teeth; it is
closely related to Neobatrachus, a
toothed genus. Of the five toothless
myobatrachine genera, Fseudophryne is
most closely related to Crinia, a toothed
genus; the other four edentate genera
form a group but are not closely related
to one another. Tooth loss occurs in the
subfamily Leptodactylinae only in Thysa-
laemus. Not all species of the genus lack
teeth; this variation has contributed to
unnatural classifications, in which some
species of the genus were placed in the
Bufonidae and others in the Cystignathi-
dae. Most of the species of the nattereri
and petersi groups of Fhysalaemus lack
teeth — one species in each group has
teeth. All species of the Fhysalaemus
cuvieri group have teeth. Batraclwphry-
nus and its Oligocene relative, Neopro-
coela, are toothless but closely allied to
the toothed genera, Caudiverbera, Tel-
matobufo, and Telmatobius. A few spe-
cies of Tehnatobius are toothless and
have been erroneously associated with
Batrachophrynus. As Vellard (1951)
and Cei and Roig ( 1968 ) pointed out,
the toothless species of Telmatobius are
closely related to toothed species. Smin-
thillus is an edentate derivative of the
West Indian complex of Eleutherodac-
tylus.

Noble (1931) contended that the

Criniinae ( =zCycloraninae -\- Myoba-
trachinae) had deeper maxillae than did
the Neotropical leptodactylids. His ob-
servations apparently were very limited
because among the Australo-Papuan lep-
todactylids, the Myobatrachinae have
shallow maxillae (little or no facial lobe
of the maxilla) whereas deep maxillae
are found in the Cycloraninae. The
maxillae of the cycloranines are no deep-
er than those seen in many Telmato-
biinae (e.g., some Eleutherodactylus,
Odontophrynus, Froceratophrys) or Lep-
todactylinae (Fleurodema). The depth
of the maxilla appears to be correlated
with the degree of reduction of ossifica-
tion. Uperoleia and Myohatrachus have
deeper maxillae than the other myo-
batrachines, and Uperoleia is the only
myobatrachine lacking an extensive fron-
toparietal fontanelle.

The alary processes of the premaxil-
lae are well developed in all leptodac-
tylid genera, except Myohatrachus, in
which they are essentially obsolete ( Fig.
11). The extensive reduction of the an-
terior portion of the skull of Myohatra-
chus has resulted in minute premaxillae.
The palatal shelf is still well developed
and similar to those of the other Myo-
bati-achinae. The reduction of the an-
terior cranial elements in Myohatrachus
is compensated for by the development
of massive palatines.

The alary processes are directed an-
terodorsally, dorsally, or posterodorsally
(Fig. 10), and the slope is apparently
correlated with the development of bur-
rowing habits; the more posterior the
slope, the greater the burrowing ten-
dencies. In several genera, most notably
members of the Elosiinae, the alary
processes have a lateral vector as well.
In general, the alary processes are rela-
tively narrow near their base in the Af-
rican and Neotropical genera (Fig. 11),
whereas the processes are very broad at
the base in the majority of the Australo-
Papuan genera (Fig. 11). The range of
variation of this character in the sub-

40

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 11. Variation in the premaxillae of lep-
todactylids. (A) median view of left premaxilla
of Ceratophnjs calcarata, JDL S-237, X 2.5;
(B ) frontal view of same; ( C ) median view of
left premaxilla of Eleiithewdactt/Ius curtipes,
JDL S-351, X 10; (D) frontal view of same;
(E) frontal view of left premaxilla of Batracho-
phnjmis macrostomus, KU 98127, X 2; (F)
frontal view of left premaxilla of Mixophijes
fasciolattts, KU 56627, X 2.5; (G) ventral view
of right premaxilla of Batrachophrynus macro-
stomus, KU 98127, X 2; (H) ventral view of
right premaxilla of Eleutherodactylus ctniipes,
JDL S-351, X 10; (I) ventral view of right
premaxilla of Batmchyla leptopus, UMMZ S-
2246, X 9; (J-K) frontal and ventral views of
bones of snout of Myohatrachus gouldi, KU
110333, X 5. Following abbreviations are used:
m — maxilla, n — nasal, s — septomaxilla, ap —
alary process, and ps — palatal shelf.

family Telmatobiinae is as great as the
variation throughout the family.

The palatal shelf of the premaxilla is
functionally absent in the Ceratophryi-
nae but present in all other leptodactylid
genera. The variation in this structure
in the non-ceratophryines involves depth
of the shelf, degree of medial dissection,
and the development of the palatal

process (Fig. 11). Deep palatal shelves
characterize Batrachophrynus and Neo-
procoela and may reflect loss of teeth,
although the shelf is relatively deep in
HoJoaclen, a toothed genus. Batraclitjh,
Crossodactijhis, and Hijlodes have very
shallow palatal shelves and long palatal
processes. Telmafobius and Thoropa,
and to a lesser extent, PhijsaJaemus, have
shallow palatal shelves, which extend
laterally to meet the maxillae. In each
case, the palatal process is either elon-
gate or large, or both. Deeply dissected
palatal shelves occur in Eleutlierodactij-
his, Hylactopliryne, Ischnocne?na, Lith-
odytes, Niceforonia, Paratehnatohius,
some Physalaemus, Proceraiophrys,
Smintliilhis, SyrrJwphus, and Tomodac-
tyhis. The shelf is most deeply dissected
in some Eleiitherodactyhis, Hylacto-
phryne, Niceforonia, and Tomodactyhis.
The depth of the palatal shelf may re-
flect stress on the anterior end of the
skull during burrowing.

The Australo-Papuan genera exhibit
almost no variation in the shape and
extent of the palatal shelf.

The maxillae show only minor varia-
tion in the depth of the facial lobe, the
degree of Haring of the maxilla, and the
development of exostosis. Megaelosia
is unique among leptodactylids in its
posterior expansion of the maxilla (Fig.
12), which is characteristically a taper-
ing bone. Exostosis is seen in Caiidiver-
hera, Cyclorana australis, the cerato-
phryine genera, some Eleutherodactylus
{cornutus and unistrigatus groups), and
some species of Proceraiophrys.

The range of variation in the palatal
shelf of the maxilla is less than that in
the premaxilla. In most genera, the pos-
terior end of the maxillary palatal shelf
is expanded (pterygoidal process) and
abuts or overlaps the anterior ramus of
the pterygoid. In some genera (e.g.,
Caudiverbera), there is a definite suture
between the pterygoidal process and the
pteiygoid (Fig. 12). This character is
apparently correlated with a more solid
skull architecture.

LYNCH: LEPTODACTYLOID FROGS

41

Figure 12. Expansion of the posterior portion
of the maxilla in (A) Megaelosia gocldi ( KU
92966, X 4). Maxilla-pterygoid articulation in
(B) Leptodachtijhis pentadactyhis (KU 68159,
X 1) and (C) Caiidiverbera caudiverbera
(FMNH9703, X 1).

Teeth, when absent on the premaxil-
la, are also absent on the maxilla. In all
toothed genera except Adelotiis, the
teeth form a continuous row from the
suture between the premaxillae to a
point near the end of the maxilla, pos-
terior to the maxilla-pterygoid junction.
In Adelotiis (Fig. 13) there is a distinct
diastema between the maxillary and pre-
maxillary teeth.

Leptodactylid teeth are of two types,
exemplified by Ceratophnjs and Odon-
tophrijnus. In Ceratophnjs and four
other genera, the teeth are fang-like,
some (e.g., Megaelosia) more so than
others, whereas in Odontophnjniis and
most other genera, the teeth are blunt
with some indications of bulbing and
cusp formation (like that described for
several hylids — Duellman and Trueb,
1966, Trueb, 1966). The teeth in Megae-
losia are six to seven times as long ( pro-
portionately) as those in most other lep-
todactylids.

Parsons and Williams (1962, 1963)
characterized the Lissamphia by numer-
ous features, one of which was jointed

or hinged teeth — a basal pedicel and a
distal crown. Reig and Limeses (1963)
pointed out that the ceratophryine gen-
era do not have hinged teeth, whereas
the genera often wrongly combined with
Ceratophnjs [Odontophrynus and Pro-
ceratophnjs (=zStomhus auctorum)] pos-
sess hinged teeth. The ceratophryines
have comparatively long teeth (here
termed "fang-like" for purposes of dis-
cussion). Among the American genera,
elongate fang-like teeth are found in
Ceratophnjs (including Chacophrys) ,
Lepidohatrachus, Megaelosia, and Tel-
matohius. The teeth of Crossodactijlus,
Hylodes, and Telmatobufo tend to be
pointed and elongate but are not com-
parable to those of members of the
"fang-like teeth" group. Distinct pedi-
cel-crown development is seen in the
"fanged" Megaelosia and Telmatobitis
but not in the Ceratophryinae (Fig. 13).

Among the toothed Australo-Papuan
leptodactylids, fang-like teeth are found
only in the cycloranine Adehtus; it and
all other toothed Australo-Papuan gen-
era and Heleophryne have pedicelate
teeth.

Goin (1959) published a note on the
number of teeth per maxilla in approxi-

FiGURE 13. (A,B) Anterior portion of the skull
of Adelotiis brevis ( KU 56242) illustrating the
diastema. (C) Maxillary teeth of Pwceratophnjs
appendiculata ( KU 93070) and (D) same of
Ceratophrys cakamta (JDL S-237). Teeth of
Pioceratophrys are pedicellate; those of Cerat-
ophrys are non-pedicellate. A and B, X 3; C
and D, X 12.5.

42

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

mately 65 species of EJeutherodactijlus
and 12 species of Leptodactyhis; the
range in the former was 33 to 111 per
maxilla and in the latter 43 to 79. Goin
asserted that, within the species groups
with which he was familiar, the species
groups could be characterized on the
basis of average number of teeth. I con-
sider the number of teeth, in most cases,
to more accurately reflect the average
size of the species studied than relation-
ships. Most species of Eleutlwrodactyliis
have between 50 and 80 teeth on each
maxilla. Reig and Limeses (1963) noted
the low tooth counts of the cerato-
phryines ( about 25 per maxilla ) as com-
pared with 50 or more per maxilla in
most other leptodactylids. They reported
the following data for the cerato-
phryines: Chacophnjs, 22 ± 2 per max-
illa; Ceratophnjs, 27 ± 3, and Lepido-
hatrachus, 28 ± 5. In Megaelosia there
are 29 to 34 teeth per maxilla (based on
four individuals); these data compare
favorably with those for the cerato-
phryines and for Adelotus (26 to 28
teeth per maxilla).

The septomaxillae are small, incon-
spicuous elements of the maxillary arch
and are intricately associated with the
nasal cava. The shape of the bones re-
flects the connecting pathways of the
cava. The septomaxillae are proportion-
ately minute in Ceratoplinjs when com-
pared with those of most leptodactylids.
The smallest species of the family have
simple U-shaped septomaxillae ( Baldauf
and Tanzer, 1965 ) . I have examined few
leptodactylids for the detail of the septo-
maxillae, but have determined that the
bones are invariably present. The septo-
maxillae of several genera of leptodacty-
lids are illustrated in Figure 14. Serious
study of the septomaxillae will probably
require study of serially sectioned ma-
terials.

Lower Jaw

The anuran mandible contains two
bones, the dentary and angular, and the
mentomeckelian cartilage (Fig. 15). No

leptodactylid has true teeth on the lower
jaw, although odontoids are developed
to varying degrees by several species.
Grypiscus was once thought to have
mandibular teeth, but Noble (1922) re-
ported these "teeth" to be a serrate ridge.
1 have not found a serrate ridge on the
mandible of any Cydoramphus (^Gry-
piscus) or any other leptodactylid frog
except large Ceratophrys. Many indi-
viduals of Ceratophrys have a sharp
odontoid on the mentomeckelian carti-
lage near the mandibular symphasis.
The only leptodactylid with large odon-
toids is Adelotus hrevis (Fig. 15). These
tusk-like odontoids are better developed
in the male than in the female. The
maxillary diastema of Adelotus is ap-
parently a spacing adaptation for the oc-

FiGURE 14. Septomaxillae of several Neotropical
leptodactylid frogs. (A-D) dorsal, ventral, lat-
eral and median views of right septomaxilla of
Leptodactyhis pentadactijlus (KU 84982, x 9.5),
(E) ventral view of right septomaxilla of
Eleutherodacttjhis flcischmanni ( KU 68158, X
9.5), (F) posteroventral view of right septo-
maxilla of Ceratophrys calcarata ( JDL S-237,
X 16), (G) lateral view of left septomaxilla of
Hylodes lateristri^ata ( KU 92878, X 9.5),
(II-I) lateral and dorsal views, respectively, of
left septomaxilla of Tchnatohius liaittholi (KU
72879, X 9.5).

LYNCH: LEPTODACTYLOID FROGS

43

Figure 15. Left mandibles of (A-B) Cerato-

phrt/s aurita ( EHT 1415, X 1), and (C-D)

Addotiis brevis (AMNH 59490, X 2.5). Lateral

and dorsal views, respectively.

elusion of the teeth of the maxillary arch
and the odontoid of the dentary.

Roof of the Skull

The following bones are included in
this category: nasals, sphenethmoid (en-
dochondral), frontoparietals, prootics,
and exoccipitals. Unlike some of the
casque-headed hylids, the casque-
headed leptodactylids always lack a der-
mal sphenethmoid. Trueb (1966, 1970)
reported a dermal sphenethmoid in some
casque-headed, co-ossified hylids (Apor^
asphenodon, Conjthomantis, some Hyla,
Osteocephalus, Tmchycephalus, and Tri-
prion petasatus). The frontoparietals
and nasals are co-ossified in Caudiver-
bera and the Ceratophryinae, and are
extensively exostosed in Ctjclorana aus-
tralis and several EleitthewdactyJus.
Functionally, the prootic and exoccipital
form a single bone (otoccipital), al-
though they are not always fused (per-
haps paedomorphic).

The nasals exhibit considerable varia-
tion in size and position relative to the
premaxillae, sphenethmoid, and fronto-
parietals. Large nasals are present in
some members of those genera with the
nasals in medial contact (e.g., Eleuthero-
dactyhis), and small nasals are usually
correlated with medial separation (e.g.,

Hylodes). In general, widely separated
nasals are correlated with a proportion-
ately large sphenethmoid; however,
large sphenethmoids may also be found
in genera with closely juxtaposed nasal
bones. Separation of the nasals does not
always correlate with the presence of a
large frontoparietal fontanelle (Fig. 16).
A large sphenethmoid usually is corre-
lated with an anterior rotation of the
alary processes of the premaxillae, pre-
sumably reflecting the forward shift of
the nasal capsule. At present, I am un-
willing to state that nasal and fronto-
parietal separation always correlates
with a reduction in the amount of bone
in the skull or that the enlargement of
the sphenethmoid correlates with the
abo\'e. I consider the variations in the
bones of the roof of the skull as dis-
cordant. A frontoparietal fontanelle is
present in juveniles of all leptodactylids
[even Ceratophnjs just before metamor-
phosis (Fig. 74)] but is retained in rela-
tively few genera ( 21 ) in the adult. The
South African Heleophryne has a large
frontoparietal fontanelle, and large fon-
tanelles are seen in the myobatrachine
genera Crinki, Glauertia, Metacrinia,
Myohotrachus, and Pseudophryne but
not in Taudactyhis or Uperoleia. The
skull roof is solid in the cycloranines
Adelotus, CycJorarm, Lechriodm, and
Mixophyes, whereas in Heleioporus,
Ki/arranm, Limnodynastes, Neobatrach-
us, Notaden, and PhUoria moderately
well-developed frontoparietal fontanelles
are present. Frontoparietal fontanelles
are lacking in the Ceratophryinae, Elo-
siinae, and all but two genera of the
Leptodactylinae (Paratehnatobius and
Pleuwdema), although the leptodactyline
PseudopaJudicola could be described as
having a long, narrow fontanelle (Fig.
17). Among the Telmatobiinae, only
two genera (BatrachyJa and Thoropa)
have moderate-sized fontanelles; small
fontanelles are found in Batmchophry-
niis, Eupsophus, Holooden, Neoprocoeh,
Tehnatobius, and Tehnatobtifo. The
skull is completely roofed in the other

44

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 16. (A) Skull
diastema, KU 68263) ;

of a leptodactylid lacking a frontoparietal fontanelle {Eleutherodachjlus
uid (B) one with a relatively large fontanelle (Thoropa hitzi, KU 92850).
Both X 6.8.

Fif;uRE 17. Separation of nasal bone.s.
ceiatoplinjs hoiei ( KU 93076, X 2.3);

(A) Pscudopahidicola salfica ( KU 93068, X 6.8); (B) Pro-
and (C) Elcutlicwdactijlus richmondi (AS 12623, X 4.5).

LYNCH: LEPTODACTYLOID FROGS

45

17 Neotropical leptodactylid genera.
Few species of Eleutlierodactyhis char-
acteristically have a frontoparietal fon-
tanelle as an adult. Boulenger ( 1882 )
reported this condition in E. cuiiipes,
but I think he confused the skeletons of
E. curtipes and E. ichijmperi.

The nasals are in median contact in
Amhhjphnjmis, Barycholos, Batracho-
phnjmis, Caiidwerbera, Crossodacty-
lodes, Cycloramphus, Eleutlierodactylus,
Hylocfophnjne, Ischnocnema, Mixo-
phyes, Fhiloria, Scythrophrys, Syrrho-
phus, Tomodactyhis, Zachaenus, in all
but two genera (Paratehnatohius and
Pseudop(dudicoJa) of the Leptodactyli-
nae, the Ceratophryinae, and are in con-
tact or very narrowly separated in all
but one genus (Taiidactylus) of the Myo-
batrachinae. The nasals are slightly to
moderately separated in Adelotus, Cy-
clorana, Eiiparkerelh, Eiipsophns, He-
leioporus, Holoaden, HyJorina, Limno-
dynastes, Megaelosia, Neohatrachus,
Niceforonia, Notaden, Odontophrynus,
Paratehnatohius, Proceratophrys, Smin-
thUhis, Tehiiatohim, Telmatohtifo, and
Thoropa. The nasals are widely sep-
arated in Batraclnjla, Crossodactylus,
Hekophryne, Hylodes, Kyarranus, Pseti-
dopahidicola and Taiidactyhis. In those
genera with slight separation of the
nasals, the separation may be due to a
general reduction of bone or to an an-
terior extension of the sphenethmoid, as
in Proceratophrys (Fig. 17). Without
developmental studies, separation of the
latter two groups is not feasible.

Contact between the nasals and fron-
toparietals is uncommon in the Lepto-
dactylidae. When these bones are in
contact, the sphenethmoid is not visible
dorsally unless there is a partial separa-
tion of the frontoparietals anteriorly (as
in Cycloramphus) or a posteromedial
separation of the nasals (as in Ambhj-
phrynus, Eleutlwrodactyhis, and Procera-
tophrys). Relatively broad nasal and
frontoparietal contact occurs in the gen-
era Caiidiverhera, Ceratophrys, Cyclo-
ramphus, Hydrolaetare, Lepidohatrach-

us, and Phdoria; tenuous contact occurs
in Glauertia, Mixophyes, and Myo-
hatrachus. The nasals and frontoparietals
of all specimens of Leptodactyhis I have
examined are separated, although Gal-
lardo ( 1964 ) reported contact in the
Leptodactyhis ocellatus group.

Ornamentation of the frontoparietals,
as expressed externally in cranial ridges
or crests, has been noted by many
authors dealing with the Leptodactyli-
dae. Development of crests has often
influenced generic recogniton (e.g.,
Eleutherodactylus hiporcatns, cornutus,
and galdi groups). Much of the early
systematic study of the Bufonidae in-
volved minor variations in the develop-
ment of such crests, and it is in Bufo
that the greatest development of crests
occurs (e.g., Bufo typhomus) . The fron-
toparietal ornamentation in leptodactyl-
ids varies from slight ridges above the
orbits (Niceforonia) to the heavy crest
development and exostosis seen in Cy-
clorana australis, Eleutlwrodactylus hi-
porcatus, E. cornutus, E. devillei, E.
galdi, E. sulcatus, and Proceratophrys
cristiceps, to the dermostosis of the skull
bones seen in the ceratophiyines and
Caudiverbera. The ornamentation be-
gins at the lateral edges of the fronto-
parietals near the posterior edge of the
orbit and proceeds posteriorly, then
medially, and finally anteriorly. Concur-
rent with the development of ornamen-
tation on the nasals, ornamentation be-
gins on the otic plate of the squamosal.

The thickening of the frontoparietal
posteriorly and/ or the involvement of
the dermis of the head in co-ossification
results in the enclosure of the carotid
artery in a bony canal. Posteriorly, this
results in a carotid foramen (Figs. 18-
19 ) . Distinct carotid foramina are found
in the ceratophryines and Caudiverbera,
but not in other Neotropical genera. The
carotid artery usually passes above the
otic region of the skull between the epi-
otic eminences and the edge of the fron-
toparietal shelf. This is true of Heleo-
phryne and all Australo-Papuan genera

46

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 18. Occipital canal and foramen devel-
opment in non-casque-headed leptodactylids.
(A) open canal, Notaden nichoUsi, KU 93582,
X L2, (B) short enclosed canal, Heleioponis
eijeri, UMMZ 124504, x 1.4, (C) long enclosed
canal, Mixophyes fasciolatiis, KU 56627, x 1-2,
and (D) Lechriodiis fletcheri, AMNH 59488,
X 1.4.

except Cylorana, Heleioponis, Kyarran-
us, Lechriodiis, Mixophyes, Neobatrach-
us, Notaden, and Philoria. In Cyclo-
rana and Notaden the carotid artery lies
in a shallow or deep, exposed channel
(Fig. 18), whereas in the other genera
the passage is overlain by bone to form
a short (Heleioponis, Kyananiis, Neo-
hatraclnis, and Philoria) to long (Lechri-
odiis and Mixophyes) canal (Fig. 18).

The crista parotica is, for purposes of
my discussion, more a region than a

B,, ->.

"--^lP^

^s^^

S5^~x

^^g

— ^^

Figure 19. Squamosal-maxillary angles in lep-
todactylid frogs. (A) Neohatrachus picttis, /_
=55° (FMNH 97281, X 1.2, (B) Myobatrachtis
goiddi, angle not subject to measurement ( KU
110333, X 2.4, (C) Eleuthewdactyhis bufoni-
formis, /=53° (KU 80621, X 0.6), and (D)
Leptodacfyhis insidarum, / =36° ( KU 41026,
XO.6).

structure. It is the otoccipital (fused '
prootic and exoccipital) lateral to the
epiotic eminences. In leptodactylids the
cristae paroticae are either short and
stocky or long and narrow. In Eleuthero-
dactyhis these two groups tend to grade
into one another.

Fusion of the otoccipital to the fron-
toparietal is known in several bufonid
groups (Tihen, 1960a, 1962a) as well as
in SyrrJwplms (Baldauf and Tanzer,
1965). Among the 57 genera of lepto-
dactylids this fusion occurs only in
EleutJierodactyhis and some of its allies
(EuparkereUa, SmintliiUus, Syrrhophus,
and Tomodactyhis). All but one species
group of West Indian EleiitJierodactyhis
{inoptatiis group) have the otoccipital
and frontoparietal fused, as do some of
the species of Eleiitlierodactyhis living in
the higher parts of the Andes in Colom-
bia, Ecuador, and Venezuela.

Schaeffer (1949) regarded the pres-
ence of a temporal arcade (articulation
between the squamosal and frontopari-
etal over the cristae paroticae) to be
unique to Caudiverbera, among lepto-
dactylids. The distribution of the char-
acter in other frog families is sporadic.
In several genera, lateral growth of the
frontoparietal is such that the fronto-
parietal almost meets the squamosal, but
fails to do so because the squamosal is
not expanded medially (Anotheca,
Eopelobates, and Pelobates, except P.
ciiltripes, where contact occurs). Sev-
eral bufonids have a suture between the
otic plate of the squamosal and the
frontoparietal, as do the following hylid
genera: AparaspJwnodon, Corijthoman-
tis, Heniipliractiis, Pternohyla (fodiens,
but not dentata), Trachycephalus, and
Triprion. The suture is present in only
three leptodactylid genera — Caudiver-
bera (including Eopliractus and Gigan-
tobatrachus) , Ceratophrys (including
Chacophrys), and Lepidobatrachus;
these genera include only a dozen spe-
cies, slightly fewer than the number of
species of hylids exhibiting sutural con-
tact between the squamosal and fronto-

LYNCH: LEPTODACTYLOID FROGS

47

parietal. The intrageneric variation in
this character in Pelobates and Pterno-
Jnjla serves as adequate warning as to
the amount of weight that can be given
this character; fortuitously, there is no
intrageneric variation in this character
in the leptodactylid genera, so they can
be diagnosed by the presence of the
suture. Expression of the sutural contact
always accompanies casque develop-
ment.

Temporal Architecture

Griffiths (1954) pointed out the pos-
sible use of the squamosal-maxillary
angle in ascertaining the relationships of
higher frogs. The squamosal-maxillary
angle is the angle formed by the body
of the ventral ramus of the squamosal
and the maxillary arch. The muscula-
ture of the region (principally the m.
depressor 7nancUbnlae) has been used to
separate the Pelobatidae from the higher
frogs and to separate the Bufonidae
(and Atelopodidae) from the Hylidae
and Leptodactylidae (see page 34). The
discussion here is concerned with the
squamosal bone and the associated an-
gle. Griffiths (1954, 1959, 1963) reported
that all leptodactylids have squamosal-
maxillary angles between 45° and 50°
and that the bufonid-atelopodid com-
plex has an angle greater than 55°. My
investigation of this character complex
does not support Griffiths' statements.
Leptodactylids have an observed range
of from 15° to nearly 90°. The curva-
ture of the posterior portion of the max-
illary arch and the inner edge of the
ventral ramus of the squamosal often
renders measurements of angles difficult
or impossible. In addition to a dorso-
ventral curvature of the maxillary arch,
there is a lateral curvature of the quad-
ratojugal in many genera. Loss of the
quadratojugal precludes measurements
of the squamosal-maxillary angle. The
peculiar rotation of the cranial elements
of Myobatrachiis has resulted in a tenu-
ous contact between the strongly curved
(dorso-ventral) quadratojugal and max-

illa (Fig. 19). Use of the squamosal-
maxillary angle in any suprageneric clas-
sification of frogs is hazardous at best.
StaiTctt (1968) came to this same con-
clusion.

The presence of an otic plate of the
squamosal in casque-headed frogs was
noted above. In general, the otic ele-
ment of the squamosal is not well de-
veloped and does not overlie the otoccip-
ital. In Caudwerhera, Ceratophnjs, and
Lepidohatrachus, the otic plate is large.
In Ceratophnjs a posttemporal fenestra
(Fig. 20) is formed by the contact of
the squamosal and frontoparietal. The
otic plate of the squamosal and laterally
expanded frontoparietal rest flush on the
otoccipital in Caudiverbera and Lepi-
dobatrachus.

Figure 20. Posterior views of skulls of (A)
Ceratophnjs calcarata (JDL S-237) and ( B)
Caudwerhera caudiverbera (FMNH 9703) il-
lustrating presence and absence, respectively, of
temporal fenestrae. The lower figure is also a
type II occipital condyle arrangement; compare
with figure 29B. X 0.5.

In lateral profile, several variations
on the "typical" squamosal are apparent
in the lengths and sizes of the zygomatic
and otic rami (Fig. 21). The myobatrach-
ine genera have short, knob-like zygo-
matic rami and elongate otic rami of the
squamosals. The otic plate, if present, is
a narrow structure and not clearly de-
fined. This pattern also occurs in the
West Indian Eleutherodacttjlus, Smin-

48

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 21. Variation in length of rami of squamosals. (A) Procciatophrys boiei ( KU 9.3076, X 4),
(B) Cijcloramphus eleuthewdactijlus ( KU 92785, X 3.5), (C) Zachaenus steinegeri (KU 92747,
dorsal view X 9), (D) Eleutliewdactyhis fitzingeri (KU 117.358, X 3.5), (F) Telmatohius marmoratus
(UMMZ 68179, X 3), and (F) Eleutherodacttjlus diastema (KU 68263, X 6.8).

t-

thillus, Syrrhophiis, Tomodactyhis, and
a few mainland Eleutlierodactylus. The
otic ramus is very short in Batraclw-
phnjnus, Neoprocoela, Telmatohius, and
TeJmatohufo; in these genera, the zygo-
matic ramus is moderate in length. The
zygomatic and otic rami of all other lep-
todactylids (except Caudiverhera, Cera-
tophnjs, Cyclorana amtralis, EJeutJiero-
dactyhis ruthae, Lepidohatrachus, Me-
gaelosia, and Proceratoplirys) are equally
developed. An otic plate is developed
in many groups. Caudwerhera, Cerato-
plirys, and Lepidohatrachus have a
broad maxillary-squamosal articulation
which completes the ventroposterior
margin of the orbit. Cyclorami australis
and Proceratoplirys have broad articula-
tions between the squamosal and maxil-
la, but in these species there is no pos-
terodorsal growth of a squamosal process
of the maxilla. In Megaelosia, there is a
strong, ligamentous contact between the
bones, but a suture is not apparent. The

zygomatic ramus of Eleutherodactyhis
rutljae is very long and in tenuous con-
tact with the maxilla. A similar develop-
ment of the squamosal was noted in
Cyclorana alhoguttata by Parker (1940).

Determination of the extent of the
maxilla and squamosal in the maxillary-
scjuamosal bridge of the ceratophryines
is impossible without the aid of a de-
velopmental series. Some authors have
figured what I consider to be fictitious
suture lines. In Ceratoplirys ornata
froglets (stage 46), development of the
bridge is clearly evident. The ventral
portion of the arch is provided by a pos-
terodorsal growth of the maxilla and the
dorsal portion is made up of the zygo-
matic ramus of the squamosal ( Fig. 74 ) .

Palate

Parker (1940) presented a complete
survey of the anterior palate in the Aus-
tralo-Papuan leptodactylids as then

LYNCH: LEPTODACTYLOID FROGS

49

Figure 22. Variation in the size of the prevomerine bones of Neotropical leptodactyhds. (A) Batra-
chophnjmi.s macwstomiis (KU 98127, X 1.2), (B ) Euparkerella brasiUetms {KU 93192, X 13.4), (C)
Synliophus pipilam nchtiJosus ( KU 59950, X 6.8), (D) Eleutherodactijlus Cochrane (AS\'8014, X
9), (D) E. atkinsi (AS V6263, x9), and (F) £. hufoniformh (KU 80621, X 2.4).

understood (Neobatraclnis, PJiiloria, and
Taudactylus were not available or not
recognized by Parker). Almost no data
have been presented on the palate of
African or Neotropical leptodactyhds
except for the papers of Mehely (1904)
and Parker ( 1927 ) concerning the palu-
dicoline leptodactyhds (Physalaeimis,
Pleurodema, and Pseiidopaludicola).

Primitively, the prevomers are large
and toothed; their reduction or loss is a
derived condition. The prevomers are
minute or absent in all members of the
Australo-Papuan Myobatrachinae and
the Neotropical genera Batrachophnjnus
and Euparkerella as well as in some spe-
cies of Eiipsophus, Niceforonia, and Syr-
rhophus. An almost complete, graded
series in the size of the prevomers ( from
large to minute) can be demonstrated
within the subfamily Telmatobiinae
( Fig. 22 ) . The prevomers are minute in
Batrachophrymis and are progressively
larger in Euparkerella, Syrrhophus pipi-

lans, SrninthiUus Ihnhatus, Syrrhophus
longipes (often toothed), Eleutherodac-
tylus cochranae, E. atkinsi, and E. bu-
foniformis.

Reduction in the size of the prevomer
does not necessarily mean that the pre-
vomerine teeth are lost. Although loss of
the prevomer appears to be the trend in
Crinia and Syrrhophus, some members
of each genus have prevomerine teeth.
The prevomerine teeth have been lost in
the following leptodactylid genera: Ba-
trachophrymis, Crinia (part), Crosso-
dactylus, Crossodactylodes, Eleuthero-
dactylus (part), Euparkerella, Eupso-
phus (part), Glauertia, Metacrinia, Myo-
hatrachus, Niceforonia (part), Physalae-
mus (occasionally present in one spe-
cies), Pseiidopaludicola, Pseudophryne,
Sminthilhis, Syrrhophus (part), Taudac-
tylus, Telmatobius (part, intraspecifical-
ly variable), and Uperoleia. Only nine
leptodactylid genera are completely
edentulous (Batrachophrynus, Glauertia,

50

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Metacrinia, Myohotroclms, seven species
of Thijsalaemus, PseudopJinjne, Smin-
tliillus, Taudactylus, and one species of
Vperoleia).

Loss of the prevomerine teeth has
occurred in all members of three other
bufonoid families ( Bufonidae, Dendro-
batidae, and Rhinodennatidae) and oc-
curs sporadically in the Hylidae. In the
Dendrobatidae, the prevomerine bones
are reduced in size and may be lost in
some groups (Fig. 23).

Figure 23. Anterior palatal region of Co-
lostethus panamensis (KU 77681, X 10).

The telmatobiine genera (Eleuthero-
dactylini ) Eleutherodactijlus, Euparker-
ella, SmintJiiUiis, Syrrlioplnis, and Tomo-
dactylus show interesting geographic
variation in the size and median separa-
tion of the prevomers. The prevomers
are relatively small and widely separated
in the West Indian Eleutherodactylus,
Etiporkerella, SmintliiUus, Syrrlioplnis,
and Tomodactylus, whereas the bones
are relatively larger and only slightly
separated or in median contact in the
Middle and South American species of
Eleutherodactylus. Large prevomers (in
median contact) are found in the South
American genera Amhlyphrynus and
Ischnocnema; these genera are probably
derivatives of Eleutherodactijlus.

The prevomers of the Cycloraninae
and Heleophryninae are large and
toothed. The median separation of the
bones varies from moderate (Notaden)
to none (Mixophyes). I recognize two
tribes of the Cycloraninae — Cycloranini
and Limnodynastini. They differ in
breeding biology and in the relative

positions of the prevomerine dentigerous
processes to the choanae. The dentiger-
ous processes in the Cycloranini lie be-
tween the choanae, anterior to the pos-
terior edge of the choanae, whereas in
the Limnodynastini, the dentigerous
processes lie posterior to the choanae.
The dentigerous processes lie anterior to
the choanae in Heleophryne — the prevo-
mers of this genus are unique ( Fig. 73 ) .

Contrary to older reports in the lit-
erature, the anuran palatine does not
bear teeth. The "palatine teeth" are
either odontoids on the palatal bones or
are the prevomerine teeth. The palatine
bones are present in all leptodactylids,
but they are very small in Crinia, Eu-
parkerella, and Pseudophryne. In Myo-
hatrachus the palatines form the major
supportive elements of the anterior part
of the skull. Palatal ridges occur in sev-
eral of the larger Eleutherodactylus, in
Batrachophrynus, Caudiverhera, Cerato-
phrys, Proceratophrys, and in Cyclorana
australis. Discrete odontoids occur on
the palatal ridges in large specimens of
Caudiverhera and Ceratophrys. The
palatines are absent in several bufonid
genera (Tihen, 1960a) and in all den-
drobatids and rhinodermatids. In the
dendrobatids, loss of the palatines is
compensated for by the development of
a large palatal shelf from the postero-
ventrolateral edge of the sphenethmoid.
The dendrobatids are clearly derived
from the Elosiinae; in the Elosiinae the
palatine bones are present, and a pos-
teroventrolateral palatal shelf develops
in Hylodes but is not especially evident
in the other genera.

The parasphenoid is an unpaired
bone covering much of the ventral sur-
face of the skull. It is tri-radiate, with
an anterior ramus and two lateral alae.
The anterior ramus rests anteriorly on
the sphenethmoid, and the alae rest on
the otic capsules. In Batrachophrynus,
the anterior ramus extends well anterior
to the palatines — the parasphenoid does
not extend farther anterior in any other
leptodactylid genus. In the Elosiinae,

LYNCH: LEPTODACTYLOID FROGS

51

the anterior ramus is very short (as it is
in dendrobatids ) . A median keel is
found on the parasphenoid in a few lep-
todactyHds — the significance of a median
keel is not apparent.

The lateral alae are short or long,
oriented at right angles to the anterior
ramus or deflected posterior, and over-
lapped or not by the median rami of the
pterygoids ( Fig. 24 ) . The alae are over-
lapped by the median rami of the ptery-
goids in the Ceratophryinae, Cyclo-
raninae, Heleophryninae, in Megaelosia,
in Hydrolaetare, Leptodactyhis {ftisciis,
melanonotus, ocellatus, and pentadacty-
hi.s groups), Limnomedusa, and Pletiro-
dema (Leptodactylinae), in Amhlyphry-
nus, Batrachophrynus, Caudiverbera,
Crossodactylodes, Cyclora7npJius, Eupso-
phus, Hylactophryne, Hylorina, Ischno-
cnema, Odontophrynus, Proceratophrys,
Thoropa, and Zachaenus (Telmatobii-
nae). In many species of Eleutherodac-
tylus, the pterygoid and parasphenoid
overlap. The bones are in tenuous con-
tact in Edalorhina, Hylodes, Physahe-
mus, and Pseudopaludicola, as well as in
some Eleutlierodactyhis. The bones are
widely separated in the Myobatrachinae,
Barycholos, Batrachyla, Crossodactylus,
some EleutherodactyJus, Euparkerella,
Holoaden, some Leptodactylus (marmo-
ratiis group), Lithodytes, Niceforonia,
Paratelmatobitis, Smintliillus, Syrrho-

phus, Tehnatobius, Telnmtobufo, and
Tomodactylus.

Non-overlap of the pterygoid and
parasphenoid seems to have been ac-
complished in two ways — by simple
shortening of both elements, or by bend-
ing of the median ramus of the ptery-
goid. The latter means is apparently the
most common in the Leptodactylinae, in
which an almost complete graded series
can be demonstrated.

The tri-radiate, paired pterygoids
articulate laterally at two points on the
maxillary arch and medially they may
(or may not) rest on the otic capsule.
The anterior ramus is always longer than
the median or posterolateral rami and
is either in ligamentous or sutural con-
tact with the maxilla (Fig. 12). Tihen
( 1962b ) used the long anterior ramus of
Neoprocoela (reaching the palatines) as
one character to synonymize the genus
with the Bufo calamita group. Palatal-
pterygoid contact occurs in five telmato-
biine genera (Batrachophrynus, Odonto-
phrymis, Proceratophrys, Tehnatobius,
and Tehnatobufo), one elosiine genus
(Hylodes), and two leptodactyline gen-
era (Pleurodema and Pseudopaludicola).
The palatines and pterygoids are nar-
rowly separated in the telmatobiine gen-
era Batracliyla, Eupsophus, and Hylo-
rina, the elosiine genus Crossodactylus,
and the leptodactyHne genera Limno-

FiGURE 24. Ventral views of the skulls of leptodactylids with a pterygoid-parasphenoid overlap
(A) Eleuthewdachihis fleischmanni, KU 68157, X 2, and one without an overlap (B) E. inop-

tatus; AS X2356, X 2.4.

52

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

medusa and Pliysalaetnus; these bones
are moderately separated in the lepto-
dactyhne genera Edalorhina and Litho-
dytes. The palatines and ptciygoids are
narrowly separated in all cycloranines
except Philoria (wide separation). Ac-
tual contact of the bones occurs as a
variation in Adelotiis, Cyclorana aus-
fralis, Heleioponis, and Neohatrachus.
They are widely separated in all other
leptodactylid genera.

A ventral flange occurs on the body
of the pterygoid ( Fig. 25 ) in a few Neo-
tropical genera — Cycloromphus, Hydro-
laetare, and Zachaenus. Principally be-
cause of the absence of this character,
Zachaenus roseus Cope was removed
from the genus Zachaenus. The holo-
type of Z. roseus is a macerated heap of
fragments, and its systematic status is
uncertain. The flange may have devel-
oped for the attachment of muscles used
in burrowing in the genera Cycloram-
phus and Zachaenus; Hydrolaetare is ap-

FiGtTRE 25. Occipital view of the skull of Cijclo-

ramphtis dnhiiis ( KU 92780, X 7.5) illustrating

the enlarged pterygoidal flange.

parently aquatic, and the flange may re-
flect the habit of burrowing in mud.

Otoccipital and Columella

The otoccipital bone is the fused
prootic and exoccipital. These bones are
occasionally separate — usually in juve-
niles and in species with many paedo-
morphic features. Dorsomedially, the
otoccipitals are bordered by the fronto-
parietals. The frontoparietals and otoc-
cipitals are fused in a few genera of the
Eleutherodactylini. Ventrally, the otoc-

cipitals rest on the parasphenoid. The
columella is associated with the otoccipi-
tal and rests on the cartilaginous oper-
culum. The shape, size, and direction of
the columella show some variation; I
have not investigated these variations.

The columellae are lacking in Crosso-
dactylodes pintoi, Euparkerella hrasili-
ensis, Eupsophus juninensis, E. monti-
coki, Holoaden, Paratehnatobius, all
species of Pseudophryne, Tehnatohius
niger, Tehnatol)ufo hidlocki, and proba-
bly in Niceforonia simonsii. I am not
willing, at present, to recognize loss of
the columella (or entire middle ear) as
being of taxonomic value at the generic
level.

Gallardo (1965) regarded the degree
of median separation of the occipital
condyles of considerable importance in
discerning intrafamilial relationships of
the Leptodactylidae. This character is
also reflected in the positions of the
atlantal cotyles. Tihen ( 1962a ) cited the
form of the occipital condyles as evi-
dence of a close relationship between
bufonids and the ceratophryine genera.
An essentially heterocoelus condition ob-
tains in Ceratophrys, Caudiverhera, and
LepidohatracJnis — the condyles are not
discrete elements. In the Cycloraninae,
Batrachophrynus, Eupsophus, Hylorina,
Heleophryne, Limnomedusa, Megaelo-
sia, Pleurodema, Proceratophnjs, Tehna-
tohius, and Tehnaiohufo, the occipital
condyles are closely juxtaposed but dis-
crete elements (Fig. 26). In the other
34 Recent leptodactylid genera, the con-
dyles are widely separated. The distinc-
tion between "closely juxtaposed" and
"widely separated" occipital condyles
may be partially size-dependent. The
small species of Leptodactyhis have
widely separated condyles whereas the
larger species (pentadactyhis group)
have the condyles less widely separated
( Fig. 27 ) . The occipital condyles are
widely separated in the largest species of
Eleutlierodactyhis and in the large-
headed And)]yphrynus. These observa-
tions suggest that some taxonomic

LYNCH: LEPTODACTYLOID FROGS

53

Figure 26. Occipital views of skulls of (A)
Ceratophrtjs calcarata (JDL S-237, X 2) and
(B) Cychrana atistralis ( KU 93550, X 4) show-
ing confluent occipital condyles as opposed to
the cond\lar arrangement typical of a type II
cenical cotylar arrangement.

weight may be accorded this character.
In small leptodactylid frogs, both condi-
tions are found; I regard this observation
as additional evidence in support of the
use of this character as a primary tool in
systematics of leptodactylids and other
bufonid frogs.

The condyles are closely juxtaposed
in the primitive frog families, several

Figure 27. Posterior views of skulls of a small
Leptodactylus (A) L. qiiadrivittatus, KU 41030,
X 4, with widely separated occipital condyles
and, a large species, (B) L. pentadactyhts, KU
68159, X 1, with comparatively narrowly sepa-
rated occipital condyles.

leptodactylid genera, Rhinoderma, and
all bufonids. The condyles are widely
separated in all other frogs.

Vertebral Column

All leptodactylids have eight pre-
sacral vertebrae, although the first (cer-
vical ) and second vertebrae are fused in
several genera. In no case is there verte-
bral deletion of presacral vertebrae
( tlirough incoi-poration into the sacrum )
as is known in several bufonids (Noble,
1926b, Tihen, 1960a, 1965). The verte-
brae are procoelus in most leptodacty-
lids, but the majority of the Australo-
Papuan and African genera have free
intervertebral discs (ectochordy) or an
unconsolidated intervertebral tissue (a
paedomorphic trait). The anuran cen-
trum has been the focus of several
lengthy discussions (see Griffiths, 1963,
for a summaiy, and Inger, 1967) and
was one of the primary characters that
Noble ( 1922 ) used in constructing his
anuran phylogeny.

Cervical vertebra. — The cervical and
first thoracic vertebrae are fused in Ade-
lotus, Heleioporus, Heleophryne, Kijar-
raniis, Limnodynastes, NeobatracJuis,
Notaden, and Fhiloria. The vertebrae
are sometimes fused in large specimens
of Ceratophrys and Lepidohatraclms,
but in these genera the fusion is variable
and most often reflects senility.

As noted above, the occipital con-
dyles are either widely separated or
closely juxtaposed. The articular sur-
faces are either separated or confluent.
In those frogs with closely juxtaposed
occipital condyles, it is difficult to de-
termine whether the articular surfaces
are narrowly separated or confluent; this
distinction can be made by examining
the cotylar arrangement of the cervical
vertebra.

The variation in cervical cotylar ar-
rangements permits the definition of
three classes (Fig. 28).

Type I: the cervical cotyles are
widely spaced (this class is the concave
atlas pattern of Gallardo, 1961, 1965).

54

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 28. Three patterns of atlantal cotylar arrangements. (A-C) Dorsal, lateral, and anterior views
of cervical vertebra of Ceratophnjs calcarata (JDL S-237), (D-F) Livwodynastes dorsalis (UMMZ
S-165), and (G-I) Eleiitherodactijlus ntgidostts (JDL S-1238); Types III, II, and I, respectively.

All X 5.

This pattern is the most widespread in
leptodactylids and advanced frogs. In
this class, the cotyles are sometimes
stalked; when they are stalked, there is
a prominent gap between the cervical
vertebra and exoccipital bridge (Fig.
29).

Type II: the cervical cotyles are nar-
rowly separated and there are two dis-
tinct articular surfaces. In many species,
there is a deep notch between the cotyles
( Fig. 28D ) . In most of the frogs with a
type II cervical, the occipital condyles
are clearly discrete articular surfaces.

Type III: the cervical cotyles' are
confluent and represent a single articular
surface. The occipital condyles are con-
fluent and the occipital-cervical articular

pattern is hererocoelus. Only two lepto-
dactylid genera (Ceratophnjs and Lepi-
dohatrachus) have a type III cervical
vertebra. Consistent fusion of the cervi-
cal and second vertebrae is found in
eight Australo-Papuan and African gen-
era with type II cervicals.

Type II cervical vertebrae occur in
AdeJotus, Batrachophninus, Caudiver-
hera, Cijclorana, Eupsophus, Heleiopor-
us, HeJeophnjne, Kijarranus, Limnody-
nastes, Me<^aelosia, Mixopliyes, Neobat-
rachus, Notaden, Philoria, Procerato-
phrys, Tehnatobius, and Tehnatobtifo.
The Oligocene Neoprocoeki has a type
II cervical. The occipital condyles are
very large in Limnomediisa, Odonto-
plirynu.s; and Pleurodema; the cervical

LYNCH: LEPTODACTYLOID FROGS

55

Figure 29. The two occipital-cervical articula-
tion types in the Leptodactylidae. (A) Eleu-
therodacttjhis sulcatus, KU 100355, X 2; and
(B) Cyclorana australis, KU 93550, X 2.

pattern is intermediate between type I
and type II cervical patterns. In all
other leptodactylid genera, the type I
cervical is uniformly present.

The Australo-Papuan Cycloraninae
all have type II cervical vertebrae, and
the Myobatrachinae all have type I cer-
vical vertebrae. One genus of the Elo-
siinae has a type II cervical (Megaelosia),
and the other two genera have type I
cervical vertebrae. The Leptodactylinae
have only type I cervical vertebrae, al-
though two genera (Limnomedusa and
Pleurodema) have an intermediate type
cervical. In the largest leptodactylid
subfamily, the Telmatobiinae, type II
cervicals are found in all of the genera
of the Telmatobiini, in two of the four
genera of the Alsodini, and in one of the
genera of the Odontophrynini. Only

type I cervicals are found in the Grypis-
cini and Eleutherodactylini. The cer-
vical vertebral type seems to be very
useful in ascertaining the relationships
of leptodactylids. The type II cervical
is probably primitive, as evidenced by
the uniform occurrence of this type in
the archaic frog families and Pelobati-
dae. Type II cervical vertebrae are also
characteristic of the Bufonidae and
Rhinodermatidae.

Presacral vertebrae. — Hecht (1960)
pointed out that the primitive frogs have
very short transverse processes on the
posterior presacral vertebrae, and prin-
cipally for this reason he associated
Eorubeta (Eocene) with the advanced
frog families. Zweifel (1956a) demon-
strated that some of the pelobatids of
the subfamily Megophryinae have long
transverse processes of the posterior pre-
sacral vertebrae (as long as the sacral
diapophyses) and very long (here
termed expanded) transverse processes
of the anterior presacral vertebrae. Some
Megophryinae have short posterior pre-
sacral vertebral transverse processes
{LeptobracJuiim, Boulenger, 1908). As
noted by Hecht (1960) the transverse
processes of the posterior presacral
vertebrae of most leptodactylids are as
long as or only slightly shorter than the
sacral diapophyses. However, in sev-
eral of the more primitive leptodactylid
genera, the transverse processes of the
posterior presacral vertebrae are short-
ened to varying degrees (Fig. 30). The
transverse processes of the anterior pre-
sacral vertebrae seem to vary in length
independently of the variation in the
length of the transverse processes of the
posterior presacral vertebrae. The short-
ening of the transverse processes of the
anterior presacral vertebrae and the
concurrent lengthening of the transverse
processes of the posterior presacral verte-
brae represent an evolutionary trend in
frogs. The transverse processes of the
anterior (second, third, and fourth) pre-
sacral vertebrae are widely expanded
(much wider than the width of the

56

MISCELLANEOUS PUBLICATIOX MUSEUM OF NATURAL HISTORY

sacral diapophyses ) in Ceratophnjs and
Lepidohatrachus. The transverse proc-
esses of the anterior presacral vertebrae
are slightly expanded in Cydorana,
Odontophnjmis, Proceratophnjs, and
Telmatobiifo. The transverse processes
of the anterior presacral vertebrae are
not wider than the width of the sacral
vertebrae in all other leptodactylids and
are shortened (their greatest width is
less than the width of the sacral dia-
pophyses) in Crinia and Pseudoplnyne
(Fig. 31).

The transverse processes of the pos-
terior presacral vertebrae are greatly
shortened (knob-like) in Neobatraclms
and Notaden (Fig. 30), very short (less
than one-half the width of the sacral

Figure 30. Posterior vertebral columns of four
genera of cycloranine leptodactylids illustrating
reduction in transverse processes in advanced
Cycloranini, and presence of lateral flanges on
urostyle in primitive genera of subfamily. (A)
Limnodtjnastes tasmaniensis, AMNH 60589, X
5; (B) Cydorana australis, KU 93550, X 2.5;
(C) Hek'iopows eijeri, UMMZ 124504, X 2.5;
and (D) Neobatraclms centralis, KU 93578,
X2.5.

diapophyses) in Crinia, Heleioporus,
LepidobatracJnts, and Pseudophryne,
and moderately to slightly shortened in
Caudiverhera, Ceratophrys, Cydorana,
Cydoramphus, Eiipsophus, Glauertia,
Heleophryne, Myobatradnis, Odonto-
pJirynus, Proceratophrys, and Uperoleia.
In all the other 39 leptodactylid genera,
the transverse processes of the posterior
presacral vertebrae are as wide as the
sacral diapophyses.

CeratopJirys and Lepidobatradnis
have a bony shield overlaying the sec-
ond, third, and fourth vertebrae (Fig.
31). In Ceratophrys, the shield is at-
tached to the neural spines by ligaments.
Among the species of Ceratophrys, there
is little variation in the size and shape of
the shield; some species of the genus
have been reported to lack a dorsal
shield, but I have not seen any adult
Ceratophrys that lack the shield, al-
though most juveniles of Ceratophrys
lack a vertebral shield. In Ceratophrys,
the shield is a loosely consolidated plate
of osteoderms. Based on limited obser-
vations, the number of elements in the
plate is reduced by fusion of adjacent
plates with increasing age. I have not
seen any specimens with only one or
two elements in the shield. Dermostosis
of the shield occurs in both genera. In
Lepidobatradnis laevis, the shield is
small and fused to the neural spines; the
shield is larger in L. asper and L. llanen-
sis and is connected to the vertebrae by
ligaments (Reig, 1960b, Reig and Cei,
1963).

Sacral vertebrae. — The sacro-coccy-
geal articulation is bicondylar in all lep-
todactylids including the African and
Australo-Papuan genera with ectochor-
dal vertebral columns. Several authors
(Noble, 1930, 1931, Schaeffer, 1949)
noted the differences in the dilation of
the sacral vertebral diapophyses. Schaef-
fer (1949) implied that all Australo-
Papuan leptodactylid genera had dilated
sacral diapophyses and all Neotropical
genera had narrow (rounded) sacral
diapophyses. Noble (1931) and Reig

LYNCH: LEPTODACTYLOID FROGS

57

(1960a) pointed out that at least some
of the Neotropical genera characteris-
tically have dilated sacral diapophyses.
The sacral diapophyses are broadly
dilated in all pelobatids, bufonids, and
most hylids. The fossil Oligocene lepto-

dactylid, Neoprocoela, has broadly di-
lated sacral diapophyses, as does the
enigmatic Chilean Rlunodenna. No ex-
tant leptodactylid genus has broadly
dilated sacral diapophyses, although it
is impossible to place lower limits on

Figure 31. Vertebral columns of (A-B) Ccratophnjs aiiiita, A has the bony shield removed ( KU

98129, X 0.9; FMNH 51704, X .9), (C) Lepidohatmchiis asper ( KU 80783, X 2.2), (D) Crinio

slgniferaiKU 56243, X 6.4) and (E) Leptodactyliis pentadactijlus (KU 68159, X 0.9).

58

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

the character of "dilated sacral dia-
pophyses." The sacral diapophyses are
moderately dilated in all of the Australo-
Papuan genera and in eight Neotropical
genera (Batrochylo, Caudiverhera, Cros-
sodactylocJes, Edalorhina, Eupsophus,
Odontophnjnus, PhijsaJaemus, and Fleu-
rodema). Any distinction between the
degree of dilation exhibited by some of
the abox'e-mentioned genera and such
leptodactylids as Ceratophnjs and Fara-
telmatobius is a very fine one and prob-
ably is not defensible. The majority of
the African and Neotropical leptodacty-
lid genera have rounded, undilated
sacral diapophyses which are either per-
pendicular to the sagittal plane or
slanted posteriorly (Fig. 32).

Figure 32. Sacral vertebrae of leptodactylids
and a bufonid illustrating the degree of expan-
sion of the sacral diapophyses. ( A ) Eletithero-
dactylus nigulosus (JDL S-I238, X 4), ( B)
Odontophnjnus cultripes ( KU 92975, X 2.5),
(C) Cijclomna dalili (UMMZ 65250, X 3), and
(D ) Bufo marmoreus ( KU 84894, X 2 ) .

Coccyx. — I have not observed zyga-
pophyseal processes on the anterior por-
tion of the coccyx of any leptodactylid,
although S. B. McDowell (in litt.) has
observed prezygapophyseal processes on
the coccyx of Metacrinia. The coccyx is
short and stocky in several genera; nerve
foramina are evident on the anterolateral
surface of the coccyx in several leptodac-
tylids. Neither of these characters shows
a clear trend but appears sporadically
among the species I have examined.
Batrachoplirynus has large transverse
processes at the anterior end of the coc-

In the Elosiinae, the neural arches
cyx (Fig. 79).

are distinctly non-imbricate and poorly
ossified. The roof of the neural canal of
the coccyx is likewise poorly ossified.

Pectoral Girdle

The primary basis for the divisions of
the advanced frogs by Cope and Boulen-
ger was the variation in the architecture
of the pectoral girdle. The two divisions,
the Arcifera and Firmisterna, differ in
the overlapping epicoracoidal cartilages
in the former and the median fusion of
the epicoracoidal cartilages in the latter.
Noble (1921, 1922, 1931) severely criti-
cized the Cope-Boulenger system after
he found several South American frogs
with intermediate types of pectoral
girdles. Griffiths (1959, 1963) redefined
the terms arciferal and firmisternal on
the basis of the presence (arciferal) or
absence (firmisternal) of free epicora-
coidal horns. Griffiths also pointed out
that the two types of pectoral girdles
have different developmental patterns.

When the primary basis of frog clas-
sification changed from the pectoral
girdle to the vertebral column and tliigh
musculature, relatively few groups
shifted between the Arcifera ( ^Procoe-
la and Bufonoidea) and the Firmisterna
(=Diplasiocoela and Ranoidea). The
Dendrobatidae, GeohatracJws, Rhino-
derma, and Sminthillus (Euparkerella,
Noblella, and SmintliiUus) were alter-
nately associated with the Bufonoidea
and the Ranoidea. Noble (1926b and
1931) considered all of these genera to
be bufonid derivatives, whereas Griffiths
( 1959 ) considered the dendrobatids to
be a Neotropical subfamily of the Rani-
dae, Geobatrachtis and Rhinoderma to
be the only members of a Neotropical
leptodactylid subfamily, and Sminthillus
to be a polyphyletic assemblage of
EleutJwrodactyliis derivatives.

Most leptodactylid frogs are clearly
arciferal (in either Cope's sense or Grif-
fiths' sense). Prior to the refinement of
the distinction between arcifery and
firmisterny, the leptodactylid frogs of
the genera SmintliiUus and Noblella were

LYNCH: LEPTODACTYLOID FROGS

59

characterized by their arcifero-firmister-
nal pectoral girdles. Noble ( 1921, 1922,
1926b, 1931 ) considered the epicoracoid-
al cartilages of SiuinthiUus to be fused
from a point at the base of the omoster-
num to the anterior edge of the coracoid,
whereas Griffiths (1959) found that the
epicoracoidal bridge was restricted to
the anterior-most portion of the carti-
lages and always restricted to a narrow
zone anterior to or between clavicles. I
have found epicoracoidal bridges in both
small and large species, but the fusion is
most often found in the smaller speci-
mens, because one tends to be less me-
ticulous in the dissection of larger speci-
mens. All leptodactyhds have discrete
epicoracoidal horns which are usually
partially concealed by the sternum.
Hoffman (1930) reported Heleophnjne
to pass from arcifery in juveniles and
subadults to firmisterny in adults. Poyn-
ton ( 1964 ) pointed out that Heleophnjne
is always arciferal and that Hoffman's
"old adult Hehophnjne" was a specimen
of a ranid (either Hijhimhates or Lepto-
pelis). All other leptodactyHd genera
are, and have always been, regarded as
arciferal in the sense of Cope and
Boulenger.^

The relative bulk of the clavicle and
coracoid, the presence of an omoster-
num, and the presence of an osseous
sternal ( post-zonal ) element are variable
characters in the pectoral girdles of lep-
todactylid frogs. Piatt (1934) distin-
guished three types of omosterna and
three types of sterna in 35 species of lep-
todactylids (Eleittherodoctyhis, Lepto-
cloctylus, and LitJiodytes) based on the
character states uniformly cartilaginous,
partly calcified, and containing an os-
seous element. I am unwilling to place
much taxonomic weight on small varia-
tions in the size of the omosternum and
sternum, or on the distinction between
uniformly cartilaginous and partly cal-

cified. Calcification of the sternum oc-
curs in large females of several species
in which the sternum of the smaller
male is uniformly cartilaginous. The
larger specimens of a given species are
more likely to have some calcification of
the sternal plate than are the smaller
specimens of the species. In several lep-
todactyHd genera, the differences in the
average adult size of the various species
of that genus are reflected in the pres-
ence and absence of calcification of the
sternum and often of the omosternum
and epicoracoidal cartilages. Although
fully cognizant of the limitations of both,
I consider the following characters to be
useful in distinguishing the genera of
leptodactylids: 1) presence of a carti-
laginous or calcified sternal plate versus
the presence of a discrete osseous ster-
nal element (usually a style) and 2)
presence of an omosternum.

The sternum in most leptodactylids
is a cartilaginous plate (Fig. 33); in the
Neotropical Leptodactylinae (Barycho-
los, EckilorJiina, Hydrolaetare, Leptodac-
tylus, Limnomedtisa, Litljodytes, Fara-
telmatohiiis, Physahemus, Fleurodema,
and Fseudopaludicola) the sternum con-
tains an osseous (or calcified) style or
an osseous plate (Paratelmatobiiis). Litli-
odytes also has an osseous style in the
omosternum; the omosternum is carti-
laginous in all of the other genera.

Parker ( 1940 ) reported that the omo-
sternum was reduced in size or absent
in all Australo-Papuan leptodactylids.
In the Australo-Papuan genera Adelotus,
Cyclorana, Lechriodus, Limnodynastes,
and Mixophyes, the omosternum is large
to moderately large ( Fig. 33 ) . The omo-

' Barrio ( 1970 ) named a new Chilean genus
(Instietophrynus) with a pseudofimiisternal pec-
toral girdle similar to that seen in some Ateloptis,
but having a well-developed prezonal element.

Figure 33. Pectoral girdles of ( A ) Heleophnjne
natalcnsis (KU 105925, X 1.5), (B) Limno-
dynastes peronii ( KU 93562, x 1.2), and (C)
Crinia signifera ( KU 56317, X 4).

60

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

sternum is absent in Mijohatrachus and
reduced in size in the other Australo-
Papuan genera of the Myobatrachinae
and the cycloranine genera Heleioporus,
Kyorranus, Neohatraclnis, Notaden, and
Philoria. The omosternum in Crinia is
usually elongate and not separable from
the anterior extent of the epicoracoidal
cartilages (Fig. 33). The omosternum
is present in all other leptodactylids that
I have examined except Lepidobotm-
dnis, OdontopJirynus, Froceratophnjs,
and EhutherodactyJus ruihae, although
the omosternum is small in several spe-
cies of the West Indian Eleutherodac-
tyhis. The omosternum is unusually
large — often larger than the sternum —

Fk;ure 34. Pectoral girdles of leptodactyline
leptodactylid (A) Pseudopaludicola ameghini
(KU 93050, X 5.5 ) and two telmatohiines ( B)
Tclmatohiiis hautholi ( KU 72879, X 2) and
(C) Eleiitherodacti/lus hogotcnsis ( KU 110409,
X4).

in a few Neotropical leptodactylids
(Ceratophrys and Telmatobufo).

The clavicles are large and strongly
arched in the Ncoti-opical genera Caudi-
verhera, Tehnatobiiis, and Tebnatobufo,
and are very slender and fragile in many
of the smaller species of leptodactylids
( Fig. 34 ) . In most species of the fam-
ily, the clavicles are neither massive nor
fragile. The clavicles are relatively
straight in most Neotropical leptodac-
tylids (except the Ceratophryinae and
Telmatobiini) and many Cycloraninae.
In most myobatrachines the clavicles are
slender and strongly arched (Fig. 33)
but in Myobatrachus the clavicles are
massive and only slightly arched.

The following generalizations con-
cerning the architecture of the leptodac-
tylid pectoral girdle are suggested: 1)
arcifery is the primitive character state
— pseudofirmisterny is derived and of
little taxonomic significance; 2) an osse-
ous element in either the pre-zonal or
post-zonal element is secondary to uni-
formly cartilaginous elements; calcifica-
tion of the sternal plate is a labile fea-
ture; and 3) primitively, the omosternum
is large — reduction in size or loss of the
omosternum is derived.

The osseous post-zonal elements of
leptodactylines are primitively broad
and become more style-like in the ad-
vanced genera (Fig. 35). Posterior bi-
furcation of the style is derived.

Pelvic Girdle

The pectoral girdle has long been
recognized as an important complex for
systematic studies of anurans whereas
few authors sought to find characters in
the simpler pelvic girdle, which is more
closely associated with the primary
adaptive shift of frogs — jumping. The
anuran pelvis is composed of three ele-
ments (ilium, ischium, and pubis) of
which only the ilium exhibits familial or
consistent generic variation. Some pale-
ontologists (e.g., Auffenberg, Chantell,
Estes, Holman, and Tihen) considered
the ilium of paramount importance in

LYNCH: LEPTODACTYLOID FROGS

61

Figure 35. Pectoral girdles of four leptodacty-
line leptodactylids. (A) Phtj.salaeiniis ])ustuIosiis
(KU 68272, X 5.5), (B) Leptodactyliis mcla-
nonotus ( KU 68276, X 3), (C) Physalaemus
signifcnis (KU 93033, X 5.5), and (D) Para-
telmatohiiis hitzi ( KU 107089, X 5.5).

the identification of disarticulated frog
fossils at the familial, generic, and, in
some cases, specific level. Brief refer-
ences were made by some of these
authors to the range of variation ex-
hibited by the leptodactylid genera.

Holman (1965) noted the variation
in development of the ilial crest and in
the size and shape of the ilial promi-
nence (termed by him, vastus promi-
nence). The nomenclature for the ilial
morphology adopted here follows that of
Chantell (1964) and is illustrated for lep-
todactylid frogs in Figure 36. The only
additional term employed here is the
"preacetabular zone" — that area of the
ventral acetabular expansion lying anter-
ior to the lip of the acetabulum and dor-
sad to the ventral lip of the acetabulum.

Owing to the diverse forms of the
ilium in leptodactylids, no familial diag-

nosis can be made at this time. The rela-
tively few genera available support this
conclusion. Skeletons of the following
genera were available: Adelotus, Barij-
chohs, BatracJwpJinjmis, Caudiverbera,
Ceratophrys, Crossodactylus, Cyclorain-
phus, Cyclorana, Edalorhina, Eleuthero-
dactylus, Eupsophus, Heleioporus, Hy-
lactophryne, Hylodes, Isclinocnema,
Kyarramis, Lechroidus, Lepidobatra-
chus, Leptodactyliis, Limnodynastes,
Limnomedusa, Megaelosia, Neobatra-
cJiiis, Notaden, Odontophrynus, PJujsa-
laemiis, Pleiirodema, Proceratophrys,
Pseiidophryne, Syrrliophus, Tebnutobius,
Thoropa, Tomodactyhis, and Zachaenus.

The two ilial types are briefly de-
scribed below.

Ceratopliryine type: ilial shaft rela-
tively short; dorsal crest not developed,
if present, in form of ilial shaft ridge;
dorsal protuberance not, or but slightly,
differentiated from spike-like dorsal
prominence; dorsal acetabular expansion
well developed, distinct dorsal vector;
preacetabular zone present, broad; ven-
tral acetabular expansion relatively

Figure 36. Nomenclature of the leptodactylid
ilium. Top is ilium of Cerafophnjs ornata (genus
lacking dorsal crest); middle is that of Lepto-
dactijlus pentadactyhis (genus with dorsal crest).
(A ) dorsal acetabular expansion; ( B ) dorsal pro-
tuberance (vastus prominence of Holman); (C)
dorsal prominence; ( D ) acetabular fossa; (E)
ventral acetabular expansion; (F) shaft; (G)
dorsal crest. The preacetabular zone is hatched
in the lower figure.

62

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

small; angle of ventral acetabular ex-
pansion greater than 90°; acetabular
fossa relatively small (Fig. 37).

Leptodactyline type: ilial shaft rela-
tively long; dorsal crest well developed,
usually higher anteriorly and in the re-
gion of dorsal protuberance; dorsal pro-
tuberance relatively large [vastus prom-
inence of Holman ( 1965 ) ] , ovoid, or
tear-drop-shaped, situated on a large
dorsal prominence which is confluent
with dorsal crest; posterior border of
dorsal prominence gently sloping; dorsal
acetabular expansion not well developed,
little or no dorsal vector; preacetabular
zone present, broad; ventral acetabular
expansion relatively large; angle of ven-
tral acetabular expansion much less than
90°; acetabular fossa relatively large
(Fig. 37).

Figure 37. Right ilia of (A) Pseudophryne
hihroni, AMNH 64511, X 5.4; (B) Adelotiis
hrevis, AMNH 59489, xL6; (C) Cydorana
(lustralis, AMNH 62228, X 1.2; (D) Liiuno-
dynastes dorsalis, KU 93553, X 1.6; (E) left
ilium of Neohatmchus centralis, KU 93578, X
1.6; and (F) right ilium of Notaden hennetti,
FMNH 97858, X 1.2.

The ilia of the two genera of the
Ceratophryinae are nearly impossible to
separate. Lepidohatrachus has a more
distinct ilial shaft ridge than does Cera-
tophnjs and a slightly less pronounced
dorsal prominence (spike); too few
specimens are available to make a gen-
eralization. Odontophnjmis and Procera-
tophrys ( Telmatobiinae, Odontophryn-
ini) are the only other Neotropical gen-
era examined which have the cerato-
phryine pattern. Both genera differ

from the Ceratophryinae in having a
more distinct dorsal protuberance, larger
ventral acetabular expansion, and less
obtuse angle of ventral acetabular ex-
pansion (Fig. 38).

Several cycloranine genera have ilia
which differ only slightly from the cera-
tophryine pattern. All Australo-Papuan
genera examined have a relatively large
angle of ventral acetabular expansion
(90-110°) and relatively small acetabu-
lar fossa. The dorsal prominence is
never spike-like and most genera have a
distinguishable dorsal protuberance
(e.g., Adelotiis, HeJeioponis, Lecliriodus,
and 'Neohatrachus) . The dorsal protu-
berances of Kijarranus and Limnody-
nastes are only slightly less differen-
tiated, but those of Cydorana, Notaden
and the myobatrachines are not distin-
guishable from the dorsal prominence.
Dorsal crests are lacking in the myo-
batrachines, and all cycloranines exam-
ined except Adelotiis, Kyarranus, and
Lecliriodus (Fig. 38).

The angle of ventral acetabular ex-
pansion is slightly greater than 90° in
Caiidiverhera, Edalorliina, Eleutliero-
dactylus, Engystomops, Eupsoplnis, Hy-
lactophryne, Limnomedtisa, Pleiirodenia,
Syrrhoplnis, Tomodactylus, and Zachae-
nus, whereas it is markedly acute in
Crossodactylus, Cycloramphus, Hylodes,
Leptodactylus, Megaelosia, and Tlioropa.
In Batrachophryniis, Ischnocnema, and
Telinatobhis it is about 90°.

The dorsal crest is absent in Edalo-
rliina, Eupsophus, Physalaemus, and
Pleurodema. In some Eletitlierodactyltis
and the Mexican Syrrhoplnis and Tomo-
dactylus, the dorsal crest is reduced to
a small area immediately anterior to the
dorsal prominence. This latter condition
is also seen in Limnomedusa. The dor-
sal crest is well developed in Caiidiver-
hera, Crossodactylus, Cycloramphus,
most Eleiitherodactylus, Hyhctophryne,
Hylodes, Ischnocnema, Leptodactylus,
Megaelosia, and Zachaenns; in Batra-
chophrynus, Telmatobius, and Thoropa

LYNCH: LEPTODACTYLOID FROGS

63

Figure 38. Right ilia of (A) Hijlodes nasus,
KU 92894, X 2.1; (B) Thoropa miliaris, KU
92856, X 1.6; (C) Eupsophus wseus, AMNH
22104, X 1.6; and (D) Odontophnjnus cultripes,
KU 92975, X 1.6.

it is low and almost ridge-like (Figs.
37-40).

The dorsal protuberance is well de-
veloped in all elosiine, leptodactyline,
and telmatobiine frogs examined except
Megaelosia and Pleuwdema. In these
J latter two genera, the muscle scar for the
vastus externus head of the m. triceps
jemoris is broad and covers the whole
of the dorsal prominence. The protuber-
ance is ovoid, round, or elongate, and no
characterization of genera can be made
owing to the variability of the protuber-
ance in Eleutherodactylus.

The degree of dorsal vector in the
dorsal acetabular expansion shows some
variation in the Neotropical leptodacty-
lids. Almost no dorsal vector is seen in
Batrachophrymis, Edalorliina, Leptodac-
tylus, Pleuwdema, Telmatobius, and
Thoropa, whereas the other genera ex-
hibit a considerable dorsal vector. Eleu-

FiGURE 39. Right iha of ( A ) Eleutherodactylus
diastema, JDL S-244, X 6.6; (B) £. inoptatus,
AS X2356, X 1.2; (C) E. nigulosus fleisch-
manni, KU 68157, X 1.2; (D) E. palmed, JDL
S-242, X 3.3; and (E) Ctjcloratiiphus fulginosus,
KU 92790, X 1.2.

therodactylus palmeri (Fig. 39), in con-
trast to the other species of the genus
examined, has no dorsal vector and the
dorsal crest appears to be lacking but is
present and curved over the body of the
ilial shaft.

The ilium seems to have some merit
in suggesting relationships among the
leptodactylid genera, but at this time
inadequate data are available in order to
incorporate the variation of the ilium
into my generic classification. In some
cases ( Ceratophryinae and Elosiinae),
the ilial variation is consistent with the
\'ariations of other character complexes,
but in other cases (Leptodactylinae),

Figure 40. Right ilia of (A) Phijsalaemus
pustulosus, KU 41031, X 5.4; (B) Pleuwdema
cinerea, KU 80836, X 1.6; (C) Leptodactylus
wagneri, KU 104390, X 1.2; (D) Caudiverhera
caudiverbera, A.M.N.H. No. 51510, X 0.6; and
(E) Batrachophrymis macrostomus, KU 98127,
XO.6.

the variation is not consistent with my
other data. Owing to these discordan-
cies, use of the ilium in a leptodactylid
classification is deferred pending a
broader survey of the variation in the
ilia of other leptodactyhd frogs.

The data now available are adequate
to permit a reassessment of the status of
a Lower Miocene Leptodactylus recently
described from Florida. Holman (1965)
named L. abavus based on five ilia from
the Arikareean horizon of Thomas Farm,
Gilchrist County, Florida. The frog he
described and figured has almost no pre-
acetabular zone, a moderately large
angle of ventral acetabular expansion,
and a small ventral acetabular expan-

64

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

sion. These characteristics are not t>'pi-
cal of Leptodoctylus but are typical of
Rana — especially the species of the R.
pipiens group. There is a marked simi-
larity between his figures of the ilia of
Leptodactyhis abaviis, Rana miocenica,
and Rana cf. R. pipiens. Leptodactyhis
ahaviis Holman, 1965, is accordingly here
removed from the family Leptodactyli-
dae and placed in the Ranidae. The
specific status of Rana abavus (Hol-
man), new combination, is beyond the
scope of this paper, but at present, there
are no apparent differences between it
and Rana miocenica, a sympatrant.

Limb Elements

To a certain extent, the mode of life
of a frog is reflected in the proportions
of the long bones (humerus, radioulna,
femur, tibiofibula, and astragulus and
calcaneum). Species living in paramos
and those with burrowing habits tend to
have heavier, stockier long bones than
do the arboreal or active terrestrial frogs,
in which the limb bones are relatively
slender. Those differences that do exist
are subtle and offer little in the way of
taxonomic utility. Because the charac-
ters seem more reasonably associated
with ecological adaptation, they are not
regarded as useful in a phylogenetic
classification. The intrageneric variation
in EJeutherodactyhis precludes the use of
relative proportions of the limb elements
for generic or suprageneric classifica-
tions.

The greatly enlarged arms of males
of some species of Leptodactyhis have
been noted by several authors. The
skeletal base for the sexual dimorphism
is considerably more striking. Large
flanges are present on the humeri of the
male, whereas the female has humeri
more typical of frogs (Fig. 41). Devel-
opment of humeral flanges is uncommon
in the Leptodactylidae, and is most pro-
nounced in the genus Leptodactyhis.
The development of humeral flanges in
LeptodactyJus appears to be species-
group variable and therefore may prove

useful in subgeneric classifications. Mod-
erate flanges are seen in the Eupsophus
nodosus group, again restricted to the
male (Grandison, 1961), in Crossodac-
tylodes, Paratehnatobius, some PJiysa-
laemus, and Telmatobius (Fig. 41).

Figure 41. Humeri of (A) female Leptodactyhis
pentadactijhis ( lateral view of right humerus,
KU 68159, X 0.8) and (B) male L. pentadacty-
lus (lateral view of left humerus, KU 84981, X
0.8); medial views of left humeri of male (C)
Physalacmiis signiferiis ( KU 93033, X 4.8),
(D) P. awieri (KU 92999, X 4.8), and (E)
Paratehnatobius httzi (KU 107089, X 4.8).

Thalan<:,eal formulae. — The primitive
phalangeal formulae of the Leptodactyli-
dae are 2-2-3-3 for the hand and 2-2-3-4-3
for the foot. Phalangeal reduction is
often regarded as a character sui generis
in amphibian classifications, because it
is a relatively uncommon phenomenon.
The only variations on the above for-
mulae in the Leptodactylidae are seen in
Etiparkerella which has a 2-2-3-2 formula
for the hands and a normal foot formula
(Figs. 42-43) and in Limnodynastes
peronii which has a 1-2-3-3 hand for-

LYNCH: LEPTODACTYLOID FROGS

65

inula (all other Limnodipiastes have a
2-2-3-3). Parker (1940) and Moore
(1961) briefly mentioned that the digits
were reduced in some of the myobatra-
chine leptodactylids (Crinia and Psetido-
phrijne). In external appearance, the
digits are short and stubby, but the
phalangeal formulae are typical for bu-
fonoid frogs (2-2-3-3, 2-2-3-4-3).

Terminal PJialanges. — The nature of
the terminal phalanges has been one of
the major character complexes in lepto-
dactylid classification. Generic diag-
noses invariably contain reference to
"simple, pointed, knobbed or T-shaped"
terminal phalanges.

Based on a literature study, there
seems to be a wide range of terminal
phalangeal types within the Leptodactyl-
idae. The phalangeal types are termed
pointed, simple, knobbed, bifurcate, Y-
shaped, and T-shaped. In the past, the
technique for observing the terminal
phalanges was to dissect the digital tip
and observe the shape of the bone. Since
the terminal phalanges are often very
small and delicate, dissection tends to
break any lateral expansions. Observa-
tions of the bones with the naked eye
often fails to discern small lateral proc-
esses. In my opinion, the only technique
acceptable for the observation of the
terminal phalanges is examination of
macerated or cleared and stained speci-
mens. My studies, presented below, are
based almost exclusively on examination
of cleared and stained material.

No leptodactylid normally has
pointed terminal phalanges. The most
simple phalanges are very slightly
knobbed (e.g., Limnodynastes and the
Pseudopaludicola falcipes complex). A
graded series can be demonstrated rang-
ing from P. falcipes to some of the ar-
boreal Eleutherodactylus with widely
expanded, delicate, T-shaped terminal
phalanges (Figs. 42-44).

The following genera exhibit definite
T-shaped terminal phalanges in all spe-
cies: Batrachyla, Crossodactylodes (see
Fig, 44, Y-shaped), Crossodactylus, Hel-

FiGURE 42. Palmer views of the skeletons of the
hands of (A) Eletitherodactylus brevicrus, KU
108982, X 10.5; (B) Euparkerella brasiliensis,
KU 93192, X 21); and (C) Eleittherodactijlus
binotatus, KU 92813, X 7.

eophryne, Hylodes, Lithodytes, Megae-
losia, Sininthillus, Syrrhoplnis, Taudac-
fyhts, Thoropa, and Tomodactijlus. Mod-
ified T-shaped terminal phalanges are
seen in Euparkerella (Figs. 42-43).
Barycholos and the Leptodactylus mar-
moratiis group have terminal phalanges
with moderately long lateral processes.
In the genus Eleutherodactylus there is
a continuum of variation in the terminal
phalanges from those of E. binotatus and
E. nigrovittatus (knob-like) to that ex-
emplified by E. brevicrus (broadly T-
shaped). The phalangeal type of all
other leptodactylids, except the Pseudo-
paludicola falcipes group and Limnody-
nastes (Fig. 45), is included in the bot-
tom of the Eleutherodactylus series [for
example, Holoaden, Leptodactylus (ex-
cept marmoratus group), Niceforonia,

66

MISCELLANEOUS PUBLICATIOxX MUSEUM OF NATURAL HISTORY

Figure 43. Plantar views of skeletons of feet of

(A) Eleutherodactyhis hogotensis, KU 110409,

X 7; and ( B ) Euparkerella brasiliensis, KU

93192, X 21.

Paratelmatohius, PseudopJiryne, Telma-
tohius, and Zachaenus (Fig. 44)].

Use of the nature of the terminal
phalanges is not invalidated by the above
data. Even when the lateral processes of
the terminal phalanges are reduced (as
in E. hinotatus and E. nigrovittatus) , the
eleutherodactyline digital groove is evi-
dent externally. The lateral expansions
of the terminal phalanges of the hands in
£. nigrovittatus, Holoaden bradei, and
Niceforonia festae are comparable, but
there are true T-shaped terminal pha-
langes in Eleutlierodactylus nigrovittatus
but merely the lateral expansion of
knobbed terminal phalanges in Holoa-
den and Niceforonia (Fig. 45). Further-
more, in Eleutherodactylus nigrovittatus,
at the tip of each digit (except the
thumb and inner toe) there is a trans-
verse terminal groove. The degree of
development of lateral expansions varies

on the different digits of the hand and
foot as well as between the hand and
foot. When reduction of lateral expan-
sions occurs, it is most readily observed
on the hand; the toes retain relatively
well developed lateral expansions. The
lateral expansions of the inner digits
show greater reduction than do those of
the outer digits.

PrepoUex and inner metacarpal. — The
presence of nuptial excrescences, asperi-
ties, or spines serves as an external in-
dicator of some modification of the first
metacarpal and /or expansion or modifi-
cation of the prepollex. These changes

F

Si

a

H

Figure 44. Third fingers of representative lep-
todactylids (A) Eleutherodactylus nigrovittatus
(USNM GOV 8108, x 17), (B) £. parvus ( KU
92834, x 17), (C) Holoaden bradei (KU 107088,
X 8.5), (D) Niceforonia festae (KU 118137, X
17), (E) Tehnatohius hautholi ( KU 72879, X
4.2), (F) Leptodacttjlus melanonotus ( KU
68275, X 5.6), (G) Pseudophryne hibroni (KU
93588, X 8.5), (H) Liiiinodyiiastes peronii (KU
93566, X 8.5), (I) Pseudopaludicola pusilla
(UMMZ 54589, X 17), (J) P. ameghini (KU
93050, X 17), (K) Zachaenus parvulus ( KU
107090, X 8.5), (L) Paratelmatohius lutzi ( KU
107089, X 8.5), and (M) Crossodactylodes
pintoi (USNM 102611, X 17).

LYNCH: LEPTODACTYLOID FROGS

67

Figure 45. Skeletons of hand of (A) Eleuthero-
dactijhis niprovittatits (USNM GOV 8108) and
feet of (B) Niceforonia fcstae ( KU 118137)
and (G) Eleutherodactiilus iiifiwvittattis (USNM
GOV 8108). All X 10.5.

are not manifest in the female and
while perhaps of some phylogenetic
importance are not employed here. Pre-
pollices are absent or greatly reduced in
size in those genera and species which
lack nuptial asperities or swelling of the
thumb. In some genera (e.g., Telmato-
hius), the prepollex is greatly enlarged
(Fig. 46), whereas in others with simi-
lar nuptial excrescences (e.g., Physalae-
mus), the inner metacarpal is modified
for support. In those genera with large,

isolated nuptial spines ( e.g., Heleioporus
and Leptodactylus) , the inner spine is
prepollical and the outer metacarpal
(Fig. 46).

Figure 46. Skeletons of first finger (thumb) and
prepollices of male leptodactylids. (A) Lepto-
dactylus pentadactyhis (KU 84981, X 1.2), (B)
L. melanonotiis (KU 68275, X 4.8), (G) Telma-
tohitis hatitholi (KU 72879, X 3.6), (D) Paratel-
matohius hitzi (KU 107089, x 15), and (E)
Physalaemus signifems (KU 93033, X 7).

SYSTEMATIC ACCOUNTS

In the following accounts those char-
acters which are only diagnostic at the
subfamily level are listed in the subfam-
ily accounts but not in the generic ac-
counts. I consider it desirable to use the
same sequence of character statements
in all of the generic accounts, and have
numbered all character or character com-
plexes consecutively throughout the fol-
lowing accounts. In some cases, where

the state of a character is not known for
a genus, the number is retained in the
diagnosis, but no statement follows it.
The following characters and character
complexes were used to diagnose the
subfamilies, tribes, and genera recog-
nized herein: 1) sternum cartilaginous,
or bearing an osseous plate or style; 2) a
dermostosed vertebral shield is present
or absent; 3) the transverse processes of

68

MISCELLANEOUS PUBLICATION' MUSEUM OF NATURAL HISTORY

the anterior presacral vertebrae are ex-
panded or not; 4) the arrangement of
the cervical cotyles; 5) cervical and sec-
ond vertebrae free or fused; 6) cranial
bones involved in dermostosis or not;
7) omosternum present or absent, large
or small; 8) degree of dilation of sacral
diapophyses; 9) maxillary arch toothed
or not, teeth pointed or blunt, pedicel-
late or not; 10) direction of and width
at the base of the alary processes of the
premaxillae; 11) shape of the palatal
shelf of the premaxilla; 12) size of facial
lobe of maxilla; 13) shape of the palatal
shelf of the maxilla, development of
pterygoid process of maxilla; 14) maxil-
lary arch complete or not, if not, quad-
ratojugal present or absent; 15) size of
nasals and degree of median contact be-
tween nasals; 16) contact of nasals and
maxillae, nasals and pterygoids; 17) con-
tact of nasals and frontoparietals; 18)
frontoparietal fontanelle — present or ab-
sent; 19) ornamentation of frontopari-
etals; 20) fusion of frontoparietal and
prootic; 21) temporal arcade present or
not; if present, temporal fenestra present
or not and temporal notch present or
not; 22) development of epiotic emi-
nences; 2.3) shape of cristae paroticae;
24) size of zygomatic ramus of squa-
mosal; 25) size of otic ramus of squamo-
sal, development of otic plate; 26) size
of squamosal-maxillary angle; 27) colu-
mella present or absent; 28) prevomers
toothed or not, in median contact or not,
and position of dentigerous processes
relative to choanae; 29) median and
lateral extent of palatines, development
of odontoids; 30) sphenethmoid divided
or entire, shape and extent anteriorly;
31) shape of anterior ramus of para-
sphenoid, keeled medially or not; 32)
degree of overlap of alae of parasphe-
noid and median rami of pterygoids, and
posterior deflection of parasphenoid
alae; 33) relative bulk of pterygoid,
length of anterior ramus and median
ramus; 34) size, shape, and arrangement
of occipital condyles; 35) development

of mandibular odontoids; 36) shape of
terminal phalanges; 37) size and shape
of alary processes of hyoid plate; 38) cri-
coid cartilage divided venti-ally or not;
39) patterns of insertion of m. petro-
liyokleiis anterior and r?j. sternolujoideiis
on hyoid plate; 40) single or double slip
of m. depressor mandibulae; 41) pupil
shape — horizontal or vertical; 42) nup-
tial asperities on thumb and /or chest or
not; type of vocal sac; 43) development
of major body glands — parotoid, flank,
lumbar, and inguinal; 44) tongue shape;

45) webbing of toes, shape of digital
tips, development of tarsal folds, and
presence of an outer metatarsal tubercle;

46) larval morphology — vent median or
dextral, tooth rows, and development of
labial papillae; 47) amplectic position;
48) egg deposition — foam nest, in water,
or terrestrial situation; 49) snout- vent
lengths of adults — abbreviated SVL; 50)
tympanum visible, concealed, or absent;
51) any unique characters that are es-
pecially diagnostic of the genus.

Characters 1), 2), and 3) do not oc-
cur in any generic account and charac-
ters 5), 6), 14), 21), and 39) occur in
only some generic accounts. Character
51) is used only when appropriate.

GENERA EXCLUDED FROM
THE LEPTODACTYLIDAE

As a result of my reassessment of the
variation of many characters used in the
classification of the Leptodactylidae,
tliree genera were found to belong to
other families. Geobatraclius is removed
from the Leptodactylidae primarily on
the basis of discussions with Charles F.
Walker concerning the systematic posi-
tion of the genus; Hylopsis, long an enig-
matic Neotropical frog genus, is a hylid
and probably inseparable from Hijh;
and Wiinoderma is excluded from the
Leptodactylidae because it exhibits no
close similarity to any of the 57 lepto-
dactylid genera, but does exhibit some
similarity to the endemic South Ameri-
can bufonid genera.

LYNCH: LEPTODACTYLOID FROGS

69

Geobatrachus Ruthven, 1915

Ruthven (1915) named Geobatrachus
walkeri as a new genus and species of
the Dendrobatidae. The species remains
rare and is apparently endemic to the
Santa Marta Mountains of Colombia.
Noble (1931) considered Geobatrachus
most closely allied to Rhinoderma and
SminthiUiis. Griffiths (1959) demon-
strated that the species of SminthiUus
are leptodactylids. He placed Geoba-
trachus and Rhinoderma in the Rhino-
dermatinae, a subfamily of the Lepto-
dactylidae; the two genera were asso-
ciated together because of their common
pseudofirmisterny and edentulousness.

Dr. Charles F. Walker is currently
engaged in a study of the relationships
of Geobatrachus and does not consider
it to be a leptodactylid genus. Teeth are
present, the pectoral girdle is firmister-
nal, the sacral diapophyses rounded, the
tarsal bones fused, and the anterior zonal
elements of the pectoral girdle are
greatly reduced or possibly absent. Geo-
batrachus is apparently a microhylid

genus.

Hylopsis Werner, 1894

Werner (1894) named Hylopsis
phtycephalus as a new genus and spe-
cies of the Cystignathidae. The type-
locality is "South America." Werner's
meager description of the holotype does
not provide many clues as to its relation-
ships but the following data are of sig-
nificance in determining the status of
Hylopsis platyceplmlus: pectoral girdle
arciferal (inferred from the family as-
signment; Werner must have examined
the pectoral girdle because he com-
ments on the omosternum ) , omosternum
cartilaginous (therefore present), pupil
horizontal, large digital pads on hands
and feet, fingers one-half webbed, and
toes fully webbed.

The webbing pattern described by
Werner is unique if Hylopsis is a lepto-
dactylid; no leptodactylid now known
has webbing between the fingers, and

very few species have fully webbed feet.
The webbing pattern is not unique in
the Centrolenidae, Hylidae, or Pseudi-
dae. Pseudids do not have digital pads
whereas most centrolenids and hylids do.
Centrolene, Centrolenelh, Hyla (part),
Osteocephalus, PJirynohyas, Smilisca,
and Sphaenorhynchus have webbing pat-
terns that are like that described for
Hylopsis. All have horizontal pupils, but
Centrolene and Centrolenella lack an
omosternum. Werner reported the pre-
vomerine teeth to be absent but did not
give the size or sex of the unique holo-
type. If prevomerine tooth patches were
absent and the specimen was half grown
or an adult, then it is unlikely that Hy-
lopsis could be associated with Osteo-
cephalus, Phrynohyas, or Smilisca. Wer-
ner's description of the holotype suggests
that the frog had a very different snout
shape from that of Sphaenorhynchus.
Hylopsis cannot be distinguished from
Hyla, although this could be an artifact
of Werner's description. If the genus is
a bufonoid genus, then it must be close
to or identical with Hyla (assuming
Werner correctly presented the charac-
ters of the holotype).

Rhinoderma Dumeril and Bibron, 1841

Rhinoderma was often placed in its
own family or subfamily by pre-Noble
systematists. Mivart (1869) and Bou-
lenger (1882) placed it in the Engys-
tomatidae. Noble ( 1931 ) argued that it
was a brachycephalid — a heterogeneous
Neotropical family group of derived bu-
fonids. Davis (1935) and Griffiths
(1954) argued that if the three brachy-
cephalid subfamilies recognized by
Noble were independently evolved from
bufonids, then the Atelopodinae, Den-
drobatinae, and Rhinodermatinae were
families of frogs, not subfamilies.

Griffiths (1959) studied the Brachy-
cephaHdae and placed the included gen-
era in four famihes: Atelopodidae (Ate-
lopus and Brachycephalus), Bufonidae
(Cacophryne, Demlrophryniscus, Didy-
namipus, and Oreophrynellu), Leptodac-

70

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

tylidae (Geohatrachus, Nohlellu, Rhino-
derma, and Sminthilhis), and Ranidae
(Dendrohates, PhijUohates, and Prosther-
apis). The three ranid genera were
placed in the subfamily Dendrobatinae.
The leptodactylid genera were placed in
the Rhinodermatinae (Geohatrachus and
Rhinoderma) and Sminthillus divided
into three genera — Euparkerella, No-
blella, and Sminthilhis, which were all
considered very closely related to Eleti-
therodactyhis.

The breeding behavior of Rhinoder-
ma darwini is unique among amphib-
ians. Amplexus is axillary. Like many
other frogs, the eggs are large (3-6 mm.
in diameter), few in number (20-40),
and pigmented. The eggs are laid in a
sheltered terrestrial situation and are
guarded by the male. After hatching,
the larvae complete their development
in the hypertrophied vocal sacs of the

Figure 47. Dorsal and lateral views of the skull

of Rhinoderma danoini ( KU 68685, X 5.5).

The anterior outline of the head is indicated by

the dashed lines.

male. The mouthparts are modified; the
tooth rows are 1/2 and a beak is present;
the labial papillae are interrupted an-
teriorly, and the vent is median.

The skull of Rhinoderma is not
heavily ossified (Fig. 47) except pos-
teriorly. The occipital condyles are
broad and close together and the cer-
vical cotylar arrangement is type II.
Teeth are lacking, the palatines and pre-
\'omers are lacking, the maxillary arch is
incomplete, but the quadratojugal is
present. The columella is present, the
squamosal-maxillary angle is about 55°,
and no otic plate is developed on the
squamosal. The m. depressor mandilm-
lae consists of the pars tympanicus only.

There are eight presacral vertebrae,
all procoelus, and the cervical and sec-
ond vertebrae are fused. The sacral dia-
pophyses are broadly dilated, and the
sacrum articulates with the coccyx by a
bicondylar articulation. The pectoral
girdle is pseudofirmisternal with free
epicoracoidal horns and an omosternum
and sternum are present and cartilagi-
nous. The organ of Bidder is lacking.
The terminal phalanges are knobbed.
Rhinoderma lacks outer metatarsal tu-
bercles.

Rhinoderma is clearly not related to
the Dendrobatidae, as claimed by Cei
( 1962a ) , nor is it closely related to any
other Neotropical gioup. It differs least
from the endemic South American Bu-
fonidae (several genera referred to the
Atelopodidae or Bufonidae by various
authors), but is clearly not a bufonid,
because it lacks an otic plate on the
squamosal, organ of Bidder, outer meta-
tarsal tubercle, and has a well developed
omosternum. The family Leptodactyli-
dae is sufficiently loosely defined that
the inclusion of Rhinoderma would not
appreciably expand the family definition.
If Rhinoderma is included in the Lepto-
dactylidae, I would recognize it as the
sole member of a subfamily. However, I
anticipate that when a more thorough
study of the Neotropical bufonids is
made, the differences between Rhino-

LYNCH: LEPTODACTYLOID FROGS

71

derma and the Bufonidae will be in part
demonstrated to be corollaries of the
reduction in ossification in Rhinoderma.
I prefer to recognize the Rhinodermati-
dae for this enigmatic Chilean genus and
define the family as follows: bufonoid
frogs with a pseudofirmisternal pectoral
girdle, cartilaginous pre-zonal and post-
zonal elements, lacking an organ of Bid-
der, palatines, prevomers, teeth, an otic
plate on the squamosal, having broadly
dilated sacral diapophyses, and a unique
life history among frogs — tadpoles de-
velop in the male vocal sacs.

KEYS TO THE GENERA OF
LEPTODACTYLID FROGS

The purpose of a key should ulti-
mately be for rapid and accurate iden-
tification of a specimen, regardless of
what information accompanies it. Keys
are especially useful in the identification
by non-herpetologists of a specimen with
no hint of its provenance. It is very dif-
ficult to prepare an effective key to a
group of organisms when one places as
a primary limitation that the characters
used in that key will be characters that
are readily accessible and/or superficial.
In the family Leptodactylidae, as in most
other groups, the members on different
continents have undergone parallel mod-
ifications in adapting to their environ-
ment or to a particular mode of life.
Therefore it is almost impossible to write
a key, based on external (or super-
ficially available internal) characters, to
the Leptodactylidae on a world-wide
level. Thus the following keys are 1) a
key to the African and Australian genera
and 2) a key to the African and Ameri-
can genera. Heleophrijne, the sole Afri-
can genus, has been included in both
keys in an attempt to make the keys
more cosmopolitan.

As superficial characters I include
several characters readily visible once
the skin in the tympanic region has been
reflected; the presence or absence of a
tympanic membrane can be observed as
can the nature of the musculus depressor

mamlibulae. Most of the other charac-
ters used are those that traditionally
have been used in keys.

Key to African and Australo-
Papuan Leptodactylids

Two subfamilies of leptodactylids oc-
cur in Australia and are not readily sep-
arated by any "key" external character.
Osteologically, they separate readily, but
since my intent in preparing the keys
was for utility in identifying preserved
specimens, I have avoided using all but
easily accessible osteological characters.
All cycloranines possess prevomerine
dentigerous processes and these proc-
esses are absent in all myobatrachines
except for some of the species of Crinia.
Outer metatarsal tubercles are absent in
almost all cycloranines and present in
all myobatrachines except some Crinia
and Taudactyhis. With the exception of
the microcephalic Mijohatrachus, all
myobati-achines are very much smaller
than any of the cycloranine leptodacty-
lids. Alosolute size however is a very
poor key character.

1. M. depressor mandihidae in a single

slip (par tympanicus) 2

M. depressor mandihulae in two
slips 4

2. Tympanic annulus absent

Pseudophryne

Tympanic annulus present 3

3. Large parotoid glands present

Philoria

Parotoid glands absent Crinia

4. Prevomerine teeth absent 5

Prevomerine teeth present 9

5. Outer metatarsal tubercle present .—
6

Outer metatarsal tubercle absent _.-
Taiidactylus

6. Anal flap present; parotoid and flank
glands present; pupil vertical in life
Uperoleia

72

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Anal flap absent; parotoid glands
absent or poorly defined; flank glands
absent; pupil horizontal 7

7. Toes webbed Glauertia

Toes devoid of webbing 8

8. Head minute compared to body

Myobatrachus

Head of normal proportions

Metacrinia

9. Maxillary teeth absent Notaden

Maxillary teeth present 10

10. Prevomerine dentigerous processes
between choanae 11

Prevomerine dentigerous processes
posterior to choanae 14

11. First finger opposable to others

Cyclorana

First fiinger not opposable to others
12

12. Outer metatarsal separated by web
Mixophyes

Outer metatarsals united 13

13. Males with diffuse nuptial pads; toes

one-third to fully webbed

Neohatrachus

Males with or without nuptial spines,
no nuptial pads; toes one-fourth or
less webbed Heleioporus

14. Toes more than two-thirds webbed
Heleophryne

Toes less than one-third webbed - 15

15. Tympanum clearly visible externally
Lechriodus

Tympanum partially or completely
concealed 16

16. Prominent odontoids on lower jaw;

diastema in tooth row on maxilla

Aclelotus

No odontoids on mandible or dia-
stema in maxillary tooth row 17

17. Fingers short, first shorter than
second Kyarranus

Fingers longer, first as long as or
longer than second __ Limnodynasies

Key to African and Neotropical
Leptodactylids"^

The following key is much less defin-
itive than the former because several
leptodactylid genera cannot be separated
solely upon the basis of external char-
acters. For example, the genera Eupso-
phus and Niceforonia cannot be sepa-
rated on external characters but are
readily separated once the nature of the
occipital condylar-cervical articulation is
known. I consider these two genera to
belong to different tribes of the Telma-
tobiinae. In the later couplets in the key
I have resorted to more and more osteo-
logical features — for the most part these
are discernible with the aid of a thin
probing needle and the figures of the
skulls for reference.

The genera Paratelmatohitis and
Pseud opahidicola will key out in two
sections of the key since the nature of
the sternal style in Pseudopaludicolu is
difficult to assess from the examination
of preserved material and because the
osseous plate in Paratelmatohius is not a
style.

As will be apparent, the genera Eiip-
soplnis and Niceforonia key out together
three times in the following key (coup-
lets 32, 34, and 42) and are never sepa-
rated. Separation of these two genera on
external characters is effectively impos-
sible unless one has a breeding male
( Eiipsophus has nuptial asperities on the

•''' Insuetophnjnus can be readily separated from
all Neotropical leptodactylids as Barrio ( 1970)
pointed out on the basis of the pectoral archi-
tecture. Presumably, the quadratojugal bone is
present, and Insuetophnjnus would key out at
couplet 32 as "Eupsophus (part) and Nicefo-
ronia (part)." Inspection of the pectoral girdle
will provide evidence for separating Insueto-
l)hnjnus, because the former two genera have
arciferal girdles in contrast to the pseudofirmi-
sternal girdle of Insuetophnjnus.

LYNCH: LEPTODACTYLOID FROGS

73

thumb whereas Niceforonia does not).
Eleutlwrodactylus and Syrrliophiis key
out together (couplet 22). Most species
may be separated by the presence of
numerous supernumerary plantar tuber-
cles in Sijrrhophus and the absence of
such tubercles in Eleutlwrodactylus.
Some species of the Beta division of
Eleutlwrodactylus have as many super-
numerary tubercles as do Syrrhophus or
more.

1. Cranial bones involved in dermo-
stosis 2

Cranial bones not involved in der-
mostosis 4

2. Toes fully webbed Caudiverhera

Toes less than two-thirds webbed _.
3

3. Pupil horizontal; outer metatarsal
tubercle absent Ceratophrys

Pupil vertical; outer metatarsal
tubercle present Lepidohatraclnis

4. Pupil vertical 5

Pupil horizontal 9

5. Sternum a broad cartilaginous
(sometimes calcified) plate 6

Sternum bearing a distinct bony or
calcified style 8

6. Toes fully webbed 7

Toes free of webbing Hylorina

7. Parotoid gland present; digital tips
simple Telmatohufo

Parotoid gland absent; digital tips
pad-like Heleophryne

8. Toes fully webbed Hydrolaetare

Toes free or basally webbed

Limnomedusa

9. Sternum with a distinct bony or cal-
cified style 10

Sternum a cartilaginous, calcified, or
osseous plate 17

10. Quadratojugal bone absent 11

Quadratojugal bone present 12

11. Antebrachial tubercles present

Pseudopaludicola

Antebrachial tubercles absent

Pleurodema

12. Prevomerine teeth absent

Physalaemus

Prevomerine teeth present, on well
defined processes 13

13. Eyelid bearing fleshy tubercles

Edalorhina

Eyelid without fleshy tubercles — _ 14

14. Sternal element style shaped 15

Sternal element irregular in outline,
plate-like Paratelmatobius

15. Sternal style bifurcate posteriorly ._
Barycholos

Sternal style not bifurcate posteriorly

16

16. Terminal phalanges distinctly T-
shaped; digital discs with circum-
ferential groove Lithodytes

Terminal phalanges knobbed or
weakly T-shaped; no circumferential

groove on digital discs

Leptodactylus

17. Dorsal surface of each digital pad
bearing a pair of scute-like glands
18

Dorsal surface of digital pad or tip
without distinct glands 20

18. Quadratojugal absent or present only
as a sliver-like bone; small frogs,
adults less than 40 mm. SVL 19

Quadratojugal massive; adults 50 to
110 mm. SVL Megaelosia

19. Males with median subgular vocal
sac; nuptial spines present; pre-
vomerine teeth usually absent; quad-
ratojugal bone absent

Crossodactylus

74

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Males with paired lateral vocal sacs
or none; nuptial spines absent; pre-
vomerine teeth usually present;
quadratojugal present Hylodes

20. Tips of at least outer digits bearing
transverse, terminal groove along
edge of digital pad 21

Tips of digits without terminal
grooves 23

21. Maxillary arch toothless

Smintliilhis

Maxillary arch toothed 22

22. Lumbar gland present

Tomodactijlus

Lumbar gland absent

__ Eleutlierodactylus and Syrrhophus

23. Terminal phalanges T-shaped or Y-
shaped 24

Terminal phalanges knobbed 26

24. Digital pads large, tympanum absent
Crossodactylodes

Digital pads relatively narrow; tym-
panum visible externally 25

25. Quadratojugal bone present

Thoropa

Quadratojugal bone absent

Batrachyla

26. Quadratojugal bone absent

Pseudopaludicola

Quadratojugal bone present 27

27. Sternum an osseous plate

Paratelmatohius

Sternum cartilaginous 28

28. Supernumerary plantar tubercles
large, numerous 29

Supernumerary plantar tubercles
absent or small 30

29. Discoidal fold present

Hyhctophryne

Discoidal fold absent - Ischnocnenia

30. Tympanic annulus absent 31 '

Tympanic annulus visible externally
or concealed beneath skin 33

31. Fingers short, especially fourth finger
Euparkerella

Fingers of normal proportions to
long 32

32. Toes fully webbed

Batrachophrynus

Toes free to two-thirds webbed

Eupsophus ( part )

and Niceforonia (part)

33. Tympanum visible externally 34

Tympanum concealed beneath skin
35

34. Large frontoparietal crests present ^
Amhlyphrynus

No cranial crest Eupsophus (part)
and Niceforonia (part)

35. Inner metatarsal tubercle spade-like
elevated 36

Inner metatarsal tubercle not ele-
vated or spade-like 37

36. Large cranial crests present; parotid

glands absent

Proceratophrys (part)

Frontoparietal region flat or edge of
frontoparietals slightly ridge-like;
parotoid glands present or absent -
Odontophrynus

37. Cranial crests present

Proceratophrys (part)

Cranial crests absent 38

38. Metatarsal tubercles flat, sub-equal
in size Holoaden

Inner metatarsal tubercle at least
twice as large as outer 39

39. Outer metatarsal tubercle minute,
less than one-fourth size of inner;
toes webbed Tehnatobius

LYNCH: LEPTODACTYLOID FROGS

75

Outer metatarsal tubercle larger,
about one-third to one-half size of
inner; toes free or webbed; if webb-
ed, outer tubercle one-half size of
inner 40

40. Skin or dorsum uniformly pustular,
toes free or webbed; large inguinal
gland present Cijcloramphus

Skin or dorsum smooth or with scat-
tered pustides; toes free of webbing;
inguinal gland absent 41

41. Inner lingers and toes very short .„^
Scijthrophnjs

Fingers and toes of normal length
42

42. Tongue free all around, anterior
edge only slightly free; if appearing
adherent anteriorly, axillary patag-
ium present Zachaenus

Tongue firmly adherent anteriorly;

axillary patagium absent

Etipsophus (part)

and Niceforonia (part)

Leptodactylidae Berg, 1896 (1838)

Cystignathi Tschudi, 1838, Classification der

Batrachier, p. 25, 37, 78. [Type-genus Cys-

tignathus Wagler, 1830].
Leptodactylidae Berg, 1896, An. Mus. Buenos

Aires, 5:161. [Type-genus Leptodactyhis

Fitzinger, 1826].

The subfamilial, tribal, and generic
accounts follow. In the generic accounts,
I have provided generic synonymies in-
cluding all synonyms and permutations
thereof and the manner of designation of
the type-species; the diagnostic defini-
tion listing the states for the 50 charac-
ters scored for each genus; a statement
of composition, and where appropriate,
the most recent revisionary works on the
group; a statement of distribution; and
a section of remarks. In the remarks
section, I have justified any new generic
synonymies or partitionings. Brief re-
marks on the relationships of the genera
are made in the remarks section.

Drawings of the skulls of all genera
of leptodactylids are included with the
exception of those genera known from so
few specimens that the only available
data are taken from stereo-radiographs.
In genera having sufficient intrageneric
variation (for example, Cyclorana and
Eletitherodactyhis) , drawings of more
than one species are included.

Cycloraninae Parker, 1940

Cycloraninae Parker, 1940:12.

In his monogiaph of the Australo-
Papuan leptodactylids, Parker (1940)
amply demonstrated the desirability of
recognizing two subfamiHes. My studies
have further substantiated Parker's divi-
sion of Noble's ( 1931 ) Criniinae into two
subfamilies. Parker (1940) suggested
that with additional study, it might be
possible to demonstrate that there was a
close relationship between some of the
Cycloraninae and some Myobatrachinae.
My studies have enhanced the differ-
ences, not mitigated them, and I con-
sider that the two subfamilies possibly
represent independently derived groups.

The following diagnostic statements
are true for all members of the subfam-
ily Cycloraninae: 1) sternum cartilagi-
nous; 2) vertebral shield absent; 3)
transverse processes of anterior presacral
vertebrae not greatly expanded; 4) cer-
vical cotylar arrangement type II; 6)
cranial bones not dermostosed; 8) sacral
diapophyses dilated; 14) maxillary arch
complete; 20) frontoparietals not fused
to prootics; 21 ) temporal arcade lacking;
27) columellae present; 36) terminal
phalanges knobbed; 37) alary processes
of hyoid plate on narrow stalks; 38)
cricoid cartilage not divided ventrally;
39) m. petrohyoideus anterior and m.
sternohyoideus insert on the lateral edges
of hyoid plate; and 47) amplexus in-
guinal in known species. All species
have dilated sacral diapophyses; Parker's
( 1940 ) statement to the contrary for
Mixophyes is in error because his obser-
vation was made on a juvenile.

76

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

The primitive members of the sub-
family exhibit some or all of the follow-
ing characteristics: omosternum rela-
tively large; cervical and second verte-
brae fused; intervertebral discs free;
transverse processes of posterior pre-
sacral vertebrae short and directed an-
teriorly; frontoparietal fontanelle lack-
ing; carotid artery enclosed in a bony
canal; outer metatarsal tubercle lacking;
pupil vertical; eggs not deposited in a
foam nest; aquatic tadpoles present;
tadpole with median vent and 3/3 to
4/3 tooth rows.

Two tiibes of the Cycloraninae are
recognized herein. In the Cycloranini,
the eggs are deposited in water without
a foam nest, on land in moist situations,
or in dry burrows with a foam nest. All
species have aquatic tadpoles. Females
lack spatulate fringes on the inner fin-
gers (Fig. 48). The prevomerine den-
tigerous processes lie between, not pos-
terior to, the choanae in all Cycloranini.
In the Limnodynastini, the eggs are de-
posited in a foam nest floating on water
or in a foam nest in moist terrestrial
situations. All species except Kyarranus
and Philoria have aquatic, feeding tad-
poles. Development is direct in the
montane Kyarranus and Thihria. Fe-
males of all Limnodynastini have spatu-
late fringes on the fingers (Fig. 48),
which are used in the construction of
the foam nest (Parker, 1940, Martin,

Figure 48. Palmar views of hands of adult

females of ( A ) Cijdorana australis ( KU 93545)

and (B) Limnodyiiastcs ))cionii (KU 93562).

Both X 3.7.

1968). The prevomerine dentigerous
processes are situated posterior to the
choanae in all Limnodynastini.

Cycloranini Parker, 1940

Five genera are included in this
tribe: Cyclorana, Heleioporiis, Mixo-
phyes, Neobatradnis, and Notaden. He-
leioporiis lays its eggs in dry burrows in
a foam nest (Lee, 1966, Martin, 1968),
but the other genera lay their eggs in
water or in moist terrestrial sites (Mar-
tin, 1968).

Cyclorana Steindachner, 1867

(Figs. 49-50)

ChiioJcptes Giinther, 1859, Cat. Bat. Sal. British
Mus., p. 34 [Type-species by monotypy,
Chiwleptes australis (Gray), 1842; pre-
occupied by Chiwleptes Kirby, 1837 ( In-
secta : Hemiptera ) ] .

Cyclorana Steindachner, 1867, Reise Novara,
Amph., p. 29 [Type-species by monotypy,
Ci/clorana tiovaehoUatidiae Steindachner,
1867].

Phractops Peters, 1867, Mtber. k. Preuss. Akad.
Wiss., Berlin, 1867:30 [Type-species by
monotypy, Phractops ahitaceus Peters,
1867.].

Mitrohjsis Cope, 1889, Bull. U.S. Natl. Mus.,
34:312 [Type-species by monotypy, Chiro-
leptes alhoguttatus Giinther, 1867].

Cheiroleptes Spencer, 1901, Proc. Roy. Soc.
Victoria (2) 13:176 [Replacement name for
Chiroleptes Giinther, 1859 (preoccupied);
hence taking the same type-species].

Diagnostic definition. — 5 ) cervical
and second vertebrae free; 7) omoster-
num present, moderate in size; 9) maxil-
lary arch toothed, teeth blunt, pedicel-
late; 10) alary processes of premaxillae
directed posterodorsally, relatively wide
at base; 11) palatal shelf of premaxilla
broad, deeply incised; 12) facial lobe of
maxilla deep, weakly exostosed in alho-
guttatus and more so in australis; 13)
palatal shelf of maxilla moderate in
width, expanded posteriorly into a ptery-
goid process; 15) nasals separated me-
dially, relatively small, exostosed in
australis; 16) nasals in broad contact
with maxillae, not in contact with ptery-
goids; 17) nasals separated from fronto-
parietals; 18) frontoparietal fontanelle

LYNCH: LEPTODACTYLOID FROGS

77

Figure 49. Dorsal and ventral views of the skull
of Cyclorana australis (KU 93550, X 2.5).

lacking; 19) lateral edges of frontopari-
etals exostosed in australis, not orna-
mented in other species; 22) epiotic
eminences prominent; 23) cristae paroti-
cae long and relatively broad; carotid
artery not enclosed in a bony canal;
24) zygomatic ramus of squamosal in
broad articular contact with maxilla in
australis and extensively sculptured; in
tenuous contact with maxilla in albo-
guttatus and not ornamented; elongate
but widely separated from maxilla in
other species; 25) otic ramus of squa-
mosal long, developed into small otic
plate; in australis, otic plate is propor-
tionately larger than that in other spe-
cies and otic ramus is large and sculp-
tured; 26) squamosal-maxillary angle
26-35°; 28) prevomers large, entire,
toothed, narrowly separately medially;

29) palatines large, deep, bearing odon-
toid ridge in australis; 30) sphenethmoid
entire, extending anterior beneath na-
sals; 31 ) anterior ramus of parasphenoid
narrow, not keeled medially; 32) para-
sphenoid alae oriented at right angles to
anterior ramus, broadly overlapped by
median rami of pterygoids; 33) ptery-
goids moderate in size, no ventral flange,
anterior rami in long contact with max-
illae, not reaching palatines; 34) occipi-
tal condyles moderate sized, not stalked,
narrowly separated medially; 35) man-
dible lacking odontoids; 40) m. depres-
sor mandibulae in two slips; 41) pupil
horizontal; 42) males with median sub-
gular vocal sac, nuptial asperities on
thumb; 43) body lacking glands; 44)

Figure 50. (Top) dorsal, (middle) lateral

views of the skull of Cyclorana dahli (UMMZ

65250 ) ; and ( bottom ) lateral view of skull of

C. australis ( KU 93550). All X 2.5.

78

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

tongue large, free posteriorly; 45) toes
webbed at base to fully webbed; outer
metatarsal tubercle absent except in in-
ennis; digital tips narrow; first finger
longer than second; 46) larvae with me-
dian or dextral vent, 2/3 tooth rows,
labial papillae interrupted anteriorly;
47) amplexus inguinal; 48) eggs laid in
gelatinous masses in temporary ponds;
49) males 35-90, females 35-100 mm.
SVL; 50) tympanum visible externally;
51) thumb opposable.

Composition. — Parker (1940) recog-
nized seven species (allwguttatus, aus-
tralis, hrevipes, cultripes, dahU, inermis,
and pJattjcepljahis). One other species
(slevini) has been described since.

Distribution. — Austialia except on the
western coast and interior. The genus
does not occur on New Guinea or Tas-
mania.

Remarks. — The generic synonymy of
Cyclorana has been stable since 1940.
The status of the nominal species of the
genus is seriously in need of review. Cy-
clorana is a generalized genus of Cyclo-
raninae. Osteologically, it is least dif-
ferent from Lechriochis and Mixophyes,
but it has no close relationship with
either of these genera.

The variation in the skulls of this
genus is striking. Cyclorana albogtittata
is intermediate between the heavily
exostosed australis and the other species
of the genus. The only other leptodac-
tylid genus with comparable variation in
the skull bones is Eleutherodactylus.

Heleioporus Gray, 1841

(Figs. 51-52)

Heleioporus Gray, 1841, Ann. Mag. Nat. Hist.,
(1)7:91 [Type-species by monotypy, Helei-
oporus alhopunctatus Gray, 1841].

Helioporus Gray, in Grey, 1841, J. Exped. Cent.
Australia, 2:447 [Emendation of Heleio-
porus Gray, 1841].

Perialia Gray, in Eyre, 1845, J. Exped. Cent.
Australia, 1:406 [Type-species by monotypy,
Perialia eijeri Gray, 1845].

Philocrijphus Fletcher, 1894, Proc. Linn. Soc.
New South Wales, (2)8:233 [Type-species
by monotypy, Philocrtjphus flavoguttattis
Fletcher, 1894].

Diagnostic definition. — 5) cervical
and second vertebrae fused; 7) omo-
sternum present, small; 9) maxillary
arch toothed, teeth blunt, pedicellate;

10) alary processes of premaxillae di-
rected posterodorsally, broad at base;

11) palatal shelf of premaxilla relatively
narrow, moderately long palatal process;

12) facial lobe of maxilla of moderate
depth, not exostosed; 13) palatal shelf
of maxilla relatively narrow, pterygoid
process present; 15) nasals small, short,
very wide, separated medially; 16) na-
sals in broad contact with maxillae, sep-
arated from pterygoids; 17) nasals not
in contact with frontoparietals; 18) fron-
toparietal fontanelle moderate in size;
19) frontoparietals not ornamented; 22)
epiotic eminences prominent posteriorly;
23) cristae paroticae long and narrow;
carotid artery enclosed in a bony canal,
roofed over; 24) zygomatic ramus of

Figure 5 1 . Dorsal and ventral views of the skull
oi Heleioporus eyeri (UMMZ 124504, X 6).

LYNCH: LEPTODACTYLOID FROGS

79

Figure 52. Lateral view of skull of Heleioponis
eyed (UMMZ 124504, x 6).

squamosal short, slightly longer than otic
ramus; 25) otic ramus of squamosal de-
veloped into small otic plate; 26) squa-
mosal-maxillary angle about 50°; 28)
prevomers large, entire, toothed, nar-
rowly separated medially; 29) palatines
large, narrowly separated medially; 30)
sphenethmoid entire, extending anterior-
ly beneath posterior half of nasals; 31)
anterior ramus of parasphenoid narrow,
not keeled; 32) parasphenoid alae ori-
ented at right angles to anterior ramus,
broadly overlapped by median rami of
pterygoids; 33) pterygoids relatively
slender, anterior rami in long contact
with maxillae, reaching palatines; 34)
occipital condyles large, not stalked, nar-
rowly separated medially; 35) mandible
lacking odontoids; 40) m. depressor
mandihulae in two slips; 41) pupil ver-
tical; 42) males with large nuptial spines
on thumb or not, never pad-like asperi-
ties; males with median, subgular vocal
sac; 43) extensive parotoid glands pres-
ent, small inguinal glands present; 44)
tongue large, notched and free pos-
teriorly; 45) feet basally webbed, outer
metatarsal tubercle absent, inner meta-
tarsal tubercle spade-like, tips of digits
narrow, first finger longer than second;
46) larvae with median or dextral vent,
2/3 to 5/4 tooth rows, labial papillae
broadly interrupted anteriorly; 47) am-
plexus inguinal; 48) eggs laid in foam
nest in dry burrow, tadpoles emerge
when nest is flooded; 49) males 38-95,
females 41-95 mm. SVL; smallest species
is psammopliilus, largest is australiacus;
50) tympanum visible externally.

Composition. — Lee (1966) recog-
nized six species (albopunctatus, aus-
traliacus, barycragus, eijeri, inornatus,
and psammophilus).

Distribution. — Southeastern and
southwestern Australia.

Rejnarks. —Farker (1940) did not
differentiate the genera Heleioporus and
Neobatrachus. Main, Lee, and Little-
john (1958) considered the ethological
and morphological differences between
the groups ample for generic distinction.
Moore (1961) and Lee (1966) con-
curred in this opinion; I consider the
differences in the sizes of the transverse
processes of the presacral vertebrae addi-
tional support for the continued separa-
tion of these genera. The two genera are
unquestionably related but are not so
similar that their combination would re-
sult in a clarification of relationships.
Neobatraclnis is more closely allied to a
third related genus, Notaden.

Mixophyes Giinther, 1864

(Figs. 53-54)

Mixophyes Giinther, 1864, Proc. Zool. Soc. Lon-
don, p. 46 [Type-species by monotypy,
Mixophtjes fasciolatus Giinther, 1864].

Diagnostic definition. — 5) cervical
and second vertebrae free; 7) omoster-
num present, relatively large; 9) maxil-
lary arch toothed, teeth blunt, pedicel-
late; 10) alary processes of premaxillae
directed posterodorsally, broad at base;
11) palatal shelf of premaxilla narrow,
palatal process elongate; 12) facial lobe
of maxilla deep with a slight squamosal
processes, not exostosed; 13) palatal
shelf of maxilla very narrow, no ptery-
goid process; 15) nasals large, in median
contact anteriorly, separated posteriorly,
exposing sphenethmoid; 16) nasals in
contact with maxillae, not in contact
with pterygoids; 17) nasals in tenuous
contact with frontoparietals; 18) fron-
toparietal fontanelle absent; 19) fronto-
parietals not ornamented; 22) epiotic
eminences prominent; 23) cristae paroti-
cae long and narrow; carotid artery en-
closed in a complete bony canal; 24)

80

MISCELLAxXEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 53. Dorsal and ventral views of skull of
Mixophyes fasciolatiis ( KU 56627, X 3.5).

zygomatic ramus of squamosal elongate,
tendon contacting squamosal process of
maxilla; 25) otic ramus of squamosal
long, developed medially into otic plate;
26) squamosal-maxillary angle 45-50°;
28) prevomers small, entire, toothed,
separated medially; 29) palatines thin,
separated medially, bearing odontoid
ridges; 30) sphenethmoid entire, extend-
ing anteriorly to anterior edge of nasals;
31 ) anterior ramus of parasphenoid nar-
row, not keeled; 32) parasphenoid alae
deflected posteriorly, overlapped lateral-
ly by median rami of pterygoids; 33)
pterygoids large, anterior rami in long
contact with maxillae, nearly reaching
palatines and nasals; 34) occipital con-
dyles moderate sized, not stalked, sep-
arated medially; 35) mandible lacking
odontoids; 40) m. depressor mandibulae
in two slips; 41) pupil vertical; 42)
males with median subgular vocal sac,
nuptial asperities on thumb; 43) body

lacking glands; 44) tongue large,
rounded, only posterior edge free; 45)
toes two-thirds webbed, outer metatar-
sal tubercle absent, inner metatarsal
tubercle not spade-like, tips of digits
narrow; 46) larvae with dextral vent,
6/3 tooth rows, labial papillae not in-
terrupted anteriorly; 47) amplexus in-
guinal; 48) eggs laid in terrestrial situa-
tions and hatch upon flooding; 49) males
and females 50-100 mm. SVL; 50) tym-
panum visible externally.

Composition. — Only one species is
now recognized, but it is apparently a
composite of two (Moore, 1961, Little-
john, 1968).

Distribution. — Mountains and coastal
areas of eastern Australia.

Remarks. — Mixophyes has been rec-
ognized as a distinctive genus since its
discovery. This distinction was based in
part on misconceptions. Boulenger
(1882) reported the sacral diapophyses
as rounded; Parker (1940) agreed.
Hecht (1960) reported the sacral dia-
pophyses were dilated. The specimens
I have examined all have dilated sacral
diapophyses. Fletcher (1889) stated
that Mixophyes exhibits axillary amplex-
us. Littlejohn (pers. comm.) informs me
that the amplectic pattern is inguinal, as
in all other Australo-Papuan leptodac-
tylids.

Figure 54. Lateral and occipital views of skull
of Mixophyes fasciolatiis ( KU 56627, X 5.5).

LYNCH: LEPTODACTYLOID FROGS

81

Neobatrachus Peters, 1863

(Figs. 55-56)

Neobatrachus Peters, 1863, Mtber. k. Preuss.
Akad. Wiss., Berlin, 1863:234 [Type-species
by monotypy, Neobatrachus pictus Peters,
1863].

Diagnostic definition. — 5) cervical
and second vertebrae fused; 7) omoster-
num minute; 9) maxillary arch toothed,
teeth blunt, pedicellate; 10) alary proc-
esses of premaxillae long, directed pos-
terodorsally, relatively wide at base; 11)
palatal shelf of premaxilla narrow, pala-
tal process long; 12) facial lobe of max-
illa deep, not exostosed; 13) palatal shelf
of maxilla narrow, no pterygoid process;
15) nasals small, separated medially; 16)
nasals in contact with maxillae, not with
pterygoids; 17) nasals not in contact
with frontoparietals; 18) frontoparietal
fontanelle moderate in size; 19) fronto-
parietals not ornamented; 22) epiotic
eminences prominent; 23) cristae paroti-

FiGURE 55. Dorsal and ventral views of skull of
Neobatrachus pictus (FMNH 97281, X 3).

Figure 56. Lateral views of the skulls of Neo-
batrachus pictus (FMNH 97281, top) and
Notaden nichollsi ( KU 93582, bottom). Both

X3.

cae long and narrow; carotid artery lies
in a deep groove, exposed dorsally; 24)
zygomatic ramus of squamosal minute;
25) otic ramus of squamosal very small,
developed medially into a small otic
plate; 26) squamosal-maxillary angle 50-
55°; 28) prevomers of moderate size,
entire, toothed, narrowly separated me-
dially; 29) palatines thin, widely sep-
arated medially; 30) sphenethmoid en-
tire, extending anteriorly beneath nasals;
31 ) anterior ramus of parasphenoid nar-
row, weakly keeled; 32) parasphenoid
alae oriented at right angles to anterior
ramus, narrowly overlapped laterally by
median rami of pterygoids; 33) ptery-
goids relatively large, anterior rami in
long contact with maxillae, not reaching
palatines; 34) occipital condyles rela-
tively small, not stalked, narrowly sep-
arated medially; 35) mandible lacking
odontoids; 40) m. depressor mandibuhe
in two slips; 41) pupil vertical; 42) males
with median subgular vocal sac; nuptial
pad (callosities) on thumb and second
finger; 43) body lacking glands; 44)
tongue large, round, free behind; 45)
toes one-fourth to fully webbed, outer

82

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

metatarsal tubercle absent, inner meta-
tarsal tubercle spade-like, tips of digits
narrow, first finger longer than second;
46) larvae with dextral vent, 3/3 tooth
rows, labial papillae interrupted an-
teriorly; 47) amplexus inguinal; 48) eggs
deposited in long strings in slow moving
streams and temporary ponds; 49) adults
not exceeding 50 mm. SVL; 50) tym-
panum indistinct externally, concealed
beneath skin.

Composition. — Five species (centra-
lis, pelohatoicJes, picttis, sutor, and tcih-
morei) are presently recognized, al-
though no one has reviewed the group
since Parker (1940).

Distrihtition. — Mainland Australia
except in northern West Australia and
along the northern coast, where Neo-
hatrachiis is apparently replaced eco-
logically by its close relative, Notaden.

Remarks. — Main, Lee, and Littlejohn
(1958) argued that the Heleioporus
pictus group (Neohatrachiis) should be
afi^orded generic status since it differs
ethologically from typical Heleioporus.
Typical Heleioporus lays the eggs in
burrows in a foam nest and when the
burrow is flooded, the tadpoles emerge
to complete their development like typi-
cal frogs. Neohatrachiis lays long egg
strings ( like Biifo ) in still or slowly mov-
ing water. The two groups also differ in
the kind of nuptial excrescence (spines
in Heleioporus, roughened pad in Neo-
hatrachiis), adult size (Heleioporus are
larger frogs), and the shape of the ster-
num (bifurcate posteriorly in Heleio-
porus). The transverse processes of the
posterior presacral vertebrae of the two
genera differ in size (Fig. 30), and there
is a slight difference in the sizes of the
zygomatic and otic rami of the squa-
mosals.

The separation of these two genera is
based more on the so-called adaptive
approach to genera than the morpho-
logical approach. Without the ethologi-
cal data, I doubt if I would have adopted
a different approach from that taken by
Parker (1940).

Notaden Glinther, 1873

( Figs. 56-57 )

Notaden Giinther, 1873, Ann. Mag. Nat. Hist.,
(4)11:349 [Type-species by monotypy,
Notaden bennetti Giinther, 1873].

Diagnostic definition. — 5) cervical
and second vertebrae fused; 7) omoster-
num absent; 9) maxillary arch eden-
tate; 10) alary processes of premaxillae
elongate, directed dorsally, narrow at
base; 11) palatal shelf of premaxilla nar-
row, palatal process relatively short; 12)
facial lobe of maxilla shallow; 13) pala-
tal shelf of maxilla absent; 14 ) maxillary
arch incomplete, maxilla not contacting
quadratojugal or premaxilla, quadrato-
jugal present; 15) nasals small and sep-
arated medially; 16) nasals not in con-
tact with maxillae or pterygoids; 17) na-
sals in tenuous contact with frontopari-
etals; 18) frontoparietal fontanelle large;
19) frontoparietals not ornamented; 22)
epiotic eminences prominent; 23) cristae
paroticae short, stocky; carotid artery
lies in a shallow groove exposed dorsal-
ly; 24-25) zygomatic and otic rami of
squamosal lacking; 26) squamosal-maxil-

FiGURE 57. Dorsal and ventral views of skull of
Notaden nichollsi ( KU 93582, X 3 ) .

LYNCH: LEPTODACTYLOID FROGS

83

lary angle about 80°; 28) prevomers
moderately large, entire, toothed, sep-
arated medially; 29) palatines reduced
in size, not contacting maxillae and wide-
ly separated medially; 30) sphenethmoid
entire, small, not extending anteriorly to
nasals; 31) anterior ramus of parasphe-
noid broad, short, not keeled; 32) para-
sphenoid alae oriented at right angles to
anterior ramus, not overlapped laterally
by median rami of pterygoids; 33 ) ptery-
goids small, anterior rami in long contact
with maxillae, usually contacting pala-
tines; 34) occipital condyles large, not
stalked, narrowly separated medially;
35) mandible lacking odontoids; 40) 7n.
depressor 7nandihiilae in two slips; 41)
pupil horizontal; 42) males with median
subgular vocal sac; nuptial pad on
thumb; 43) dorsum covered with at least
two ill-defined glands, less discrete but
more extensive than in Heleioporus; 44)
tongue large, round, not free behind;
45) toes one-half to two-thirds webbed,
outer metatarsal tubercle absent, inner
metatarsal tubercle spade-like, tips of
digits narrow, first finger about as long
as second; 46) larvae with median vent,
3/3 tooth rows, labial papillae inter-
rupted anteriorly; 47-48); 49) adults to
70 mm. SVL; 50) tympanum concealed.

Composition. — Hosmer (1962) rec-
ognized three species — hennetti, mela-
noscaplnis, and nichollsi.

Distribution. — Northwestern, north-
central, and southeastern Australia west
of the dividing range.

Remarks. — Notaden has been recog-
nized as distinct since its description
chiefly because it lacks maxillary teeth.
The loss of teeth reflects a general re-
duction in the amount of bone; in Nota-
den, the reduction is striking. The loss
of the upper portions of the squamosals
is unique in the family, as is the free
floating maxilla. Because Notaden lives
in an arid environment, Parker ( 1940 )
thought it might exhibit direct develop-
ment. Slater and Main (1963) found

tadpoles in ponds and suggested that de-
velopment was rapid, but definitely was
not abbreviated.

Notaden is closely related to Heleio-
porus and Neohatrachus. All have free
intervertebral discs, fused cervical and
second vertebrae, short transverse proc-
esses of the posterior presacral vertebrae,
moderate to large frontoparietal fonta-
nelles, deep skulls, and large, spade-like
inner metatarsal tubercles. Notaden has
a horizontal pupil and lacks maxillary
teeth, whereas the other two genera have
vertical pupils and have maxillary teeth.
Notaden and Heleioporus have extensive
glandular areas on the dorsum; Neo-
hatrachus lacks glands. Neohatrachus
and Notaden have nuptial pads, and He-
leioporus has large spines on the thumb.
The tadpoles of the three genera are
similar and cannot be separated generi-
cally owing to the intrageneric variability
in vent position and tooth rows in He-
leioporus.

Limnodvnastini New Tribe

Limnodynastine leptodactylids build
a foam nest in which the eggs are de-
posited. The few observations that have
been made indicate that the female uses
the spatulate fringes on the inner fingers
(Fig. 48) to build the nest. Develop-
ment is direct in two of these genera
(Kyarranus and Philoria).

Martin (1968) proposed an evolu-
tionary course of terrestrial foam nest
development. He cited Australo-Papuan
genera where germane, but used Neo-
tropical genera for certain stages of de-
velopment. The foam nest is deposited
in open water by some Limnodijnastes,
in water-filled burrows adjacent to open
water in other Limnodijnastes, in pud-
dles adjacent to open water in Leptodac-
tylus pentad actijlns, in dry burrows
which will later flood in Heleioporus,
and in moist, terrestrial situations in
Kyarranus and Philoria.

84

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Adelotus Ogilby, 1907

(Figs. 58-59)

Cryptotis Giinther, 1863, Ann. Mag. Nat. Hist.,
(3)11:27 [Type-species by monotypy,
Cryptotis brevis Giinther, 1863; preoccupied
by Cryptotis Poniel, 1848 (Mammalia: In-
sectivora ) ] .

Adelotus Ogilby, 1907, Proc. Roy. Soc. Queens-
land, 20:32 [Replacement name for Crypt-
otis Giinther, 1863 (preoccupied); hence
taking same type-species].

Diagnostic defi n itio n. — 5 ) cervical
and second vertebrae fused; 7) omoster-
num present, moderate sized; 9) maxil-
lary arch toothed, teeth elongate, pedi-
cellate; distinct diastema in tooth row;
10) alary processes of premaxillae di-
rected posterodorsally, elongate, rela-
tively broad at base; 11) palatal shelf of
premaxilla narrow with a moderately
large palatal process and long process
lying along maxilla; 12) facial lobe of
maxilla deep; maxilla deep for entire
length, not exostosed; 13) palatal shelf
of maxilla narrow, pterygoid process well
developed; 14) quadratojugal expanded;
15) nasals wide and short, small, sep-

FiGURE 58. Lateral and dorsal views of the skull
of Adelotus brevis (KU 56242, X 2.5).

Figure 59. Ventral view of the skull of Adelo-
tus brevis (KU 56242, X 2.5).

arated medially; 16) nasals in contact
with maxillae, not with pterygoids; 17)
nasals widely separated from frontopari-
etals; 18) frontoparietal fontanelle lack-
ing; 19) frontoparietals not exostosed
but bearing large sagittal crest; 22) epi-
otic eminences obsolete; 23) cristae
paroticae short but relatively broad;
carotid artery not enclosed in an open or
closed canal; 24) zygomatic ramus of
squamosal slender, of moderate length;
25) otic ramus of squamosal short, de-
veloped medially into prominent otic
plate; 26) squamosal-maxillary angle
about 45°; 28) pre vomers toothed, en-
tire, separated medially, dentigerous
ramus greatly elongated; 29) palatines
very broad, lacking odontoid ridge, sep-
arated medially; 30) sphenethmoid en-
tire, relatively small, rounded anteriorly,
not reaching nasals; 31) anterior ramus
of parasphenoid broad, not keeled, an-
terior edge deeply serrated; 32) para-
sphenoid alae deflected posteriorly,
short, overlapped laterally by median
rami of pterygoids; 33) pterygoids rela-
tively large, anterior rami in long con-
tact with maxillae, nearly reaching pala-
tines; 34) occipital condyles large, close-
ly approximated medially; 35) mandible
bearing elongate tusk (Fig. 15), longer
in male than in female; 40) m. depressor
mandihulae in two slips; 41 ) pupil hori-

LYNCH: LEPTODACTYLOID FROGS

85

zontal; 42) males with median subgular
vocal sac; no nuptial asperities on
thumb; 43) body lacking glands; 44)
tongue large, posterior edge free; 45)
toes with basal webbing, outer meta-
tarsal tubercle present, digital tips nar-
row, first finger as long as second; 46)
larvae \\ith dextral vent, 2/3 tooth rows,
labial papillae narrowly interrupted an-
teriorly; 47) amplexus inguinal; 48) eggs
laid in foam nest in ponds or running
water (streams); 49) males to 50 mm.,
females to 40 mm. SVL; 50) tympanum
usually completely concealed.

Composition. — A single species, A.
brevis.

Distribution. — Coastal southeastern
Australia.

Remarks. — Since its discovery, the
tusked frog has been recognized as ge-
nerically distinct. The large mandibular
tusks are characters sui generis. The
species was originally named Cryptotis
brevis but the generic name, Cryptotis
Giinther, 1863, is a junior homonym of
Cryptotis Pomel, 1848, the generic name
of some American shrews. Adelotus is
not closely related to any other genus of
the Limnodynastini, but is least different
from Limnodynastes.

Lechriodus Boulenger, 1882
(Fig. 60)

Batrachopsis Boulenger, 1882, Cat. Batr. Sal.
British Mus., p. 439 [Type-species by mono-
typy, Asterophrys melanoptjga Doria, 1875;
preoccupied by Batrachopsis Fitzinger, 1843
(Amphibia: Caudata)].

Lechriodus Boulenger, 1882, Cat. Batr. Grad.
British Mus., p. 116 [Replacement name for
Batrachopsis Boulenger, 1882 ( preoccupied )
and hence taking same type-species].

Phanerotis Boulenger, 1890, Proc. Linn. Soc.
New South Wales, (2)5:594 [Type-species
by monotypy, Phanerotis ftetcheri Boulen-
ger, 1890].

Diagnostic definition. — 5) cervical
and second vertebrae free; 7) omoster-
num present, moderate sized; 9) maxil-
lary arch toothed, teeth blunt, pedicel-
late; 10) alary processes of premaxillae
directed posterodorsally, wide at base;
11) palatal shelf of premaxilla moderate

Figure 60. Dorsal and ventral views of skull of
Lechriodus fletcheri (AMNH 59488, X 3).

in width, deeply incised; 12) facial lobe
of maxilla deep, not exostosed; 13) pala-
tal shelf of maxilla relatively narrow,
small pterygoid process; 15) nasals mod-
erate sized, apparently in median con-
tact; 16) nasals in contact with maxil-
lae, not with pterygoids; 17) nasals not
in contact with frontoparietals; 18) fron-
toparietal fontanelle lacking; 19) fronto-
parietals not ornamented; 22) epiotic
eminences poorly defined; 23) cristae
paroticae long and relatively broad;
carotid artery enclosed in long, roofed,
bony canal; 24) zygomatic ramus of
squamosal slightly shorter than otic
ramus; 25) otic ramus of squamosal of
moderate length, expanded medially into
small otic plate; 26) squamosal-maxillary
angle about 50°; 28) prevomers entire,
toothed, large, dentigerous rami in tenu-
ous median contact; 29) palatines large,
narrowly separated medially; 30) sphen-
ethmoid entire, extending anteriorly
beneath posterior edge of nasals; 31) an-

86

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

terior ramus of parasphenoid narrow, not
keeled; 32) parasphenoid alae oriented
at right angles to anterior ramus, over-
lapped laterally by median rami of
pterygoids; 33) pterygoids relatively
large, anterior rami in long contact with
maxillae, nearly reaching palatines; 34)
occipital condyles moderately large, not
stalked, narrow separated medially; 35)
mandible lacking odontoids; 40) m.
depressor mancUhiiloe in two slips;
41) pupil horizontal; 42) males with
median subgular vocal sac; nuptial as-
perities of many small spines on thumb;
43) body lacking glands; 44) tongue
round, posterior edge free; 45) toes lack
webbing, outer metatarsal tubercle ab-
sent, digital tips narrow, first finger as
long as second; 46) larvae with dextral
vent, 2/3 tooth rows, labial papillae
broadly interrupted anteriorly; 47) am-
plexus inguinal; 48) eggs laids in foam
nest in temporary ponds and puddles;
49) males 40-70, females 40-96 mm.
SVL; 50) tympanum visible externally.

Composition. — Parker (1940) recog-
nized four species (fetcheri, melanopij-
gus, papuanus, and platyceps). The spe-
cies of this genus appear to be sorely in
need of review.

Distribution. — Eastern New Guinea,
the Aru Islands, and the eastern edge of
Australia southward to New South
Wales.

Remarks. — Lechriodus was long-
thought to be the only Australo-Papuan
pelobatid genus. Noble (1926a) demon-
strated that Lechriodus was a leptodac-
tylid (bufonid in his terminology). In
many respects, Lecliriodus is the most
primitive genus of Limnodynastini — the
skull is completely roofed, and the maxil-
lary dentition is similar to that seen in
the Cycloranini. In terms of breeding
biology, it is only slightly more advanced
than the generalized Limnodynastes.
The firmly ankylosed intervertebral discs
and the non-fusion of the cervical and
second vertebrae possibly are advanced
characters.

Limnodynastes Fitzinger, 1843

(Figs. 61-62)

Limnodynastes Fitzinger, 1843, Syst. Rept., 31
[Type-species by original designation, Cys-
ti^nathiis peronii Dumeril and Bibron,
1841].

Wa^leria Girard, 1853, Proc. Acad. Nat. Sci.
Philadelphia, 6:421 [Type-species by mono-
typy, Cystignathus peronii Dumeril and Bib-
ron, 1841].

Platyplectntm Giinther, 1863, Ann. Mag. Nat.
Hist., (3)11:27 [Type-species by monotypy,
Phityplectrinn marmoratum Giinther, 1863].

PlatypJectwn Peters, 1863, Mtber. k. Preuss.
Akad. Wiss., Berlin, 1863:235 [Emendation
of Platyplectrum Giinther, 1863; hence tak-
ing .same type-species].

Lymnodynastcs Cope, 1865, Rev. Nat. Hist.,
5:113 [Emendation of Limnodynastes Fitz-
inger, 1843; hence taking same type-
species].

Opisthodon Steindachner, 1867, Reise Novara,
Amph., p. 9 [Type-species by mono-
typy, Opisthodon frauenfeJdti Steindachner,
1867].

Heliorana Steindachner, 1867, Ibid., p. 32
[Type-species by monotypy, Heliorana
graiji Steindachner, 1867].

Ranaster MaCleay, 1878, Proc. Linn. Soc. New
South Wales, 2:135 [Type-species by mono-
typy, Ranaster convexiuscidus MaCleay,
1878].

Diagnostic definition. — 5) cervical
and second vertebrae fused; 7) omoster-
num present, moderately large; 9) maxil-
lary arch toothed, teeth blunt, pedicel-
late; 10) alary processes of premaxillae
directed posterodorsally, wide at base;
11) palatal shelf of premaxilla relatively
narrow, palatal process present; 12) fa-
cial lobe of maxilla deep, not exostosed;
13) palatal shelf of maxilla narrow,
pterygoid process small; 15) nasals rela-
tively small and widely separated medi-
ally; 16) nasals in broad contact with
maxillae, not contacting pterygoids; 17)
nasals separated from frontoparietals;
18) frontoparietal fontanelle present,
moderate-sized; 19) frontoparietals not
ornamented; 22) epiotic eminences
prominent; 23) cristae paroticae rela-
tively short, stocky; carotid artery not
enclosed in bony canal or groove; 24)
zygomatic ramus of squamosal as long
as otic ramus, widely separated from
maxilla; 25) otic ramus of squamosal

LYNCH: LEPTODACTYLOID FROGS

87

Figure 61. Dorsal and ventral views of skull of
Limnochjnastes dorsalis ( KU 93553, X 2.2).

moderate length, developed medially in-
to small otic plate; 26) squamosal-maxil-
lary angle about 50°; 28) prevomers
large, entire, toothed, narrowly separated
medially; 29) palatines broad, narrowly
separated medially; 30) sphenethmoid
entire, extending anteriorly to center of
nasals; 31) anterior ramus of parasphe-
noid of moderate width, not keeled; 32)
parasphenoid alae oriented at right an-
gles to anterior ramus, broadly over-
lapped laterally by median rami of
pterygoids; 33) pterygoids slender and
large, anterior rami in long contact with
maxillae, not reaching palatines; 34) oc-
cipital condyles large, not stalked, nar-
rowly separated medially in dorsalis
group, moderately separated medially in
pewnii group; 35) mandible lacking
odontoids; 40) m. depressor mandibulae
in two slips; 41) pupil horizontal; 42)
males with median subgular vocal sac;
males of convexiusculus, dorsalis, orna-
tiis, spenceri, and tasrnaniertsis have nup-
tial pads on thumb, whereas males of
fietcheri, peronii, salmini, and the Eids-

vold tasmaniensls lack nuptial asperi-
ties; 43) tibial glands in dorsalis, other
species lacking glands; 44) tongue large,
slightly notched, free posteriorly; 45)
toes lacking web to one-half webbed,
inner metatarsal tubercle spade-like in
the dorsaUs group, normal in the peronii
group, outer metatarsal tubercle absent
in all species except tasmaniensis, digi-
tal tips narrow, first finger shorter than
to longer than second; 46) larvae with
median vent, 3/3 to 4/3 tooth rows, la-
bial papillae broadly interrupted an-
teriorly; 47) amplexus inguinal; 48) eggs
laid in foam nest floating in open water
or in water-filled holes adjacent to the
main body of water; 49) males 32-87,
females 32-83 mm. SVL; 50) tympanum
visible externally or indistinct.

Composition. — Parker (1940) recog-
nized eight species — convexiusculus, dor-
salis, fietcheri, ornatus, peronii, salmini,
spenceri, and tasmaniensis. Moore
( 1961 ) suggested that spenceri was a
subspecies of ornatus; he discussed a
population of tasmaniensis-Vike frogs and
considered them subspecifically or spe-
cifically distinct but did not name or
otherwise treat them taxonomically.
Littlejohn (1968) noted northern and
southern call races of tasmaniensis.
Parker (1940) recognized five subspecies
of dorsalis, but Moore (1961) did not
discuss their status; Littlejohn (1968)
recognized the fom* that occur in south-
eastern Australia and Tasmania but
stated that they hybridize (not inter-
grade ) wherever their ranges meet there-
by suggesting that the five are species.
The nominal forms are dorsalis, dumerili,
grayi, insularis, and interior is.

Distribution. — Australia and Tas-

FiGURE 62. Lateral view of skull of Limno-

dynastes dorsalis ( KU 93553, X 2 ) .

88

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

mania; the genus is apparently absent
from most of the western Eyrean sub-
region.

Remarks. — The two species-groups
of Limnodynastes are readily separated.
The dor sails group (dor sails, ornatus,
and spenceri) is composed of medium
sized, stocky frogs with large, spade-like
inner metatarsal tubercles. The peronii
group is composed of smaller species
with a Rana-\ike habitus and "normal"
inner metatarsal tubercle. Frogs of the
dorsalis group are burrowers, whereas
those of the peronii group seldom bur-
row. The data presently available are
sufficient to assert that the genus Lim-
nodynastes does not contain enough
variability to justify recognition of sub-
genera.

Cope (1866) combined Limnody-
nastes with Borhorocaetes (an emenda-
tion of Borborocoetes Bell; principally
refers to Eupsophus). In an earlier
paper Cope ( 1865 ) had considered them
closely related but separable on the basis
of the lateral extension of the prevo-
merine tooth rows. Limnodynastes and
Eupsophus differ in many respects —
principally characters with which Cope
was not familiar. The breeding biology
of the two is quite different, and the
vertebral column of Limnodynastes does
not resemble that of Eupsophus; both
are relatively generalized leptodactylids.
The superficial skull bones of the two
are very similar. Among the most im-
portant characters in Cope's system were
the separation of the prefrontals ( =na-
sals) and the presence of a frontopari-
etal fontanelle — Eupsophus and Limno-
dynastes agree in these two characters.

Kyarranus Moore, 1958

(Fig. 63)

Kyarranus Moore, 1958, Amer. Mus. Novitates,
1919:1 [Type-species by original designa-
tion, Kyarranus sphagnicolus Moore, 1958
{=Kyarranus sphagnicola)].

Diag,nostic definition. — 5) cervical
and second vertebrae fused; 7) omoster-
num present, small; 9) maxillary arch

Figure 63. Lateral ( X 2.6) and dorsal ( X 2.3)

views of the skull of Kyarranus sphagnicola

(KU 110331).

toothed, teeth blunt, pedicellate; 10)
alary processes of premaxillae directed
dorsally, relatively wide at base; 11)
palatal shelf of premaxilla relatively nar-
row, palatal process large; 12) facial
lobe of maxilla deep, not exostosed; 13)
palatal shelf of maxilla of moderate
width, pterygoid process small; 15) na-
sals small, widely separated medially;
16) nasals in contact with maxillae, not
with pterygoids; 17) nasals not in con-
tact with frontoparietals; 18) frontopari-
etal fontanelle large; 19) frontoparietals
not ornamented; 22) epiotic eminences
obsolete; 23) cristae paroticae short and
stocky; carotid artery enclosed in short,
roofed, bony canal; 24) zygomatic ramus
of squamosal relatively long, widely sep-
arated from maxilla; 25) otic ramus of
squamosal slightly shorter than zygo-
matic ramus, expanded medially into
moderately large otic plate; 26) squa-
mosal-maxillary angle about 65°; 28)
prevomers entire, toothed, widely sep-
arated; 29) palatines wide, narrowly
separated medially; 30) sphenethmoid
entire, extending anteriorly to anterior
edge of nasals; 31) anterior ramus of
parasphenoid broad, not keeled; 32)
parasphenoid alae oriented at right
angles to anterior ramus, overlapped
laterally by median rami of pterygoids;

LYNCH: LEPTODACTYLOID FROGS

89

33) pterygoids slender, anterior rami in
long contact with maxillae, nearly reach-
ing palatines; 34) occipital condyles
large, not stalked, narrowly separated
medially; 35) mandible lacking odon-
toids; 40) m. depressor mandihuJae in
two slips; 41) pupil horizontal; 42)
males with nuptial callosities on thumb;
median, subgular vocal sac; 43) body
lacking glands; 44) tongue large, notched
posteriorly, posterior edge free; 45) toes
free of webbing, outer metatarsal tuber-
cle lacking, digital tips narrow, first fin-
ger shorter than second; 46) develop-
ment direct; 47) amplexus inguinal; 48)
eggs laid underground in sphagnum or
loose earth in a foam nest; 49) males 21-
35, females 30-34 mm. SVL; 50) tym-
panum concealed.

Composition. — Two species (love-
ridgei and sphagnicola).

Distribution. — The New England
Range of eastern Australia and a dubi-
ous record from Warragul, Victoria
( lowlands ) .

Remarks. — Parker (1940) named
loveridgei as a species of FJnJoria; Moore
(1958) reexamined the holotype of Vhi-
loria frosti and concluded that loveridgei
was generically different from frosti. He
included loveridgei with a new species
from the southern part of the New Eng-
land Range in a new genus, Kijarranus.
The principal differences between Ki/ar-
ronus and Limnodijnastes, according to
Moore ( 1958, 1961 ) , were in breeding
biology — the former exhibits direct de-
velopment and the latter has tadpoles.
Littlejohn ( 1963 ) questioned the separa-
tion of Kijarranus and Philoria after he
found that Philoria also exliibits direct
development. The two genera differ in
the development of the parotoid glands
and in the arrangement of several skull
bones. Kijarranus has a large frontopari-
etal fontanelle and widely separated na-
sals; in Fhiloria, the fontanelle is small
and the nasals large and broadly in con-
tact.

Philoria Spencer, 1901

(Fig. 64)

Fhiloria Spencer, 1901, Proc. Roy. Soc. Victoria,
13(2) :176 [Type-species by monotypy,
Fhiloria frosti Spencer, 1901].

Diagnostic definition. — 5) cervical
and second vertebrae fused; 7) omoster-
num present, small; 9) maxillary arch
toothed, teeth blunt, pedicellate; 10)
alary processes of premaxillae directed
posterodorsally, wide at base; 11) pala-
tal shelf of premaxilla of moderate width,
palatal process small; 12) facial lobe of
maxilla deep anteriorly, shallow posteri-
orly; 13) palatal shelf of maxilla of mod-
erate width, large pterygoid process pos-
teriorly; 15) nasals large, in broad medi-
an contact; 16) nasals not in contact
with maxillae or pterygoids; 17) nasals
in contact with frontoparietals; 18) fron-
toparietal fontanelle present, relatively
small; 19) frontoparietals not orna-
mented; 22) epiotic eminences poorly
defined; 23) cristae paroticae relatively
short and stocky; carotid artery enclosed

Figure 64. Dorsal and ventral views of skull of
Philoria frosti (KU 50699, X 4).

90

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

in a short, complete, canal; 24) zygo-
matic ramus of squamosal of moderate
length, widely separated from maxilla;
25) otic ramus of squamosal long, large,
expanded medially into very large otic
plate; 26) squamosal-maxillary angle
about 58°; 28) pre vomers moderate-
sized, entire, toothed, widely separated
medially; 29); palatines large, widely
separated medially; 30) sphenethmoid
entire, large, extending anteriorly to mid-
dle of nasals; 31 ) anterior ramus of para-
sphenoid broad, not keeled; 32) para-
sphenoid alae oriented at right angles to
anterior ramus, slightly overlapped lat-
erally by median rami of pterygoids; 33)
pterygoids relatively small, anterior rami
in long contact with maxillae, not reach-
ing palatines; 34) occipital condyles
large, not stalked, separated medially;
35) mandible lacking odontoids; 40) m.
depressor manclilmlae in one slip — pars
tympanictis; 41) pupil horizontal; 42)
males with median subgular vocal sac
but lacking nuptial asperities; 43) very
large parotoid gland present; 44) tongue
large, oval, free posteriorly; 45) toes free
of webbing, outer metatarsal tubercle
absent, inner of normal shape and size,
first finger shorter than second; 46) de-
velopment probably direct in nature; in
laboratory-raised clutch, the tadpoles
hatched but did not feed; had median
vent, and mouth parts greatly reduced
(Littlejohn, 1963); 47) amplexus ingui-
nal; 48) eggs laid in foam nests in bur-
rows and in holes dug in sphagnum; 49)
males 41-46, females 47-51 mm. SVL;
50) tympanum concealed.

Composition. — Monotypic, P. frosti.

Distribution. — Known only from
Mount Baw Baw, Victoria, Austraha.

Remarks. — Philoria is related to
Kyarranus and Limnodynastes. The
three genera are readily distinguished on
the basis of body glands and skeletons.
Kyarranus and Philoria are superficially
very similar in that their breeding biolo-
gies are identical, both are genera of
stubby frogs with short fingers, and the
tympani are concealed. The similarity.

in part, is probably due to adaptation to
montane environments because several
Neotropical genera living in the Andes,
bear superficial resemblance to these
Australian frogs. Kyarranus and Pluloria
probably represent independent diver-
gences from a Limnodynastes-\ike an-
cestor.

Myobatrachinae Schlegel, 1850

Myobatiachidae Schlegel, 1850:10.
Uperoliidae Giinther, 1859a: 39.
Criniae Cope, 1866:89.
Criniinae: Noble, 1931:496.
Myobatrachinae: Parker, 1940:64.

The Myobatrachinae, as presently
conceived, are restricted in distribution
to the Australo-Papuan Region. The fol-
lowing diagnostic statements are true for
all members of the subfamily Myobatra-
chinae: 1) sternum cartilaginous; 2)
vertebral shield absent; 3) transverse
processes of anterior presacral vertebrae
not greatly expanded; 4) cervical cotylar
arrangement type I; 5) cervical and sec-
ond vertebrae free; 6) cranial bones not
dermostosed; 8) sacral diapophyses di-
lated; 14) maxillary arch complete; 20)
frontoparietals not fused to prootics; 21 )
temporal arcade lacking; 30) spheneth-
moid usually divided; 35) mandible
lacking odontoids; 37) alary processes of
hyoid plate broad and wing-like; 38 ) cri-
coid cartilage divided ventrally; 39) m.
petroliyoideus anterior and m. sterno-
lu/oideus insert on the body of the hyoid
plate, near the midline; 45) first finger
shorter than second; 47) amplexus in-
guinal in known species. All Myobatra-
chinae exhibit free intervertebral discs
and non-imbricate neural arches. The
transverse processes of all vertebrae are
shortened in some genera. Metacrinia
sometimes has prezygapophyses on the
anterior end of the coccyx. The presence
of free intervertebral discs and small
prevomerine bones are the most diagnos-
tic osteological characteristics of the sub-
family. Within the subfamily, there is
an evolutionary trend toward the reduc-
tion of bone in the skull; this reduction

LYNCH: LEPTODACTYLOID FROGS

91

is reflected in the frontoparietal fonta-
nelle, size of the nasal bones, massive-
ness of the maxillary arch, divided
sphenethmoid, and reduction in the size
of the bones of the anterior palate.

Two groups of myobatrachine genera
tend to be segregated on the bases of
several characters. Uperoleia is the
only genus with vertical pupils, and the
only one lacking a frontoparietal fon-
tanelle. Glauertia and Uperoleia have
two large, compressed, metatarsal tuber-
cles— in the other myobatrachine genera,
both metatarsal tubercles are small and
not compressed, or the outer absent
(Crinia: darlingtoni, haswelli, laevisjeai,
rosea, and victoriana; and Taiidactyliis
acutirostris). The quadratojugal bones
are deep (Figs. 65, 69, and 71) in Crinia,
Glauertia, and Uperoleia, but more slen-
der in the other genera. The nasal bones
are comparatively large in Glauertia,
Metacrinia, Myohatrachtis, and Upero-
leia. In these genera, the occipital con-
dyles are larger than those of Crinia,
Pseudophryne, and Taudactylus. The
zygomatic rami of the squamosals are
relatively short in Glauertia, Metacrinia,
Myohairachus, and Uperoleia; in Crinia,
PseudopJiryne, and Taudactylus the zygo-
matic rami are longer. Loss of the teeth
of the maxillary arch is characteristic of
all genera of the subfamily, except Crin-
ia and Taudactylus. Uperoleia rugosa
(two species are included under this
name, see Littlejohn, 1968) and U. mjoe-
bergi have teeth on the maxillary arch,
but U. marmorata lacks maxillary teeth.
All myobatrachines except some species
of Crinia lack prevomerine teeth. Crinia
and Pseudophryne differ from the other
myobatrachines in having no pars scapu-
laris (m. depressor mandibulae); all
other myobatrachines have two slips in
the m. depressor mandibulae.

Myobatrachine tadpoles have labial
papillae only lateral to the tooth rows
and beak — unlike other leptodactylids,
the myobatrachines have an anterior and
a posterior interruption of the labial
papillae. The tooth rows are 1/3 to 2/3

in the known taxa. About one-half of the
species of Crinia {laevis complex) and
Glauertia and Uperoleia lay their eggs in
water and have normal tadpole develop-
ment. In these species the eggs are small
(Martin, 1968). In the Crinia signifera
complex, Metacrinia, Myobatrachus, and
Pseudophryne the eggs are large and
laid on land, part or all of the larval de-
velopment occurring in the absence of
water ( Martin, 1968 ) . Data are unavail-
able for Taudactylus, which probably
also lays its eggs on land. At least some
of the "terrestrial breeders" of the gen-
era Crinia and Pseudophryne hatch as
tadpoles when the nest is inundated and
rapidly begin feeding and behaving like
normally aquatic tadpoles. Some Crinia
and probably Metacrinia nichoUsi and
Myobatrachus gouldii have completely
terrestrial development (Martin, 1968).

Noble ( 19.30 ) named a new genus,
Indobatrachus, for Rana pusillus Owen,
a Lower Eocene frog from the Intertrap-
pean beds of Bombay in peninsular In-
dia. He considered the fossil closest to
Crinia and therefore placed it in the
group now designated as Myobatrachi-
nae. Noble argued that the presence of
a toothless bufonid in the early Cenozoic
of the northern land masses indicated a
northern origin for the group. Parker
(1940) andHecht (1963) suggested that
the remains of Indobatrachus were over-
interpreted, but failed to say why or how
they reached that conclusion. Noble's
data are not incompatible with the osteo-
logical data I have for the Recent Myo-
batrachinae. However, Indobatrachus
could also be interpreted as not being
very distant from Nesomantis and Soo-
glosstis, which may prove to be myo-
batrachines as well; these subjects are
discussed more fully in the generic ac-
count of Indobatrachus.

Crinia Tschudi,

(Fig. 65)

1838

Crinia Tschudi, 1838, Mem. Soc. Sci. Nat.
Neuchatel, 2:38 [Type-species by mono-
typy, Crinia georgiana Tschudi, 1838].

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Ranidella Girard, 1853, Proc. Acad. Nat. Sci.
Philadelphia, 6:421 [Type-species by mono-
typy, Crinia (Ranidella) si<inifcra Girard,
1853].

Pternophrynus Liitken, 1863, Vid. Meddel.
Foren., 1862:302 [Type-species by mono-
typy, Pternophrynus verrucosus Liitken,
1863].

Camariolius Peters, 1863, Mtber. k. Preuss.
Akad. Wiss., Berlin, 1863:236 [Type-species
by monotypy, Camariolius varius Peters,
1863].

Pternophryne Giinther, 1867, Ann. Mag. Natur.
Hist., (3)20:53 [Replacement name for
Pternophrynus Liitken, 1863 (preoccupied
by Pternophrynus Gill, 1863 — Osteichthys)
and hence taking the same type-species].

Diagnostic definition. — 7 ) omoster-
num present, small and elongate; 9)
maxillary arch toothed, teeth blunt, pedi-
cellate; 10) alary processes of premaxil-
lae directed posterodorsally, narrow at
base; 11) palatal shelf of premaxilla
broad, moderately deeply incised; 12)
facial lobe of maxilla deep, not exos-
tosed; 13) palatal shelf of maxilla
broadest anteriorly, narrowing posterior-

ly, small pterygoid process; 14) quad-
ratojugal deep; 15) nasals small, short,
narrowly separated medially; 16) nasals
not in contact with maxillae or ptery-
goids; 17) nasals widely separated from
frontoparietals; 18) frontoparietal fon-
tanelle large; 19) frontoparietals not
ornamented; 22) epiotic eminences obso-
lete; 23) cristae paroticae short and
stocky; carotid artery passes above skull
bones; 24) zygomatic ramus of squa-
mosal short, knob-like; 25) otic ramus of
squamosal long, not developed into an
otic plate; 26) squamosal-maxillary an-
gle about 65° if measured along main
axis of maxilla; 27) columella present;
28) prevomers widely separated medial-
ly, fragmented, dentigerous processes
present or not, bearing teeth or not; 29 )
palatines narrow, sliver-like, not reach-
ing maxillae; 30) sphenethmoid small,
divided; 31) anterior ramus of para-
sphenoid narrow, not keeled; 32) para-
sphenoid alae oriented at right angles to

Figure 65. Lateral, dorsal, and ventral views of skulls of Crinia si^nifcra. (A) KU 56243, X 8; (B-C)

KU 5636 1, X 5.

LYNCH: LEPTODACTYLOID FROGS

93

anterior ramus, shortened and widely
separated from median rami of ptery-
goids; 33) pterygoids small, anterior
rami in short contact with maxillae; 34)
occipital condyles small, stalked, widely
separated medially; 36) terminal pha-
langes knobbed; 40) m. depressor man-
dibnloe in one slip, pars tijmpanicus;
41) pupil horizontal; 42) males with
median, subgular vocal sac; nuptial as-
perities on thumb; 43) body lacking
glands; 44) tongue narrow, posterior
edge free; 45) toes lacking webbing, but
with prominent lateral fringes in has-
welli, tasmaniensls, and the signifera
complex; outer metatarsal tubercle pres-
ent in all species except darlingtoni, has-
iceUi, loevis, leai, rosea, and mctoriana;
digital tips narrow; 46) development di-
rect in some species, "normal" in others;
many species lay terrestrial eggs and
have intraoval tadpoles, if the nest is
inundated, the tadpole emerges and con-
tinues development as a free swimming
tadpole; tadpoles have dextral vent, 2/3
tooth rows, labial papillae broadly in-
terrupted anteriorly, narrowly inter-
rupted posteriorly; 47) amplexus ingui-
nal; 48) eggs small and numerous, laid
in water, or eggs small and numerous
and laid in terresti-ial situations, or eggs
large and few in number and laid in ter-
restrial situations; 49) adults small,
males 14.5-30.0, females 19.0-32.5 mm.
SVL; 50) tympanum concealed or visi-
ble externally.

Composition. — Parker (1940) recog-
I nized 13 kinds of Crinia one of which,
acutirostris, is now placed in another
genus. Moore (1961) recognized 16
species including acutirostris. Parker
(1940), Main (1957), Moore (1961),
and Martin (1968) recognized two spe-
cies groups of Crinia, but included acu-
tirostris in one of these groups. The
loevis group contains six species — dar-
lingtoni, laevis, leai, lutea, rosea, and
victoriana. The signifera group includes
eleven species — georgiana, glauerti, has-
weUi, insignifera, parinsignifera, pseud-
insignifera, riparia, signifera, sloanei.

tasmaniensis, and tinnula. As Moore
(1961) pointed out, only a small part of
Crinia signifera (sensu Parker) has been
studied for sibling species, and the num-
ber of species in this complex and the
genus should be expected to increase
significantly in the next decade.

Distribution. — Australia ( except for
most of the western Eyrean subregion),
eastern New Guinea, and Tasmania.
Three species occur on Tasmania, one of
which is endemic (tasmaniensis), and
another very wide-spread and the only
species to reach Papua (signifera, sensu
Parker).

Remarks. — Perhaps the most inter-
esting facet of the genus Crinia is the
presence of several morphologically very
similar species (signifera complex) with
different calls, breeding biology, and /or
tadpoles. In vitro crosses of these spe-
cies led to the discovery that they were
biospecies in the sense of Mayr (1963).

I have been able to study only one
species osteologically; when additional
studies are made on the skeletons of
other species of Crinia, my diagnosis
may be shown to be incomplete or in
error. Considerable variation is evident
in foot morphology and the expression of
prevomerine teeth. Parker (1940) di-
vided the genus into two groups on the
basis of the division (dentigerous ramus
separated from the part of the bone sur-
rounding the choanae) of the prevo-
merine bones. The prevomerine bones
are small but complete in most of the
species of the laevis group, and divided
in most species of the signifera group ex-
cept haswelli. Parker (1940) suggested
that hasiveUi might be the most primi-
tive species of the genus on the basis of
the pectoral girdle; the prevomerine
anatomy of haswelli is not contrary to
this suggestion. The two species groups
of Crinia are separated on the basis of
whether the skin of the venter is smooth
(laevis group) or granular (signifera
group). The species of the laevis group
lack outer metatarsal tubercles and toe
fringes, whereas the species of the sig-

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

nifera group have both outer metatarsal
tubercles and lateral fringes on the toes
(except georgiana and glanerti, which
lack toe fringes, and hasweUi, which
lacks an outer metatarsal tubercle). Al-
though the use of the presence or ab-
sence of prevomerine teeth as a major
taxonomic character has been challenged
by Main (1957), the character is useful
as a secondary character in Crinia. Well
developed prevomerine dentigerous
processes and teeth occur in leai, liitea,
rosea, and usually in laevis and victori-
ana, all of which are members of the
laevis group. The only member of the
laevis group which usually lacks prevo-
merine teeth is darlingtoni. Three spe-
cies of the signifera group usually have
well developed prevomerine dentigerous
processes and teeth (georgiana, ghuerti,
and hasicelli); the other species of the
group almost invariably lack prevomer-
ine teeth and prevomerine dentigerous
processes.

Martin (1968) pointed out the vari-
ability in breeding habits in Crinia. Two
species (lutea and rosea) have terrestrial
development with the tadpole living in
the jelly mass until metamorphosis. In
several other species (laevis group and
riparia) the eggs are laid out of water
and the tadpole usually enters water
sometime after completing early devel-
opment. In most species of the signifera
group and some of the laevis group, the
eggs are laid in water ( in small clumps )
and the developmental pattern is that
typified by Rana. Martin (1968) sug-
gested that the different patterns of ovi-
position have little taxonomic signifi-
cance in ascertaining species-group rela-
tionships.

Pseudophyryne Fitzinger, 1843
(Fig. 66)

Pseudophnjne Fitzinger, 1843, Syst. Rept., p.
31 [Type-species by original designation,
Phryniscus atistralis (paiiim) Dumeril and
Bibron, 1841 ( =Pseudophrtjne setnimar-
morata Lucas, 1892); see Parker (1940:
94) for a comment on this type-species
designation].

BufoneUa Girard, 1853, Proc. Acad. Nat. Sci. !
Philadelphia, 6:424 [Type-species by mono- '
typy, BufoneUa cnicigcra Girard, 1853].

Diagnostic definition. — 7) omoster-
num present, small and narrow; 9) max-
illary arch edentate; 10) alary processes
of premaxillae directed dorsally, broad
at base; 11) palatal shelf of premaxilla
relatively narrow, palatal process elon-
gate; 12) facial lobe of maxilla shallow;
13) palatal shelf of maxilla of moderate
width, pterygoid process small; 14)
quadratojugal shallow; 15) nasals small,
narrow, with slight median separation;
16) nasals not in contact with maxillae
or pterygoids; 17) nasals not in contact
with frontoparietals; 18) frontoparietals
fontanelle large; 19) frontoparietals not
ornamented; 22) epiotic eminences ob-
solete; 23) cristae paroticae stocky,
broad; carotid artery passes above skull
bones; 24) zygomatic ramus of squa-
mosal very short, knob-like; 25) otic
ramus of squamosal elongate, not ex-
panded into otic plate; 26) squamosal-
maxillary angle about 50°; 27) columel-
la absent; 28) prevomers absent; 29)
palatines reduced to splinter-like bones,
not in contact with maxillae, widely sep-
arated medially; 30) sphenethmoid di-
vided; 31) anterior ramus of parasphe-
noid broad posteriorly, narrowing to-
wards palatines; 32) parasphenoid alae
short, deflected posteriorly, not in con-
tact with median rami of pterygoids;
33) pterygoids small, median and pos-
terior rami short, anterior rami in short
contact with maxillae, not reaching pala-
tines; 34) occipital condyles large,
stalked, widely separated medially; 36)
terminal phalanges knobbed; 40) m. de-
pressor mandilndae in one slip — pars
tympanicus; 41) pupil horizontal; 42)
males with median, subgular vocal sac;
males lacking nuptial asperities; 43) par-
otoid, flank, and sometimes femoral
glands present; 44) tongue long, narrow,
posterior edge free; 45) toes free of web-
bing and lateral fringes, outer metatarsal
tubercle present; 46) larvae with dextral
vent, 2/3 tooth rows, labial papillae

LYNCH: LEPTODACTYLOID FROGS

95

Figure 66. (A) Lateral, (B) dorsal, and (C) ventral views of a skull of Pseiidophnjne bihroni (KU

93588). (A, X 8; B-C, x 5).

broadly interrupted anteriorly, narrow-
ly interrupted posteriorly, development
occurs in either water or in jelly mass in
terrestrial situations; 47) amplexus in-
guinal; 48) eggs laid singly in perma-
nent water or in soil or moist sphagnum
moss, number per clutch varies from 12
to over 100; 49) adults small, males 20-
28, females 20-34 mm. SVL; 50) tym-
panum absent.

Composition. — Ten species are pres-
ently recognized: australis, hibroni, co-
riacea, corrohoree, dendyi, donglasi,
guentheri, major, occidentalis, and semi-
marmorata. Parker's (1940) study is
the most recent review, although
Moore's (1961) study of the frogs of
eastern New South Wales treats sev-
eral of the species of the genus.

Distribution. — Northwestern and
southwestern Australia; and east of
the Dividing Range from east-central
Queensland to the Capital Territory;
Tasmania.

Remarks. — Pseudophryne and Crinia

are closely related and differ primarily in
maxillary dentition, loss of the middle
ear, and coloration and body glands.
The two genera were placed in different
families by taxonomists before Noble,
because Pseudophryne lacks teeth and
Crinia has teeth on the maxillary arch.
Metacrinia is superficially similar to
Pseudophryne, but differs markedly in
the skull architecture and temporal mus-
culature.

Taudactylus Straughan and Lee, 1966

(Fig. 67)

Taudactylus Straughan and Lee, 1966, Proc.
Royal Soc. Queensland, 76:63 [Type-species
by original designation, Taudactylus diurnus
Straughan and Lee, 1966].

Diagnostic definition. — 7) omoster-
num absent; 9) maxillary arch toothed,
teeth blunt, pedicellate; 10) alary proc-
esses of premaxillae directed very
slightly posterodorsally, narrow at base;
11) palatal shelf of premaxilla relatively
broad laterally, narrow medially, bearing

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

greatly elongated palatal process; 12) fa-
cial lobe of maxilla shallow; 13) palatal
shelf of maxilla of moderate width, nar-
rowing posteriorly, pterygoid process
minute; 14) quadratojugal shallow, long
and thin; 15) nasals very small, widely
separated medially; 16) nasals not in
contact with maxillae or pterygoids; 17)
nasals not in contact with frontopari-
etals; 1(S) frontoparietal fontanelle pres-
ent, small and narrow; 19) frontopari-
etals not ornamented; 22) epiotic emi-
nences obsolete; 23) cristae paroticae
short, stocky; carotid artery passing dor-
sal to skull bones; 24) zygomatic ramus
of squamosal short, thin, about one-third
length of otic ramus, therefore propor-
tionately longer than zygomatic rami of
other myobatrachines; 25) otic ramus of
squamosal long, not expanded medially
into otic plate; 26) squamosal-maxillary
angle about 55°; 27) columella present;
28) prevomers minute, fragmented, den-
tigerous rami absent, restricted to medial
edges of choanae; 29) palatines narrow,

widely separated medially; 30) sphe-
nethmoid divided; 31) anterior ramus of
parasphenoid long, narrow, reaching
level of palatines, not keeled; 32) para-
sphenoid alae short, deflected posterior-
ly, not overlapped by median rami of
pterygoids; 33) pterygoids comparative-
ly large, anterior rami in short contact
with maxillae, not reaching palatines;
34) occipital condyles small, stalked,
widely separated medially; 36) terminal
phalanges T-shaped; 40) m. depressor
mandiJmlae in two slips; 41) pupil hori-
zontal; 42) males with median, subgular
vocal sac; diffuse nuptial pad on thumb;
43) body lacking glands; 44) tongue
long, narrow, posterior edge free; 45)
toes not webbed, bearing distinct lateral
fringes, outer metatarsal tubercle absent;
46-48); 49) males 24-27, females 27-31
mm. SVL; 50) tympanum concealed.

Composition. — Monotypic, Taudac-
tijltis acutirostris ( Andersson ) .

Distribution. — Known only from four

Figure 67. Lateral, dorsal, and ventral views of the skull of Taudactylus acutirostris (KU 124233,

X5).

LYNCH: LEPTODACTYLOID FROGS

97

localities in the Great Dividing Range in
Queensland.

Remarks. — Parker (1940) included
aciitirostris in the Crinia signifera group.
He was familiar with the osteology of
the anterior palate, but did not notice
the T-shaped terminal phalanges and
the widely separated nasal bones. In
external appearance, Taudacfylus is not
especially distinctive. The pad-like digi-
tal tips and the pointed snout (which
overhangs the jaw) are the only distinc-
tive external features of the monotypic
genus. The type-species (diurmis) is not
distinguishable from Crinia acutirostris,
and therefore acutirostris is placed in the
genus Taudactylus (Straughan and
Main, 1966).

Taudactijlus acutirostris has lateral
fringes on the toes and lacks an outer
metatarsal tubercle like Crinia hasioelli;
C. haswelU differs from Taudactijlus in
having narrow digital tips knobbed
terminal phalanges), a rounded snout,
and prevomerine teeth. Although I con-

sider Crinia and Taudactijlus related, I
do not believe that any of the currently
known species of Crinia (see page 93)
will prove to be Taudactylus. Taudac-
tylus most closely resembles Crinia and
Fseudophryne osteologically but is ob-
viously different in its lack of a large
frontoparietal fontanelle.

Glauertia Loveridge, 1933

(Fig. 68)

Glauertia Loveridge, 1933, Occ. Papers Boston
Soc. Nat. Hist., 8:89 [Type-species by orig-
inal designation, Glauertia russelli Love-
ridge, 1933].

Diagnostic definition. — 7) omoster-
num present, small; 9) maxillary arch
edentate; 10) alary processes of premax-
illae directed very slightly anterodorsal-
ly; 11) palatal shelf of premaxillae of
moderate width with elongate palatal
process; 12) facial lobe of maxilla rela-
tively shallow; 13) palatal shelf of max-
illa of moderate width, pterygoid process
small; 14) quadratojugal very deep; 15)

Figure 68. Lateral, dorsal, and ventral views of a skull of Glauertia russelli ( Western Australia

Museum R22920, X 5).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

nasals moderate sized, narrowly sepa-
rated medially; 16) nasals not in con-
tact with maxillae or pterygoids; 17) na-
sals not in contact with frontoparietals;
18) frontoparietal fontanelle large; 19)
frontoparietals not ornamented; 22)
epiotic eminences poorly developed;

23) cristae paroticae short, stocky; caro-
tid artery passes dorsal to skull bones;

24) zygomatic ramus of squamosal mi-
nute; 25) otic ramus of squamosal rela-
tively long, no otic plate; 26) squamosal-
maxillary angle about 70°; 27) columella
present; 28) prevomers absent; 29) pala-
tines broad, not in contact with maxillae,
widely separated medially; 30) sphe-
nethmoid entire, not extending anterior-
ly to nasals; 31) anterior ramus of para-
sphenoid broad, lacking median keel;
32) parasphenoid alae oriented at right
angles to anterior ramus, short and
broad, widely separated from median
rami of pterygoids; 33) pterygoids rela-
tively small, anterior rami in short con-
tact with maxillae; 34) occipital con-
dyles moderately large, stalked, widely
separated medially; 36) terminal pha-
langes knobbed; 40) m. depressor man-
dibulae in two slips; 41) pupil horizon-
tal; 42) males with median, subgular
vocal sac; lacking nuptial asperities on
thumb; 43) body lacking glands or with
only very diffuse flank glands; 44)
tongue narrow, free posteriorly; 45 ) toes
one-fourth to one-half webbed, outer
metatarsal tubercle present and as large
as, and compressed like, inner metatar-
sal tubercle, digital tips narrow; 46)
larvae aquatic, not described; 47) am-
plexus inguinal; 48) eggs numerous,
small, laid in ponds; 49) adults 22-32
mm. SVL; 50) tympanum concealed.

Co77iposition. — Parker (1940) recog-
nized three species — mjoehergi, orien-
talis, and russeUi. I consider mjoehergi a
species of Uperoleia, and tentatively rec-
ognize orientaUs and russelli as specific-
ally distinct members of Glauertia.

Distrilmtion. — The two norminal spe-
cies of Glauertia are known from a half-

dozen localities in the northern terri-
tories of Australia.

Remarks. — Parker (1940) named
Glauertia orientaUs as the second species
of the genus, and placed Pseudophryne
mjoehergi Andersson in Glauertia as
well. Harrison (1927) suggested that
uijoehergi might be a synonym of Uper-
oleia marmorata, but Parker (1940) dis-
missed the suggestion with the remark
". . . Fseudophryne mjbhergi . . . appears
to be referable to the genus Glauertia."
Parker did not offer evidence for or
against this statement. Parker's ( 1940 )
diagnosis of Glauertia is clearly based
only on orientaUs and russelli. Loveridge
(1933) considered Glauertia related to
Pseudophrytie and Uperoleia, and later
(1935) stated that Glauertia was most
closely related to Pseudophryne. Parker
(1940) did not comment on the rela-
tionships of Glauertia other than "The
species of this genus appear ... to be
descended from some Criniine stock."
He apparently considered Glauertia
merely webbed Crinia. Glauertia re-
sembles Crinia only in the very large
frontoparietal fontanelle; however, the
large frontoparietal fontanelle is char-
acteristic of not only Crinia and Glau-
ertia, but also of Metacrinia, Myohatra-
chus, and Pseudophryne. In external
characters, Glauertia is most like Uper-
oleia; the two genera differ in pupil
shape, body glands, degree of digital
webbing, and in the size of the fronto-
parietal and nasal bones. The similarity
in foot morphology and in the squamosal
architecture of Glauertia and Uperoleia
strongly suggest that they are closely
related. The digits are fringed in Uper-
oleia and webbed (less than one-half)
in Glauertia — this difference is one of
degree. Uperoleia has well defined paro-
toid and flank glands whereas Glauertia
has diffuse glands along the flank. The
metatarsal tubercles in both genera are
enlarged and compressed; in no other
myobatrachine frogs is there comparable
development of the metatarsal tubercles.

Pseudophryne mjoehergi is more sim-

LYNCH: LEPTODACTYLOID FROGS

99

Figure 69. Dorsal and ventral views of skull of Uperoleia rugosa ( KU 109861, x 5).

ilar to Uperoleia rugosa than to GJauer-
tia russelli and is therefore placed in
Uperoleia. Until the pupil shape (in
life) and cranial osteology of mjoebergi
are known, this action should be re-
garded as tentative. The relationships
of the species of these two genera are
seriously in need of study; the six spe-
cies included in the two genera seem
more closely interrelated than is indi-
cated by the generic classification.

Uperoleia Gray, 1841

(Figs. 69-70)

Uperoleia Gray, 1841, Ann. Mag. Natur. Hist.,
(1)7:90 [Type-species by monotypy, Uper-
oleia inarmorata Gray, 1841].

Hijperolia Cope, 1865, Rev. Natur. Hist., 5:108
[Emendation of Uperoleia Gray, 1841,
hence taking same type-species. As pointed
out by Parker ( 1940 ) , it is best to retain
Gray's .spelling].

Diagnostic definition. — 7) omoster-
num small; 9) maxillary arch toothed or
not, if toothed, teeth blunt, pedicellate;
10) alary processes of premaxillae di-
rected dorsolaterally, wide at base; 11)
palatal shelf of premaxilla of moderate
width, palatal process elongate; 12) fa-
cial lobe of maxilla shallow; 13) palatal
shelf of maxilla of moderate width,
pterygoid process small; 14) quadrato-
jugal very deep; 15) nasals large, in
broad median contact; 16) nasals not in
contact with maxillae or pterygoids; 17)

nasals in tenuous contact with fronto-
parietals; 18) frontoparietal fontanelle
absent; 19) frontoparietals not orna-
mented or only slightly roughened pos-
teriorly; 22) epiotic eminences obsolete;
23) cristae paroticae short, stocky; caro-
tid artery lies on skull in shallow groove
median to epiotic eminences; 24) zygo-
matic ramus of squamosal minute; 25)
otic ramus of squamosal long, no otic
plate; 26) squamosal-maxillary angle
about 50°; 27) columella present; 28)
prevomers reduced to minute fragments
lying on inner edge of choanae; 29 ) pala-
tines broad, widely separated medially,
not in contact with maxillae; 30) sphe-
nethmoid usually entire, extending an-

FiGURE 70. Lateral views of the skulls of (top)
Mefacrinia nichoUsi ( KU 110332, X 4.8) and
(bottom) Uperoleia rugosa (KU 109861, X 3.2).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

teriorly beneath posterior edge of nasals;
31) anterior ramus of parasphenoid
broad, not keeled; 32) parasphenoid alae
short, deflected posteriorly, not over-
lapped by median rami of pterygoids;
33) pterygoids relatively massive, an-
terior rami in short contact with maxil-
lae; 34) occipital condyles relatively
small, not stalked, widely separated me-
dially; 36) terminal phalanges knobbed;
40) m. depressor mandihiilae in two
slips; 41) pupil vertical in life, rhom-
boidal in preserved specimens; 42) male
with median, subgular vocal sac; thumb
of male swollen in breeding season, lack-
ing nuptial asperities; 43) large paro-
toid and flank glands present; 44 ) tongue
large, round, posterior one-third free;
45) toes fringed, not webbed, outer
metatarsal tubercle present, metatarsal
tubercles enlarged, compressed, digital
tips narrow; 46) larvae with dextral
vent, 1/3 tooth rows, labial papillae
broadly interrupted anteriorly, narrowly
interrupted posteriorly; 47) amplexus
inguinal; 48) eggs small, laid singly in
ponds, marshes; over 200 eggs per
clutch; 49) males 20-29, females 21-33
mm. SVL; 50) tympanum concealed;
51) anal flap fimbriated, at least in
females.

Composition. — Loveridge (1935) and
Moore ( 1961 ) considered Uperoleia to
be monotypic. Parker (1940) recog-
nized two species, the toothed marrno-
rata and the toothless rugosa. Littlejohn
(1968) recognized two sibling species
under the specific name Uperoleia ru-
gosa. I consider Gkiuertia mjoebergi to
be a Uperoleia. Therefore, the genus
Uperoleia contains four recognized spe-
cies— marmorata, mjoehergi, and two
kinds of rugosa.

Distribution. — Littlejohn (1968) char-
acterized both species of rugosa as being
part of the "wide-ranging component."
Knowledge of the distribution of Upe-
roleia is spotty, but the genus seems to
range through all coastal areas of the
continent. It does not occur on New
Guinea or Tasmania.

Remarks. — The inclusion of mjoe-
hergi in the genus Uperoleia does not
require an expansion of the generic
definition.

Metacrinia Parker, 1940

(Figs. 70-71)

Metracrinia Parker, 1940, Novitates Zool., 42:
93.

Metacrinia Parker, 1940, Novitates Zool., 42:
93 [Type-species by original designation,
Pseudophrijne nichoUsi Harrison, 1927].

Diagnostic definition. — 7) omoster-
num small; 9) maxillary arch edentate;
10) alary processes of premaxillae di-
rected dorsally, wide at base; 11) palatal
shelf of premaxilla narrow, palatal proc-
ess elongate; 12) facial lobe of maxilla
shallow; 13) palatal shelf of maxilla nar-
row, no pterygoid process; 14) quadrato-
jugal of moderate depth; 15 ) nasals rela-
tively small, in median contact; 16) na-
sals not in contact with maxillae or
pterygoids; 17) nasals not in contact
with frontoparietals; 18) frontoparietal
fontanelle large; 19) frontoparietals not
ornamented; 22) epiotic eminences obso-
lete; 23) cristae paroticae broad, stocky;
carotid artery passes dorsal to skull
bones; 24) zygomatic ramus of squa-
mosal short, bulbous; 25) otic ramus of
squamosal long, no otic plate; 26) squa-
mosal-maxillary angle about 65°; 27)
columella present; 28) prevomers mi-
nute, fragmented, dentigerous rami
lost, widely separated medially; 29)
palatines narrow, not in contact with
maxillae, widely separated medially; 30)
sphenethmoid entire, extending anterior-
ly to middle of nasals; 31) anterior ra-
mus of parasphenoid broad, not keeled,
extending anteriorly to between pala-
tines; 32) parasphenoid alae short, de-
flected posteriorly, not overlapped later-
ally by median rami of pterygoids; 33)
pterygoids small, median rami short, an-
terior rami long but not reaching pala-
tines; 34) occipital condyles large, not
stalked, widely separated medially; 36)
terminal phalanges knobbed; 40) m. de-

LYNCH: LEPTODACTYLOID FROGS

101

Figure 71. Dorsal and xentral views of the skull of Metacrinia nichoUsi ( KU 110332, X 7).

pressor mandihtilae in two slips; 41) pu-
pil horizontal; 42) males with median,
subgular vocal sac; lacking nuptial as-
perities; 43) body lacking glands; 44)
tongue narrow, free posteriorly; 45) toes
lacking webbing or lateral fringes, outer
metatarsal tubercle present, digital tips
narrow; 46) development direct; 47-48);
49) adults less than 25 mm. SVL; 50)
tympanum concealed.

Composition. — The type-species is
the only known species.

Distribution. — W e s t e r n Australia,
west of the Darling Range and south of
Geographe Bay (Parker, 1940).

Remarks. — Metacrinia nichoUsi looks
like a drab species of Pseud opJiryne.
Parker ( 1940 ) named the genus when he
discovered that nichoUsi has a middle
ear. The genus differs from Pseudo-
phryne in the nature of the m. depressor
mandihulae, size of the nasal bones, and
the size of the zygomatic ramus of the
squamosal. The large occipital condyles
of Metacrinia suggest a relationship to
the Glauertia-U peroleia complex or to
Myohatrachus. Metacrinia apparently
has completely terrestrial development
(Parker, 1940, Martin, 1968), but obser-
vations on the life history of the species
are entirely lacking.

Myobatrachus Schlegel, 1850

(Fig. 72)

Myohatrachus Schlegel, 1850, Proc. Zool. Soc.
London, p. 9 [Type-species by monotypy,
Myobatrachus paradoxus Schlegel, 1850].

Myiohatrachus Schlegel, 1858, Handlied. Dierk.,
2:59 [Emendation of Myohatrachus Schle-
gel, 1850].

Chehjdohatrachus Giinther, 1859, Cat. Batr.
Sal. British Mus., p. 63 [Type-species by
monotypy, Breviceps gouldii Gray, 1841].

Diagnostic definition. — 7) omoster-
num absent; 9) maxillary arch edentate;
10) alary processes of premaxillae mi-
nute, knob-like; 11) palatal shelf of pre-
maxilla narrow, palatal process elongate;
12) facial lobe of maxilla minute; 13)
palatal shelf of maxilla narrow, ptery-
goid process large; 14) quadratojugal in
tenuous contact with maxilla, relatively
shallow; 15) nasals large, in median con-
tact; 16) nasals narrowly separated from
maxillae, broadly separated from ptery-
goids; 17) nasals in tenuous contact with
frontoparietals; 18) frontoparietal fon-
tanelle large; 19) frontoparietals not
ornamented; 22) epiotic eminences ob-
solete; 23) cristae paroticae short,
stocky; carotid artery passes dorsal to
skull bones; 24) zygomatic ramus of
squamosal absent; 25) otic ramus of
squamosal large, but not forming otic
plate; 26) squamosal-maxillary angle not

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

measurable; 27) columella present; 28)
prevomers absent; 29) palatines massive,
bearing odontoid ridges, articulating
with maxillae, widely separated medial-
ly; 30) sphenethmoid divided dorsally,
complete ventrally; 31) anterior ramus
of parasphenoid broad, short, not keeled;
32) parasphenoid alae short, oriented at
right angles to anterior ramus, widely
separated from median rami of ptery-
goids; 33) pterygoids small and heavy,
median rami short, anterior rami in long
contact with maxillae, reaching pala-
tines; 34) occipital condyles large, not
stalked, widely separated medially; 36)
terminal phalanges knobbed; 40) m. de-
pressor mandibidae in two slips; 41) pu-
pil horizontal; 42) males with median,
subgular vocal sacs; nuptial asperities
lacking; 43) body lacking glands; 44)
tongue long, narrow, free posteriorly;
45) toes lacking webbing and lateral
fringes, outer metatarsal tubercle pres-
ent, digital tips narrow; 46) develop-
ment apparently direct; 47-48); 49)
adults to 60 mm. SVL; 50) tympanum

concealed; 51) head disproportionately
small compared to body, limbs short.

Composition. — Monotypic; Schlegel's
paradoxus is a synonym of Gray's goxddii.

Distribution. — Western Australia.

Remarks. — Schlegel (1850) named
the type-species of MyobatracJnis with
the specific epithet paradoxus; Parker
(1940) characterized the body shape as
"globose." On first impression, Myo-
batrachus is a bizarre frog. The head is
incongruously small compared to the
size and shape of the body. The great
reduction in the size of the head is prob-
ably an adaptation to a myrmecophag-
ous livelihood. The external adaptations
are no less striking than the adaptations
of the skull. The anterior portion of the
skull is reduced in size whereas the
posterior portion of the skull is not. The
alary processes of the premaxillae are
minute (Fig. 11) and the maxillae are
very small anteriorly. The nasal bones
form an arch over the nasal capsules.
The reduction in the size and strength of
the anterior skull bones is compensated

Figure 72. Lateral, dorsal, and ventral views of skull of Myobatiachiis gouldii (KU 110333, X 7.5).

LYNCH: LEPTODACTYLOID FROGS

103

for by the enlargement of the palatines
which serve as the major supportive ele-
ment of the anterior portion of the skull.
Judging from the size of the body and
the size of the posterior portion of the
skull, Mijohatrachus was probably a
large frog with a normal-sized head be-
fore it adapted to an ant-eating habit.

Mijohairachus seems less distantly
removed from GImtertia, Metacrinia, and
Uperoleia, than from the criniine gen-
era. In spite of the reduction in the
size of the anterior cranial bones, the
skull of Mijohatrachus is primitive in
many respects. The occipital condyles
are large and massive and the nasals
are large and in median contact. The
size of the nasals, however, could more
accurately reflect a post-reduction
strengthening of the skull.

tindobatrachus Noble, 1930

Indobatrachus Noble, 1930, Amer. Mus. Novi-
tates, 401:2 [Type-species by original desig-
nation, Rana pusilla Owen, 1847].

Parker (1940) and Hecht (1963) sug-
gested that the fossils of IndohatracJms
pusilhis were "over-interpreted." With-
out actually saying so, they implied that
its familial relationships are in doubt.
The numerous figures in Noble's ( 19.30 )
paper treat most of the character com-
plexes used in family-level classification
of frogs. The bicotylar coccyx, dilated
sacral diapophyses, shortened transverse
processes of the presacral vertebrae, and
apparently free intervertebral discs all
point to myobatrachine relationships for
IndohatracJms. Noble (1930) argued
that the strongly arched clavicles indi-
cated that the pectoral girdle was arcif-
eral; strongly arched clavicles are also
found in some of the small African ranids
( Arthroleptinae), which have firmister-
nal pectoral girdles in the strict sense.
What details of the skull are preserved
are not inconsistent with the cranial
osteology of Crinia, Nesomantls, Soo-
glossus, or the arthroleptine ranids.

Before the systematic importance of
Indohatraclius can be fully appreciated,

more detailed study must be made of
the osteology of the sooglossids and the
arthroleptine ranids. The available evi-
dence is adequate to associate the fossil
with the Myobatrachinae. When our
knowledge of the osteology of the Rani-
dae is improved, the fossil may prove
significant in evaluating the primitive-
ness of some ranid subfamilies and clari-
fy the supposed relationship between
the Microhylidae and Ranidae.

Heleophryninae Noble, 1931

Heleophryninae Noble, 1931:498.
Heleophrynidae : Hoffman, 1935:2.

Heleopliryne is the sole African rep-
resentative of the Leptodactylidae. Most
authors have accorded Heleophryne sub-
familial recognition and a few have
argued that it is a monotypic family.
Noble ( 1931 ) associated it with the bu-
fonoid frogs, and has been followed by
most subsequent authors. Reig (1958)
accorded Heleophryne familial rank in
his "Superfamilia B" ( =Ranoidea of
Davis, 1935). The evidence used in
associating Heleophryne with ranoid
frogs is meager. Hoffman ( 1930 ) de-
scribed what he believed to be the pec-
toral girdle of an adult Heleophryne
regis, and contended that the pectoral
girdle of this genus was arciferal in juve-
niles and subadults, but firmisternal in
old adults. Du Toit (1930) described
the internal and external cranial anat-
omy of Heleophryne regis; in external
cranial anatomy the skull was ranid-like.
Hoffman's old adult Heleophryne was a
ranid of the genus Hylamhates, as was
the specimen du Toit used to describe
the external cranial anatomy (du Toit,
1931). Latsky (1930) reported that the
sacral vertebra of Heleophryne is di-
plasiocoelus and the thigh musculature
is intermediate between the pelobatid
and bufonid patterns. My investigation
of the vertebral column in Heleophrytie
resulted in the conclusion that the verte-
bral centra exhibit a paedomorphic con-
dition, in which the tissue of the inter-
vertebral body is not divided by invad-

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

ing arcs of connective tissue. A similar
situation was reported by Griffiths
(1960) for Nesomantis and Sooglossus.
Therefore, all the evidence thus far ad-
vanced in support of ranoid relationships
of Heleophryne is erroneous.

Although Heleophryne has several
pelobatid-like traits, it is clearly a mem-
ber of the Bufonoidea. The genus is not
clearly distinct from the primitive Aus-
tralo-Papuan Cycloraninae in that it has
a type II cervical cotylar an-angement,
fused cervical and second vertebrae, and
a vertical pupil, and lacks an outer meta-
tarsal tubercle. The relationship of the
prevomerine dentigerous processes and
choanae of Heleophryne is unique
among leptodactylid frogs, as is its tor-
rent-adapted tadpole lacking horny
beaks. Amplectic data are not available
for Heleophryne; I anticipate that it
will be found to engage in inguinal am-
plexus. The thigh musculature of Heleo-
phryne is, in several respects, inter-
mediate between that of pelobatids and
bufonoids, suggesting an independent
origin of the Heleophryninae from the

Pelobatidae, though not demanding that
this was the evolutionary course.

Because only a single genus is in-
cluded in this subfamily, I have not sep-
arated subfamily characters from generic
characters. All 50 characters are listed
in the generic account (insofar as they
are known).

Heleophryne Sclater, 1898
(Fig. 73)

Heleophrijne Sclater, 1898, Ann. South African
Mus., 1:110 [Type-species by monotypy,
HeleopJiryne purcelli Sclater, 1898].

Diagnostic definition. — 1 ) sternum
cartilaginous; 2) vertebral shield lack-
ing; 3) transverse processes of anterior
presacral vertebrae not expanded; 4)
cervical cotylar arrangement type II; 5)
cervical and second vertebrae fused; 6)
cranial bones not dermostosed; 7) omo-
sternum present, moderately large; 8)
sacral diapophyses round; 9) maxillary
arch toothed, teeth blunt, pedicellate;
10) alary processes of premaxillae di-
rected dorsolaterally, relatively broad at
base; 11) palatal shelf of premaxilla nar-

FiGURE 73. Lateral, dorsal, and ventral \'iews of skull of Hclcuplinjue nataJensis (KU 105925, X 2.5).

LYNCH: LEPTODACTYLOID FROGS

105

row, palatal process large; 12) facial
lobe of maxilla relatively shallow; 13)
palatal shelf of maxilla of moderate
width, pterygoid process large; 14) max-
illary arch complete, quadratojugal pres-
ent, shallow; 15) nasals small, widely
separated medially; 16) nasals not con-
tacting maxillae or pterygoids; 17) na-
sals widely separated from frontopari-
etals; 18) frontoparietal fontanelle large;
19) frontoparietals not ornamented; 20)
frontoparietal not fused to prootic; 21)
temporal arcade lacking; 22) epiotic
eminences obsolete; 23) cristae paroticae
long, slender; carotid artery passes dor-
sal to skull bones; 24) zygomatic ramus
of squamosal short, about as long as otic
ramus; 25) otic ramus of squamosal
short, no otic plate; 26) squamosal-max-
illary angle about 50°; 27) columella
present; 28) prevomers entire, large,
toothed, dentigerous processes lie an-
terior to choanae; 29) palatines widely
separated medially, narrow; 30) sphe-
nethmoid entire, extending anteriorly be-
neath posteromedial corners of nasals or
not reaching nasals; 31) anterior ramus
of parasphenoid broad, not keeled, ex-
tending anteriorly to point between pala-
tines; 32) parasphenoid alae oriented at
right angles to anterior ramus, broadly
overlapped laterally by median rami of
pterygoids; 33) pterygoids relatively
small, anterior rami in short contact with
maxillae, widely separated from pala-
tines; 34) occipital condyles moderately
large, not stalked, narrowly separated
medially; 35) mandible lacking odon-
toids; 36) terminal phalanges T-shaped;
37) alary processes on hyoid plate on
narrow stalks; 38) cricoid cartilage not
divided ventrally; 39) m. petrolujoideus
anterior and m. sternohyoideus insert on
lateral edge of hyoid plate; 40) m. de-
pressor mandihulae in two slips; 41) pu-
pil vertical; 42) males with median, sub-
gular vocal sac; 43) body lacking glands;

44) tongue large, posterior edge free;

45) toes at least one-half webbed, outer
metatarsal tubercle lacking, digital tips
bearing large pads, first finger shorter

than second; 46) larvae with median
vent, 4/11-4/17 tooth rows, labial papil-
lae not interrupted, beak absent; 47-48);
49) adults 35-65 mm. SVL; 50) tym-
panum visible externally.

Composition. — Poynton (1964) rec-
ognized three species — natalensis, pur-
celli, and rosei. At present, purcelli has
three recognized subspecies.

Distribution. — South Africa from the
Cape Peninsula to eastern Transvaal.

Remarks. — Excepting the consider-
able knowledge of the tadpoles, nothing
is known about the life history of Heleo-
pliryne. Pre- and post-oviposition be-
havior and oviposition sites are useful
systematic characters (at least in lepto-
dactylid classification ) .

Ceratophryinae Tschudi, 1838

Ceratophrydes Tschudi, 1838:81.
Ceratophrydidae: Cope, 1863:50.
Ceratophryinae: Parker, 1935:511.
Ceratophryiiinae: Reig, 1960a: 117.
Ceratophrynidae: Reig and Cei, 1963:181.
Ceratophryidae : Tihen, 1965:313.

The concept of a family group for
CeratopJnys and its relatives has been
advocated for more than 100 years. Dur-
ing this period the content and the rank
of the group varied considerably. Most
South American authors accord the
group familial recognition, whereas most
North American and European authors
insist that the group is at best subfamil-
ially distinct. The South American
authors consider the Ceratophryidae to
be more closely related to the Bufonidae
than to the Leptodactylidae. Noble
(1931) included the cerotaphryine gen-
era in his Pseudinae ( =Pseudidae, Cera-
tophryinae, and Telmatobiinae as used
here). Parker (1935) pointed out that
Pseudis was not related to the group but
belonged to a monotypic subfamily of
the Hylidae. He used the next oldest
family group name for the remaining
members of the group (Ceratophryinae)
The content of the gi-oup was not sig-
nificantly changed, inasmuch as it con-
tinued to embrace most of the Neotropi-
cal leptodactylid genera.

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

The following characters are diag-
nostic for the subfamily and consistent
within the included genera: 1) sternum
cartilaginous; 2) vertebral shield pres-
ent, dermostosed; 3) transverse proc-
esses of anterior presacral vertebrae
widely expanded; 4) cervical cotylar ar-
rangement type III; 5) cervical and sec-
ond vertebrae usually free, fused super-
ficially in old specimens; 6) cranial bones
dermostosed; 8) sacral diapophyses
rounded; 9) maxillaiy arch toothed,
teeth fang-like, non-pedicellate, few in
number; 14) maxillary arch complete,
maxillae and quadratojugals fused; 18)
frontoparietal fontanelle lacking; 19)
frontoparietals extensively ornamented
with dermostoic pattern; 20) frontopari-
etals fused to prootics; 21) temporal ar-
cade complete; 27) columella present;
34 ) occipital condyles large, closely jux-
taposed or confluent; 36) terminal pha-
langes knobbed; 37) hyoid plate lacking
alary processes, cornu very broad; 38)
cricoid cartilage not divided ventrally;
39) m. petroliijoicleiis anterior and m.
sternohyoideiis insert on lateral edge of
hyoid plate; 46) larvae with median
vent, labial papillae not interrupted; 47)
amplexus axillary; 48) eggs laid in gelat-
inous masses in water, eggs small and
numerous.

Three Recent genera are usually in-
cluded in this subfamily — Ceratophrys,
Chacophrys, and Lepidobatrachus. The
genus Chacophrys is not considered
valid by me, because the diagnostic char-
acteristics are all expressions of the pae-
domorphic skeleton of the type-species,
C. pierotti; Clmcoplirys is here placed as
a synonym of Ceratophrys.

Boulenger (1882, 1919) and Cochran
(1955) included members of several
genera in their Ceratophrys. Boulenger
combined the species of Ceratophrys,
Lepidobatrachus, Odontophrynus, and
Stombus in his Ceratophrys. Neither
author nomenclatorally recognized the
Ceratophryinae, although Cochran (1955)
used the term "broad-headed leptodac-
tylids" in the sense of Miranda-Ribeiro's

(1926) Ceratophrydidae. Boulenger and
Cochran recognized two groups of Cera-
tophrys, one with a dorsal shield {Cera-
tophrys in the strict sense) and one
lacking the shield (Stombus auctorum).

The genera of the "broad-headed lep-
todactylids" (the Ceratophryinae con-
cept of Boulenger, Cochran, and Mi-
randa-Ribeiro) were separated on the
basis of the development of a fleshy
"horn" on the upper eyelid and the de-
gree of webbing of the toes. The genera
included in this group by previous au-
thors are placed in two subfamihes by
me, the Ceratophryinae and Telmato-
biinae. The genera belonging to the lat-
ter subfamily are placed in three tribes.
The subfamihal and tribal separation of
these genera is supported by osteological
data. Cei (1965) and Cei, Erspamer,
and Roseghini ( 1968 ) demonstiated that
Ceratophrys and Lepidobatrachus dif-
fered from Odontophrynus and Sto^nbus
auctorum in skin proteins, and that the
latter two genera were like leptodacty-
lids insofar as skin proteins are con-
cerned. Cei ( 1965) separated the Lepto-
dactylidae, as here used, into two fami-
lies, the Ceratophryidae and the Lepto-
dactylidae, but Cei, Erspamer, and
Roseghini (1968) included Odontophry-
nus in the Ceratophrydidae.

Limeses ( 1964, 1965 ) presented data
that she regarded as suggestive of a
closer alliance between bufonids and
ceratophryids than between the latter
and leptodactylids. She was unaware of
the variability of the m. depressor man-
dibulae in the Leptodactylidae. Her
data perhaps are useful in analyzing
trends within subfamilies but they are
not convincing enough to serve as diag-
nostic characteristics of subfamilial or
familial classifications.

The Ceratophryinae occurs over
much of South America east of the
Andes and south to the Chacoan region
of Argentina. Ceratophrys stolzmanni
occurs in the arid region of Pacific Ecua-
dor and adjacent Peru, and one or two
species of the genus inhabit the Carib-

LYNCH: LEPTODACTYLOID FROGS

107

bean versant of the Guyana massif and
the Santa Marta Range of Colombia.

Ceratophrys Wied, 1824

(Figs. 74-75)

Phnjnocenis Rafinesque, 1815, Anal. Nat., p. 78
[Nornen imdinn].

Ceratophrys Wied, 1824, Isis (Okeii), 1824:
672 [Type-species by monotypy, Cerato-
phrys varius Wied, 1824].

Stombus Gravenliorst, 1825, Isis (Oken), 1825:

920 [Type-species by present designation,
Rana cornuta Linne, 1758].

Phrynoceros Bibron m Tscliudi, 1838, Classif.
Batr., p. 82 [Type-species by monotypy,
Phrynoceros vaillanti Bibron, 1838].

Trigonophrys Hallowell, 1856, Proc. Acad. Nat.
Sci. Philadelphia, 8:298 [Type-species by
monotypy, Trigonophrys rugiceps Hallowell,
1856].

Chacophrys Reig and Limeses, 1963, Physis,
24:125 [Type-species by original designa-
tion, Ceratophrys pierotti Vellard, 1948].

Figure 74. (A-C) dorsal, ventral, and lateral views of skull of an adult Ceratophrys aurita (KU 98129,
X 1). (D-F) lateral, dorsal, and xentral views of skull of a post-metamorphic (stage 46) C. ornata

(UMMZ 109753, x 2.5).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 75. (A) lateral and (B) dorsal views of cleared and stained skull of a tadpole (Stage 41) of

Cemtophnjs ornata (UMMZ 98834, X 1.4).

Diagnostic definition. — 7) omoster-
num very large; 10) alary processes of
premaxillae directed posterodorsally,
wide at base; 11) palatal shelf of pre-
maxilla lacking; 12) facial lobe of maxil-
la very deep; 13) palatal shelf of maxilla
not evident anteriorly, forming large
pterygoid process posteriorly which is
fused to pterygoid in adult; 15) nasals
large, fused medially; 16) nasal fused to
maxilla, nasal-pterygoid relationship not
evident in adult; 17) nasals broadly
fused to frontoparietals in adult; 21)
temporal arcade forming supratemporal
fenestrae, temporal arcade notched pos-
teriorly; 22) epiotic eminences promi-
nent, concealed dorsally by temporal
arcade; 23) cristae paroticae long, nar-
row; carotid artery enclosed in dermo-
stosed cranial bones; 24) zygomatic ra-
mus of squamosal fused with squamosal
process of maxilla, forming broad post-
orbital bar; 25) otic ramus of squamosal
long, expanded medially into broad otic
plate; 26) squamosal-maxillary angle 40-
50°; 28) prevomers large, entire, toothed,
fused to sphenethmoid; 29) palatines not
distinguishable in adult, palatine area
bearing large odontoids on odontoid
ridge; 30) sphenethmoid entire in post-
metamorphic specimens; 31 ) anterior ra-
mus of parasphenoid narrow, long, not

keeled; 32) parasphenoid alae oriented
at right angles to anterior ramus, broad-
ly overlapped laterally by median rami
of pterygoids; 33) pterygoids large, no
ventral flange, extent of anterior rami
not evident in adults; 35) mandible bear-
ing small odontoids at symphysis, rela-
tively sharp odontoidal ridge along most
of angular; 40) m. depressor mandibidae
in two slips dorsally, ventrally in one
large muscle block; 41) pupil horizon-
tal; 42) males with median, subgular
\'Ocal sac; nuptial excresences on thumb;
43) body lacking glands; 44) tongue
large, posterior edge free; 45) toes ba-
sally webbed, outer metatarsal tubercle
present, inner metatarsal tubercle spade-
like, digital tips narrow, first finger long-
er than second; 46) larval tooth rows
6/7, horny beak large; 49) adults 40-170
mm. SVL; 50) tympanum visible ex-
ternally or concealed.

Composition. — The following nomi-
nal species are included in the genus:
aiirita (dorsata and varia are synonyms),
calcarata, cormita, ornata, pierotti, stolz-
manni, and testudo. Dr. Oswaldo Reig
is currently engaged in a revision of this
genus.

Distribution. — South America east of
the Andes and south to the Argentine
Chaco; dry Pacific lowlands of Ecuador

LYNCH: LEPTODACTYLOID FROCS

109

and adjacent Peru; Santa Marta Moun-
tains of Colombia; Caribbean slopes of
Andes of Venezuela.

Remarks. — The skulls of adult Cera-
tophnjs contain only three pairs of bones
that are not fused into an akinetic mass,
the columellae, premaxillae and septo-
maxillae. The septomaxillae are propor-
tionately minute. The premaxillae of
ceratophryines are unique in that the
palatal shelf is lost (Fig. 11). In dry
skeletons a calcified ball is often found
lying behind the upper-most part of the
alary process of the premaxilla; this ball
apparently is formed from the alary
cartilage. Some of the bones of the skull
of Ceratophnjs (nasals, frontoparietals,
squamosals, and maxillae) are heavily
encrusted with exostoic processes, and
the suture lines are obliterated. The
suture lines in the rest of the skull are
not evident in adults, presumably owing
to the extensive calcification of cartilage
in the skull, and the concommitant fu-
sion of all skull bones (except the colu-
mellae, premaxillae, and septomaxillae).
In order to clarify the relationships of
the topographic regions of the skull, an
advanced tadpole (stage 41, fide Gos-
ner, 1960) and a post-metamorphic spec-
imen (stage 46) were cleared and
stained. The tadpole (Fig. 75) is in an
extremely advanced osteological condi-
tion for a bufonoid tadpole. The nasals,
maxillae, premaxillae, and septomaxillae
are relatively well developed, and teeth
for the maxilla are evident. There is a
small center of ossification in the area of
the prootic.

The post-metamorphic specimen is
likewise precociously developed osteo-
logically (Fig. 74). The juvenile skele-
ton is described below, and the descrip-
tive phrases are arranged in the same
manner as a diagnostic definition. The
description is based on UMMZ 109753.
1) sternum a cartilaginous plate; 2) ver-
tebral shield absent; 3) transverse proc-
esses of anterior presacral vertebrae
widely expanded; 4) cervical cotylar ar-
rangement type III; 5) cervical and sec-

ond vertebrae not fused; 6) cranial bones
not involved in dermostosis; 7) omoster-
num large; 8) sacral diapophyses round-
ed; 9) maxillary arch toothed, teeth
long, not pedicellate; 10) alary processes
of premaxillae directed posterodorsally;
11) palatal shelf of premaxilla absent
except for small palatal process; 12) fa-
cial lobe of maxilla deep, heavily exo-
stosed; 13) palatal shelf of maxilla ab-
sent anteriorly, posteriorly represented
by large pterygoid process; 14) maxillary
arch complete, squamosal process large,
completing squamosal-maxillary bridge;
15) nasals large, in broad median con-
tact; 16) nasals in broad articular con-
tact with maxillae, not contacting ptery-
goids; 17) nasals and frontoparietals in
broad contact; 18) frontoparietal fon-
tanelle very small; 19) frontoparietals
exostosed, lacking crests; 20) frontopari-
etals not fused with prootics; 21 ) fronto-
parietals are expanded posterolaterally
to cover median portion of cristae paroti-
cae and form temporal arcade; supra-
temporal fenestrae present; 22) epiotic
eminences prominent; 23) cristae paro-
ticae long, slender; carotid artery en-
closed in canal; 24) zygomatic ramus of
squamosal elongate, forming upper half
of squamosal-maxillary bridge; 25) otic
ramus of squamosal large with large otic
plate which is in contact with fronto-
parietal; 26) squamosal-maxillary angle
50°; 27) columella present; 28) prevo-
mers relatively small, entire, separated
medially, dentigerous processes lie pos-
terior to choanae; 29) palatines long,
separated medially; 30) sphenethmoid
entire, not visible dorsally; 31) anterior
ramus of parasphenoid long and narrow;
32) parasphenoid alae oriented at right
angles to anterior ramus, broadly over-
lapped laterally by median rami of ptery-
goids; 33) pterygoids moderate-sized,
anterior rami long, not reaching pala-
tines; 34) occipital condyles large, close-
ly approximated, articular surfaces sep-
arate; 35) mandible lacking odontoids;
36) terminal phalanges knobbed. In

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

contrast to the adult condition, the sep-
tomaxillae are large.

The principal difficulty surrounding
Ceratophnjs has been the definition of
and, consequently, the composition of,
the genus. Few authors confused Lepi-
dohatrachus or Odontophnjnus with
Ceraiophnjs, but many non-South Amer-
ican authors failed to perceive the many
differences between Ceratophnjs and the
generic group currently referred to un-
der the invalid generic name Stombus
Gravenhorst, 1825.

Gravenhorst (1825) proposed the
generic name Stombus and included
three species in the genus ( Rana cornuta
Linne, R. megastoma Spix, and R. scu-
tata Spix); he did not designate a type-
species. He later (1829) published an-
other paper with the combination Stom-
bus boiei; most authors have cited this
paper as the one in which the generic
name was proposed and cite boiei as the
type-species; Cope (1866) designated
boiei as the type-species. South Ameri-
can herpetologists, unlike most Euro-
pean and North American herpetologists,
have realized that the species similar to
Ceratophrys cornuta are clearly generi-
cally distinguishable from those similar
to Stombus boiei. However, the South
American authors have used the invalid
generic Stombus for this group. Of the
three species originally included in
Stombus, two (cornuta and megastoma)
are conspecific and are members of the
genus Ceratophrys; the third, scutata, is
an older specific name for the type-spe-
cies of a hylid genus, Hemiphractus
spixii Wagler, 1828. Selection of the
type-species of Stombus should depend
on the avenue of least instability. If
scutata is selected, then Stombus is the
valid generic name for the hylid genus
presently called Hemiphractus; if either
cornuta or megastoma is selected, then
Stombus is a synonym of Ceratophrys.
In either case, the generic name for the
group now called Stombus must change.
The hylid generic name is entrenched in
the literature, and any attempt to change

it would promote instability. Therefore,
Rami cornuta Linne, 1758, is selected as
the type-species of Stombus Graven-
horst, 1825. The generic group previ-
ously called Stombus has only one pri-
mary synonym, Proceratophrys Miranda-
Ribeiro, 1920, which is now the valid
generic name (see pp. 133-34). Cerato-
phrys and Proceratophrys differ in the
extent of the cranial bones, dermostosis
of the cranial bones, presence of a verte-
bral shield, expansion of the transverse
processes of the vertebrae, the develop-
ment of supernumerary plantar tuber-
cles, webbing of the toes, and tadpole
morphology. Proceratophrys is a genus
of the subfamily Telmatobiinae and the
tribe Odontophrynini.

Reig and Limeses (1963) named
Chacophrys for Ceratophrys pierotti and
distinguished the genus from Cerato-
phrys solely by an assortment of charac-
ters reflecting the less extensive ossifica-
tion of pierotti. The skeletal characteris-
tics of Chacophrys are identical with
those of the post-metamorphic (stage
46) Ceratophrys ornata (see fig. 74 and
description on pp. 107-8). I am not
recognizing Chacophrys, because its dis-
tinction rests on paedomorphic charac-
ters. The characters given by Vellard
( 1948) and Reig and Limeses (1963) for
pierotti do not distinguish that taxon
from juvenile C. ornata, but I recognize
pierotti as a valid species, because Jose
Cei assures me that it is different from
ornata. Furthermore, the number of the
chromosomes in the two species is strik-
ingly different. I am not familiar with
either species in the field.

Lepidobatraclius Budgett, 1899

(Fig. 76)

Lepidohatrachits Budgett, 1899, Quart. J. Micr.
Sci., (2)42:329 [Type-species by present
designation, Lepidobatraclius asper Budgett,
1899].

Diagnostic definition. — 7) omoster-
num absent; 10) alary processes of pre-
maxillae directed posterodorsally, long
and narrow; 11) palatal shelf of pre-

LYNCH: LEPTODACTYLOID FROGS

111

maxilla lacking except for small palatal
process; 12) facial lobe of maxilla deep,
exostosed, in broad contact with nasal;
13) palatal shelf of maxilla lacking an-
teriorly, pterygoid process developed
posteriorly; 15) nasals in median contact,
small; 16) nasals in broad contact with
maxillae, narrow contact with squa-
mosals below eye, not in contact with
pterygoids; 17) nasals in broad contact
with frontoparietals; 21) supratemporal
fenestrae absent, temporal notch absent;
22) epiotic eminences absent; 23) cris-
tae paroticae broad; carotid artery en-
closed in bony canal; 24) zygomatic ra-
mus of squamosal contacting maxilla
and nasal below eye; 25) otic ramus of
squamosal short, but expanded medially
into broad otic plate which covers crista
parotica; 26) squamosal-maxillary angle
35-55°; 28) prevomers entire, toothed,
separated medially, dentigerous proc-
esses situated between posterior borders
of choanae; 29) palatines absent; 30)
sphenethmoid entire, not visible dorsally;
31) anterior ramus of parasphenoid nar-
row, lacking median keel; 32) parasphe-
noid alae oriented at right angles to an-
terior ramus, broadly overlapped lateral-
ly by median rami of pterygoids; 33)
pterygoids massive, anterior rami not

reaching palatine area; 35) mandible
lacking odontoids; 40) m. depressor
tnandi]?idae in single slip — pars tijni-
panicus; 41) pupil vertical; 42) males
with median, subgular vocal sac, nuptial
excrescences on thumb; 43) body lack-
ing glands; 44) tongue large, posterior
edge free; 45) toes two-thirds to three-
fourths webbed, outer metatarsal tuber-
cle lacking, inner metatarsal tubercle
spade-like, digital tips narrow, first fin-
ger as long as second; 46) larvae lacking
horny beak and denticles; 49) males 46-
98, females 44-123 mm. SVL; 50) tym-
panum visible externally.

Composition. — Three species are rec-
ognized— asper {salinicola is a synonym),
laevis, and Uanensis. The genus was re-
vised by Reig and Cei (1963) and Bar-
rio (1968).

Distribution. — The Gran Chaco of
Argentina and Paraguay.

Remarks. — Boulenger (1919) synony-
mized LepidobatracJnis with Cerato-
phyrs. With the discovery of the tadpole
of laevis, Parker (1931) suggested that
when the anatomy of the adult was more
fully known, the genus Lepidobatrachus
would be recognized. Most subsequent
authors continued to use Lepidobatra-
chus as a valid generic name probably

Figure 76. Lateral, dorsal, and ventral views of skull of Lepidohatrachus aspcr ( KU 80783, X 2).

No attempt was made to illustrate the dermosteotic pattern.

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

because Nieden ( 1923 ) recognized the
genus. Nieden ceased to gather Htera-
ture data in 1915 and pubUcation of his
work was delayed by World War I; it
seems likely that he never saw Boulen-
ger's ( 1919 ) paper. Subsequently ade-
quate data have accumulated to substan-
tiate the generic distinction of Lepido-
hatrachus (Reig, 1960b, Reig and Cei,
1963, and Cei, 1968a ) . Lepidohotrachus
differs from Cerafoplirys in hax'ing a
vertical pupil, no outer metatarsal tuber-
cle, greater webbing of the toes, no
supratemporal fenestrae, and no horny
beak or denticles in the larvae. Several
published illustrations of Lepidohatra-
cJ}US illustrate a more or less rhomboidal
pupil (Cochran, 1961a, Reig and Cei,
1963), whereas others illustrate a verti-
cal slit (Cei, 1955).

fWawelia Casamiquela, 1963

Wawelia Casamiquela, 1963, Ameghiniana, 3:
144 [Type-species by original designation,
Wawelia gerholdi Casamiquela, 1963].

Casamiquela (1963) named Waivelia
gerholdi on the basis of a single, incom-
plete specimen of an Upper Miocene
frog from Argentina. The fossil consists
of a small piece of the dermostosed skull
roof, parts of the pectoral girdle, the
complete vertebral column, the left
ilium, and left leg. Casamiquela placed
the fossil in the Ceratophrydidae and
considered it allied to Ceratophnjs. Cei
( 1968a ) considered Wawelia to be di-
rectly in the phyletic line of Lepido-
hotrachus, but presented no data sup-
porting his conclusion.

The widely expanded transverse
processes of the anterior presacral verte-
brae of Waicelia are exactly like those of
Ceratophrys and Lepidohatrachus (see
Fig. 32). The characters evident on the
ilium of Wawelia are not sufficiently
diagnostic to separate the ilia of Wawe-
lia and the other ceratophryines. The
skull of Wawelia was not preserved; it
is therefore not possible to tell if the fos-
sil had a supratemporal fenestra. Watve-
lia is tentatively recognized as a valid

genus, although it is not possible to dis-
tinguish the fossil from either Cerato-
phrys or Lepidohatrachus.

Telmatobiinae Fitzinger, 1843

Telmatobii Fitzinger, 1843:31.
Hylodidae Giinther, 1859a: 90.
Alsodina Mivart, 1869:290.
Cacotina Mivart, 1869:290.
Grypiscina Mivart, 1869:295.
Telmatobiidae: Miranda- Ribeiro, 1926:14.
Pseudinae (part): Noble, 1931:499.
Ceratophryinae (part): Parker, 1935:511.
Telmatobiinae Vellard, 1951:21.
Cyclorhamphinae Lutz, 1954:175.
Eleutherodactylinae Lutz, 1954:175.
Calyptocephalellinae Reig, 1960a: 126.
Batrachylinae Gallardo, 1965:83.
Cycloramphinae: Gallardo, 1965:84.

The Telmatobiinae is the largest
leptodactylid subfamily and contains one
fossil and 24 Recent genera. I divide the
Telmatobiinae into five tribes — Telma-
tobiini, Alsodini, Odontophrynini, Gryp-
iscini, and Eleutherodactylini. The gen-
era of the Telmatobiini and Alsodini are
the most primitive of the Telmatobiinae
and are distributed in temperate South
America. These two tribes and the
Odontophrynini exhibit orthodox breed-
ing habits in that the eggs are laid in
water (except Batrachyla), and an aquat-
ic tadpole stage is present (all genera).
In the more advanced genera (tribes
Grypiscini and Eleutherodactylini) there
is a decided tendency towards direct
development.

The genera of the Telmatobiini and
some of the genera of the Alsodini and
Odontophrynini share several characters
with the Cycloraninae and Heleophryni-
nae: cervical cotylar arrangement type
II, occipital condyles large and closely
approximated, a tendency for the sacral
diapophyses to be dilated, comparatively
short transverse processes of the pos-
terior presacral vertebrae (relative to
those of the advanced genera), and
heavy-boned skulls. In addition, two
primitive non-osteological characters,
which occur in many of the old World
leptodactylids, are found in some of the
genera of these tribes. These characters

LYNCH: LEPTODACTYLOID FROGS

113

are vertical pupils and the absence of
outer metatarsal tubercles. The primi-
tive Neotropical genera also have medi-
an vents in the tadpoles. The frogs of
the Eleutherodactylini, the most success-
ful tribe, range north through Central
America and the West Indies to the
southern United States. The diversity of
morphology and biology of the genera of
this subfamily is reflected in the few
characters that are common to the 24
Recent genera. The following charac-
teristics are consistent in all of the 24
genera of the Telmatobiinae: 1) sternum
cartilaginous; 2) vertebral shield lack-
ing; 3) transverse processes of anterior
presacral vertebrae not widely expand-
ed; 35) mandible lacking odontoids;
38) m. petrolnjoideiis anterior and m.
sternohijoideus insert on lateral edge of
hyoid plate; 48) eggs laid in water, in
terrestrial situations, or in bromeliads.
When present, the maxillary teeth are
blunt and pedicellate 9).

Telmatobiini Fitzinger, 1843

Telmatobii Fitzinger, 1843:31.
Telmatobiidae: Miranda-Ribeiro, 1926:14.
Telmatobiinae Vellard, 1951:21. Burger, 1954:

195-96. Gallardo, 1965:83.
Calyptocephalellinae Reig, 1960a: 126.

The tribe Telmatobiini is used for
five genera (Batrachophnjnus, Caiidiver-
hera, Neoprocoeki, Telmatohius, and
TelmotoJnifo). The group is distributed
in the Ancles from southern Ecuador to
southern Chile, and in the lowlands of
southern and central Chile. Oligocene
and Miocene records of the group are
known from central Argentina. At first
glance, the group seems to be highly
heterogeneous, but the genera can be
arranged in a linear series in which the
adjacent genera are more closely related
to each other than to any non-adjacent
genus: Caudiverhera — Telmatobufo —
Telmatohim — Neoprocoela — Batra-
chophnjnus.

The genera of the Telmatobiini share
the following diagnostic characters
which are not repeated in the generic

accounts: 4) cervical cotylar arrange-
ment type II; 14) maxillary arch com-
plete; 20) frontoparietals not fused
to prootics; 36) terminal phalanges
knobbed; 42) males with nuptial asperi-
ties on thumb; some species of Telma-
tohius also have patches of small spines
on the chest; 47) amplexus axillary; 48)
eggs numerous, small, laid in gelatinous
masses in ponds and sloughs. All gen-
era of the tribe have massive and
strongly curved clavicles (see Fig. 34)
and large omosterna and sterna. The
sternum tends to calcify in old adults of
all of the genera.

Caudiverhera is the most divergent
genus of the tribe and most of its unique
characters are reflections of the casqued
head. Unlike the other genera of the
tribe, Caudiverhera lacks a frontopari-
etal fontanelle and has a large otic plate
on the squamosal.

Fitzinger (1843) included Tehnato-
hius in this family group. Vellard ( 1951 )
and Gallardo (1962) included only
Batrachophrynus and Tehnatobius. Gal-
lardo ( 1962 ) considered Telmatobufo a
synonym of Aruncus and most closely re-
lated to Caudiverhera. Schmidt (1952)
suggested that Telmatobufo was closely
related to Telmatohius. Schaelfer (1949)
considered the Oligocene Neoprocoela
most closely allied to Batrachophrynus
and Telmatohius. Tihen (1962b) erro-
neously synonymized Neoprocoela with
Bufo. Neoprocoela is regarded by me as
a direct ancestor to Batrachophrynus.

Barbour and Noble (1920) provided
the current generic separation of Cyclo-
ramphus and Telmatohius. Earlier au-
thors had confused the two genera and
regarded them as very closely related
( Boulenger, 1907, Miranda-Ribeiro,
1920). Because some species of Cyclo-
ramphus have extensive webbing of the
toes and Telmatohius has fully webbed
toes, many modern herpetologists regard
them related. The differences between
the two genera are so trenchant that I
place them in different tribes. Osteo-
logically, there is little in common be-

114

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

tvveen the two, and the only external
characters in which they resemble one
another are the amount of webbing of
the toes (not all species of Cijdoram-
phus) and horizontal pupils.

Caudiverbera Laurenti, 1768

(Figs. 77-78)

Caudiverbera Laurenti, 1768, Synop. Rept., p.
43 [Type-species by monotypy, Caudiver-
bera peruviana Laurenti, 1768 ( zizLacerta
caudiverbera Linne, 1758)].

Peltocephalus Bibron, in Tschudi, 1838, Classif.
Batr., p. 81 [Type-species by monotypy,
Peltocephalus quoyii Bibron, in Tschudi,
1838; preoccupied by Peltocephalus Dmne-
ril and Bibron, 1835 (Reptilia: Chelonia)].

Cahjptocephalus Dumeril and Bibron, 1841,
Erpetologie generale, 8:447 [Type-species
by monotypy, Cahjptocephalus gayi Dume-
ril and Bibron, 1841; preoccupied by
Cahjptocephalus Gray, 1832 (Insecta:
Coleoptera ) ] .

Cahjptocephala Nieden, 1923, Das Tierreich,
46:371 [Replacement name for Cahjpto-
cephalus Dumeril and Bibron, 1841 (pre-
occupied) and hence taking same type-
species; preoccupied by Cahjptocephala De-
jean, 1835 (Insecta: Coleoptera)].

Cahjptocephalella Strand, 1928, Ark. Natur-
gesch., 92A:55 [Replacement name for
Cahjptocephalus Dumeril and Bibron, 1841
(preoccupied) and for Cahjptocephala
Nieden, 1923 (preoccupied) and hence
taking same type-species].

Eophractus Schaeffer, 1949, Bull. Amer. Mus.
Nat. Hist., 93:49 [Type-species by original
designation, Eophractus casamaijorensis
Schaeffer, 1949].

Gigantobatrachus Casamiquela, 1959, Rev.
Asoc. Geol. Argentina, 13:174 [Type-species
by original designation, Gigantobatrachus
parodii Casamiquela, 1959].

Diagnostic definition. — 3) transverse
processes of anterior presacral vertebrae
very slightly expanded, those of pos-
terior presacral vertebrae somewhat
shortened; 5) cervical and second verte-
brae free; 6) cranial bones involved in
extensive dermostosis; 7) omosternum
very large; 8) sacral diapophyses dilated;
9) maxillary arch toothed, teeth elon-
gate, pedicellate; 10) alary processes
of premaxillae directed posterodorsally,
relatively broad at base; 11) palatal shelf
of premaxilla relatively broad, .slightly
indented, palatal process short; 12) fa-

FiGURE 77. Lateral and dorsal views of skull of
Caudiverbera caudiverbera (FMNH 9703, X I)-

cial lobe of maxilla deep, heavily exo-
stosed; 13) palatal shelf of maxilla of
moderate width, pterygoid process large;
15) nasals large, in broad median con-
tact; 16) nasals in broad contact with
maxillae, separated from pterygoids; 17)
nasals in broad contact with frontopari-
etals; 18) frontoparietal fontanelle lack-
ing; 19) frontoparietals exostosed, exten-
sive denticulate ornamentation; 21) tem-
poral arcade complete, not notched, su-

FiouHE 78. Ventral view of skull of Caudi-
verbera caudiverbera (FMNH 9703, X 1)-

LYNCH: LEPTODACTYLOID FROGS

115

pratemporal fenestra absent; 22) epiotic
eminences not apparent; 23) cristae
paroticae broad and long; carotid artery
enclosed in bony canal; 24) zygomatic
ramus of squamosal large, in broad con-
tact with squamosal process of maxillary
forming complete post-orbital bridge;
25) otic ramus of squamosal large, ex-
panded medially into broad otic plate
which is in articular contact with fronto-
parietal; 26) squamosal-maxillary angle
about 20° in adults, larger in juveniles;
27) columella present; 28) prevomers
relatively small, entire, toothed, narrow-
ly separated medially; 29) palatines
broad, separated medially by width of
parasphenoid, bearing numerous odon-
toids in large individuals; 30) spheneth-
moid large, entire, extending anteriorly
to middle of nasals, completely con-
cealed dorsally by nasals and frontopari-
etals; 31) anterior ramus of parasphe-
noid long, narrow, not keeled, extending
anteriorly between palatines to level of
dentigerous processes of prevomers; 32)
parasphenoid alae large, broad, oriented
at right angles to anterior ramus, broadly
overlapped laterally by median rami of
pterygoids; 33) pterygoids relatively
large, anterior rami not reaching pala-
tines, small ventral Hange on body of
pterygoids; 34) occipital condyles large,
narrowly separated medially; 37); 40);
41) pupil vertical; 42) male with medi-
an, subgular vocal sac; 43) body lacking
glands; 44) tongue large, circular, not
notched, posterior edge free; 45) toes
three-fourths webbed, outer metatarsal
tubercle lacking, inner metatarsal tuber-
cle not spade-like, digital tips narrow,
first finger as long as second; 46) larvae
with median vent, 3/3 tooth rows, labial
papillae interrupted anteriorly; 49)
adults usually 80-220 mm. SVL; fossils
are known to reach a length of more
than 300 mm. SVL; 50) tympanum visi-
ble externally.

Composition. — One Recent and one
fossil species are recognized here. Elev-
en names have been proposed for the
single Recent species: caudiverbera

Linne, 1758, peruviana Laurenti, 1768,
cristatus Daudin, 1802, feuiUaei Dumeril
and Bibron, 1836, quoyii Bibron, 1838,
gayi Dumeril and Bibron, 1841, ater
Philippi, 1902, coxi Philippi, 1902, rufa
Philippi, 1902, canqueli Schaeffer, 1949,
and parodii Casamiquela, 1959. The last
two names were proposed for Oligocene
and Miocene fossils respectively. Eo-
phractus casammjorensis Schaeffer is
tentatively recognized as a valid species
of Caudiverbera although it bears strik-
ing resemblance to the types of Giganto-
bat radius parodii. Caudiverbera casa-
mmjorensis is known only from the Low-
er Eocene of Chubut, Argentina. Fossils
that are assignable to the Recent species
are known from the Upper Oligocene
and Middle and Upper Miocene of Chu-
but, Rio Negro, and Santa Cruz, Argen-
tina.

Distribution. — In Recent times, the
species is known from localities in Chile
between 30° and 42° S latitude. The
fossils are known from central Argentina.

Remarks. — Myers (1962) pointed out
that the correct generic and specific
name for this large Chilean frog was
Caudiverbera caudiverbera. Cei (1962a)
and Gorham (1966) objected to this
usage, arguing that Caltjptocephalella
should be used to promote stability.
When "stability" is used as an argument
for preserving or rejecting a name, some
effort should be made to document the
implied statement that this name or that
name has been more frequently used.
Because the species is restricted in dis-
tribution, relatively little literature on it
has accumulated in the past 50 years.
The majority of "name usages" for the
species are simple inclusions in lists of
one sort or another. Such usages are not,
in my opinion, equal in weight to a dis-
cussion of biology, systematics, or no-
menclature. Fewer than a half-dozen
significant papers have appeared in the
last 50 years treating this species; in
these, the generic names used were Ca-
hjptocephalu, Cahjptocephalella, and
Caudiverbera. Thus, an argument based

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

on "stability" is capricious. Myers'
(1962) arguments are irrefutable; Coudi-
verhera is here used as the valid generic
name, and Linne's caudiverhera is used
in preference to Dumeril and Bibron's

Eophracfus and Gigantohatrachus
are here placed in the synonymy of
Caudiverhera for the first time. Both
genera were separated from Caudiver-
bera on the basis of dermostoic patterns,
and Gigantohatrachus further separated
on the basis of size. Eophractus was
diagnosed, in part, on the presence of a
shallow groove on the maxilla. This
groove is present in the largest specimen
of Caudiverhera that I have examined,
but not in the four smaller specimens.
In the larger specimens of Caudiverhera,
the denticles of bone on the skull begin
to fuse to adjacent denticles. In both
Eophractus and Gigantohatrachus, the
exostosed bones have a dermosotic pat-
tern in which the bone appears pitted.
The pitting is a result of fusion of adja-
cent denticles. I consider these charac-
ters to be due to age difference and
therefore not applicable to generic sep-
aration. Young specimens of the Recent
population have pitted dorsal surfaces
of the exostosed bones.

The skull of Caudiverhera is super-
ficially similar to those of the cerato-
phryines, but differs in the forward shift
of the eyes, pedicellate teeth, and the
presence of well defined sutures. The
vertebral column of Caudiverhera dif-
fers from those of the ceratophryines in
having non-expanded transverse proc-
esses of the anterior presacral vertebrae,
in lacking a dermostosed vertebral
shield, and in having dilated sacral dia-
pophyses. The vertebral column of Cau-
diverhera is more similar to that of
Tehnatohufo than to those of the other
genera of the tribe. As in the other
Telmatobiini, the pectoral girdle is mas-
sive with strongly arched and massive
clavicles and has a large omosternum
and large sternum (Fig. 34).

Reig (1960a) placed Cahjptocepha-

lella (z=CaudiverI)era) in a separate sub-
family Calyptocephalellinae. The sep-
aration was based on the peculiarities of
the skull of Caudiverhera. However,
Caudiverhera shares many osteological,
external, and non-osteological characters
with Batrachophrymis, Telmatohius, and
Tehuatolnifo, and differs from them in
having a completely roofed skull and the
skin of the head involved in dermostosis
with the skull bones. Reig (1960a) did
not compare Caudiverhera with Tehna-
tohufo, and therefore was unaware of
the numerous similarities between them.
Tehnatohufo is intermediate between
Caudiverhera and Tehnaiohius in many
of the distinguishing characters of Reig's
Calyptocephalellinae.

Telmatobufo Schmidt, 1952

Tehnatohufo Schmidt, 1952, Fieldiana, Zool.,
34:11 [Type-species by original designa-
tion, Tehnatohufo hullocki Schmidt, 1952].

Diagnostic definition. — .3) transverse
processes of posterior presacral vertebrae
shortened; 5) cervical and second verte-
brae apparently fused; 6) cranial bones
not involved in dermostosis; 7) omoster-
num very large; 8) sacral diapophyses
dilated; 9) maxillary arch toothed, teeth
pointed, slightly elongated, pedicellate;

10) alary processes of premaxillae di-
rected posterodorsally, broad at base;

11) palatal shelf of premaxilla broad,
apparently notched; 12) facial lobe of
maxilla deep, not exostosed; 13) palatal
shelf of maxilla relatively broad, ptery-
goid process lacking or poorly defined;
15) nasals not in median contact, rela-
tively small; 16) nasals not in contact
with maxillae or pterygoids; 17) nasals
not in contact with frontoparietals; 18)
frontoparietal fontanelle small; 19) fron-
toparietals not ornamented; 20); 21)
temporal arcade lacking; 22) epiotic
eminences poorly defined; 23) cristae
paroticae short, stocky; 24) zygomatic
ramus of squamosal of moderate length,
pointed; 25) otic ramus of squamosal
very short, not expanded to form otic
plate; 26); 27) columella apparently ab-

LYNCH: LEPTODACTYLOID FROGS

117

sent; 28) prevomers small, entire, sep-
arated medially, bearing teeth; 29) pala-
tines relatively narrow, long, separated
medially, lacking odontoid ridge; 30)
sphenethmoid entire, extending anterior-
ly to posterior edge of nasals; 31); 32)
parasphenoid alae not overlapped by
median rami of pterygoids; 33) anterior
rami of pterygoids in long contact with
maxillae, extending to palatines; 34) oc-
cipital condyles large, not stalked, close-
ly juxtaposed; 37-39); 40) pars scapu-
laris of m. depressor mandihulae mi-
nute, confined to a few muscle slips, pars
tympanicus massive; 41) pupil vertical;
42) males apparently lacking nuptial
asperities and vocal sac; 43) distinct
parotoid glands present; 44) tongue
large, rounded, posterior edge free; 45)
toes fully webbed, outer metatarsal tu-
bercle lacking, inner metatarsal tubercle
not spade-like, digital tips narrow, first
finger shorter than second; 46-48)''; 49)
two adults examined were 63 and 64
mm. SVL; 50) tympanum absent; 51)
fifth toe very broad, much broader than
in any other leptodactylid.

Composition. — Monotypic.

Distriljution. — Known only from the
Cordillera de Nahuelbuta, M allow,
Chile.

Retuarks. — The osteological charac-
ters (nos. 1-38) of Tehnatolnifo hid-
locki were studied by use of stereo-
radiographs of the holotype and para-
type. The observation of some charac-
teristics is very difficult and the state-
ments listed above are subject to rein-
terpretation. For example, the first two
vertebrae appear to be fused; this may
not be the case, but judging from prior
experience with skeletons and radio-
graphs, I feel the present interpretation
is probably correct. Similar arguments
and qualifications can be made for sev-

" A second species of Telmatobufo and larvae
are being described by Ramon Formas Cortes.
The genns has stream-adapted larvae with a
tooth row formula of 2/3, a \'entral mouth sug-
gestive of that in Ascaphus, Heleophryne, and
Staiirois, and papillae completely surrounding
the mouth.

eral other characteristics involving skull
bones. The transverse process of the pos-
terior presacral vertebrae of Telmato-
hiifo are shortened as in Caudiverhera,
CeratopJjrys, ProceratopJirys, and a few
other Neotropical leptodactylid genera.
Except for the dilated sacral diapophy-
ses, the vertebral column of Telmato-
bufo looks like that of Proceratoplirys
(Fig. 79). I consider Telmatobufo to be
more closely related to Caudiverbera
than to the other Telmatobiini. Caudi-
verbera and Tebiiatobufo share three
characters which are not exhibited by
the other Telmatobiini: shortened trans-
verse processes of the posterior presacral
vertebrae, vertical pupil, and absence of
an outer metatarsal tubercle. The two
genera differ in the casquing of the skull
of Caudiverbera; the skull of Tehnato-
biifo is identical, insofar as my observa-
tions will permit, with that of Tebnatobi-
us. In several respects Tebnatobufo is
intermediate between Neoprocoela and
Tebnatobius and fits the generalized pat-
tern of Telmatobiini.

Schmidt (1952) suggested that Tel-
matobufo was closely allied to, but
noticeably distinct from, Tebnatobius.
The only subsequent author to discuss
the validity and relationships of Telma-
tobufo was Gallardo (1962, 1965), who
suggested that Tebnatobufo bullocki
was identical with Aruncus valdivianus
PhiHppi. He further suggested (1962)
that Aruncus was not related to Tebna-
tobius but to CalyptocephaleUa (=Cau-
diverbera) but did not substantiate his
opinions with data.

Cei's (1958) reproductions of the
long-lost and unpublished plates for
Philippi's (1902) work provide much
more evidence concerning the identities
of the myriad of names proposed by
Philippi than do Philippi's brief descrip-
tions. The plates provide adequate
grounds for rejecting Gallardo's conten-
tion that Telmatobufo Schmidt is a syn-
onym of Aruncus Philippi. The plate for
Aruncus valdivianus (plate I in Cei,
1958) indicates that the frog had little,

118

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 79. Vertebral columns of five genera of the Telmatobiinae. (A) Proceratophrtis cristiceps
(KU 106273, X 2), (B) Odontophriimts cultripes ( KU 92975, X 2), (C) Ciidoramphus pinderi
(KU 92807, X 2), (D) Thowpu miliaris (KU 92856, X 2), and (E) Batrachophnjnm macrostonnis

(KU 96127, X 1).

if any, webbing between the toes, where-
as the specimens of Telmatohufo bul-
locki have fully webbed feet. The fin-
gers of Arunctis are proportionately
much longer than is the case for Tel-
matohufo. However, the most convinc-
ing data is the illustrated tympanum in
Aruncus valclivianus and the absence of
the tympanum in TelmatoJ)iifo huUocki.
Cei's ( 1958 ) suggestion that the figures
of Aruncus valclivianus represent a poor
rendition of Bufo spinulosus is reason-
able. Cei ( 1958 ) objected to the inclu-
sion of Aruncus valclivianus in the syn-
onymy of Bufo spinulosus because the
figures are not sufficiently accurate to
permit an assignment to subspecies, and
the synonymy would affect the applica-
tion of subspecific names. I suggest that
Aruncus vaklivianus Philippi, 1902, be
considered a nomen cluhium for the
present.

Telmatobius Wiegmann, 1835

( Figs. 80-81 )

Telmatobius Wiegmann, 1835, Nova Acta Acad.
Leop. -carol, 17:262 [Type-species by mono-

typy, Tehnatobitis peruvianus Wiegmann,
1835].

Pseudobatrachus Peters, 1873, Mtber. k. Preuss.
Akad. Wiss., Berlin, p. 414 [Type-species
by monotypy, Pseudobatrachus jelskii
Peters, 1873].

Cophaeus Cope, 1889, Bull. U.S. Natl. Mus.,
34:312, 381 [Apparently a replacement or
substitute name for part or all of Boulen-
ger's ( 1882 ) Telmatobius, which Cope con-
sidered not equal to Tehuatobius of Wieg-
mann (1835)].

Diagnostic definition. — 3) transverse
processes of posterior presacral vertebrae
not shortened; 5) cervical and second
vertebrae not fused; 6) cranial bones not
involved in dermostosis; 7) omosternum
large; 8) sacral diapophyses dilated; 9)
maxillary arch toothed, teeth pointed,
pedicellate; maxillary arch toothless in
a few populations; 10) alary processes of
premaxillae directed posterodorsally,
broad at base; 11) palatal shelf of pre-
maxilla narrow with relatively long pala-
tal process; 12) facial lobe of maxilla
relatively deep, not exostosed; 13) pala-
tal shelf of maxilla relatively narrow,
pterygoid process lacking; 14) maxillary

LYNCH: LEPTODACTYLOID FROGS

119

Figure 80. Dorsal and ventral views of skull of

Telmatobius hautholi ( KU 72879, X 5 ) . Right

septomaxilla lost in preparation.

arch complete in most species, quad-
ratojugal lacking in edentulous pata-
gonicus; 15) nasals separated medially
and small in dentate species, larger and
narrowly separated in patagonicus; 16)
nasals not in contact with maxillae or
pterygoids; 17) nasals widely separated
from frontoparietals; 18) frontoparietal
fontanelle small, except in patagonicus
in which it is large; 19) frontoparietals
not or but slightly ornamented; 21 ) tem-
poral arcade absent; 22) epiotic emi-
nences well defined; 23) cristae paroticae
short and stocky; carotid artery not en-
closed in a canal, sometimes a shallow
groove is present on the frontoparietal
and otoccipital; 24) zygomatic ramus of
squamosal relatively long, blunt, widely
separated from maxilla; 25) otic ramus
of squamosal very short, no otic plate;
26) squamosal-maxillary angle about
40°; 27) columella present, absent in
patagonicus; 28) provomers present,
usually toothed, sometimes edentate, en-
tire, in contact medially or narrowly
separated; 29) palatines long and nar-
row, narrowly separated medially; 30)

sphenethmoid entire, extending anterior-
ly to posterior edge of nasals; 31) an-
terior ramus of parasphenoid relatively
narrow, lacking median keel, extending
anteriorly between palatines; 32) para-
sphenoid alae oriented at right angles to
anterior ramus, not overlapped laterally
by median rami of pterygoids; 33) ptery-
goids relatively small, anterior rami
elongate, nearly reaching palatines; 34)
occipital condyles relatively large, not
stalked, narrowly separated medially;
37) alary processes of hyoid plate on
narrow stalks; 40) m. depressor man-
dibulae in two slips; 41) pupil horizon-
tal; 42) males lacking vocal sac; nuptial
asperities or clusters of spines on thumb
and sometimes on chest; 43) body lack-
ing glands; 44) tongue large, rounded,
posterior edge free; 45) toes usually
completely webbed, outer metatarsal
tubercle present, inner metatarsal tuber-
cle not enlarged, digital tips narrow;
46) larvae with dextral vent, 2/3 tooth
rows, labial papillae interrupted an-
teriorly; 49) adults to about 60 mm.
SVL; 50) tympanum small or concealed.
Composition. — Vellard (1951) recog-
nized 19 species of the genus. Schmidt

Figure 81. Lateral view of skull of Telmatobius

marmoratus (UMMZ 68179, X 4) and dorsal

view of skull of Tehnatohius patagonicus (KU

80781, X 7.3).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

( 1954a ) recognized 21 and Vellard
( 1960 ) modified his earlier account and
recognized 23 species. Batrachophnjnus
patagonicus is a species of Telmatohius
and one additional species has been de-
scribed since Vellard's last paper. Fifty-
one populations of Telmatohius are
presently afforded nonienclatorial recog-
nition. The following species of Telma-
tohius are recognized; the number of sub-
species is included in parentheses: alhi-
ventris (4), arequipensis (2), atacamen-
si-s, brevipes, hrevirostrls (3), cinereus,
craicfordi (2), culeus (6), haJli (2),
hauthaU (2), ignavtis, intermedius, jel-
skii (4), laevis, marmoratiis (7), mon-
tanus, niger, oxycephalus, patagonicus,
penwianus, praehasalticus, reverherii,
rimac (2), simonsi, somuncurensis, vel-
lardi, and verrucosus."'

Distribution. — South American Andes
from southern Ecuador (1° S) to central
Chile and Argentina (32° S). The great-
est diversity occurs in southern Peru.

Remarks. — As presently constituted,
Telmatohius is one of the larger lepto-
dactylid genera. Vellard (1951, 1953,
1955, 1960) placed the 23 species known
to him in three species groups: penwi-
anus group (stream frogs), marmoratus
group (primarily or totally aquatic
frogs), and jelskii group ( semiterrestrial
frogs). However, several authors have
pointed out that the closest relatives of
some aquatic species are semiterrestrial
species. Loss of the maxillary or prevo-
merine teeth has been a major taxonomic
character in the study of the relation-
ships of the frogs of this genus. Maxil-
lary teeth are lost by some members of
the peruvianus and marmoratus groups.
Prevomerine teeth are lost by some pop-
ulations of all species groups.

I have seen few skeletons of this

' The species list given here is incomplete. Cei
and Vellard have increased the niunher of taxa
known from Argentina. Gallardo (1970) and
Cei (in litt.) suggest that T. montanus is more
closely related to certain species here assigned
to Eupsophus. Studies on tire osteology of many
species of Telmatohius are in progress and when
completed should provide some clarification.

genus, and therefore my characterization
of it will probably undergo some altera-
tion with the acquisition of additional
material in the future. The paedomor-
phic T. patagonicus is strikingly different
in cranial osteology (Fig. SI) from the
other species I examined.

tNeoprocoela Schaelfer, 1949

Neoprococla Schaeffer, 1949, Bull. Amer. Mus.
Nat. Hist., 93:57 [Type-species by original
designation, Neoprocoela edentatus Schaef-
fer, 1949 (=N. edentata). Lower Oligo-
cene].

Schaeffer (1949) described and named
a relatively well preserved bufonoid frog
from the Lower Oligocene of Patagonia
as a new genus and new species of Lep-
todactylidae. The fossil is edentate (as
is reflected in Schaeffer's choice of a
trivial name), has a moderately small
frontoparietal fontanelle, and dilated
sacral diapophyses. Schaeffer did not
consider the fossil to be a bufonid be-
cause to do so ". . . would require the
presence of the Bufonidae (sensu stricto)
in South America by no later than
the early Oligocene, an occurrence
which is not supported by the known
paleontological facts." Exactly what
facts these were was not explained by
Schaeffer. Instead, he assigned the fos-
sil to the Leptodactylidae, and charac-
terized it as having a number of primi-
tive, criniid-like traits. Schaeffer sug-
gested that Neoprocoela was related to
Batrachophnjnus and Telmatohius.

The discovery of a Miocene toad
(Bufo marinus) from Colombia (Estes
and Wassersug, 1963) clearly estabhshes
the Bufonidae in South America from at
least the Eocene because South America
was isolated from Middle America be-
tween the Eocene and Pliocene (Lloyd,
1963). Therefore, Schaeffer's objections
to placing Neoprocoela in the Bufonidae
can be seriously questioned. Tihen
( 1962b ) placed Neoprocoela in the syn-
onymy of Bufo and considered the type-
species as a species of the B. calamita
group. Tihen associated the fossil with

LYNCH: LEPTODACTYLOID FROGS

121

Bufo for the following reasons: 1) in
edentulous leptodactylids, the prevomers
are greatly reduced in size, 2) the alary
processes of leptodactylids are directed
dorsally or laterally, not toward the mid-
line as in Neoprocoela edeniata and bu-
fonids, 3) the sphenethmoid is entire in
Neoprocoela, 4) the shape of the squa-
mosal suggests a large squamosal-maxil-
lary angle, 5) the broad nasals are in
median contact, 6) there is a long max-
illary-pterygoid contact, 7) the maxillae
are very broad, 8) the atlantal cotyles
are closely approximated ventrally, 9)
the first transverse processes are directed
anterolaterally and are expanded ( ^di-
lated), and 10) the sacral diapophyses
are expanded (^dilated). Tihen cited
two other characters of Neoprocoela
which are uncommon in bufonids but
characteristic of the Btifo calamita group
— large frontoparietal fontanelle and
very short otic ramus of the squamosal.
Tihen argued that while each of the ten
characteristics listed above can be dem-
onstrated to occur in one or more lepto-
dactylid groups, the simultaneous occur-
rence of all ten is not known for any
leptodactylid, but is the case in Bufo.
Tihen's "leptodactylid condition" for his
comparison of Neoprocoela, leptodactyl-
ids, and bufonids, must have been based
on the skeletons of some of the Australo-
Papuan leptodactylids. Characters 3)
and 5) of Neoporcoela are very much
unlike the conditions seen in edentulous
Australo-Papuan leptodactylids, but are
like the conditions seen in some edentu-
lous Neotropical leptodactylids. With
the exception of characters 1), 2), and
10), Neoprocoela agrees completely
with Batrachophrijmis. The fossil agrees
with Bairachophnjnus in the two char-
acteristics cited by Tihen that are "un-
usual" for Bufo. The dilation of the
sacral diapophyses of Neoprocoela is not
as great as in Bufo. The sacral dia-
pophyses are round in Batrachophrymis
(Fig. 79). The dilation of the sacral
diapophyses of Neoprocoela is no greater
than that seen in Telmatobufo, a close

relative of Batrachoplirynus. The length
of the transverse processes of the pos-
terior presacral vertebrae is not evident
in the single fossil of Neoprocoela; this
character is very different in Batracho-
phrijnus and the Bufo calamita group.

Tihen recorded the alary processes
(ascending processes of Tihen) of the
premaxillae as directed toward the mid-
line in Neoprocoela. This is not a lepto-
dactylid trait but clearly a bufonid trait.
Perusal of Schaeffer's figures clearly in-
dicates that the skull of Neoprocoela
was crushed and distorted. The skull
apparently was crushed from the left to
the right side. The premaxillae are dis-
torted with an anterior rotation at their
median suture; this results in a deflection
toward the midline of the alary proc-
esses. As in Batrachophrymis, the alary
processes are long and thin.

The large prevomers of Neoprocoela
readily distinguish it from Batracho-
phrymis, in which the prevomers are
minute. In Caudiverhera, Telmatohius,
and Telmatohnfo, the prevomers are
large and toothed. It is only logical to
assume that the ancestral stock of Batra-
chophrymis had large and toothed pre-
vomers. In the course of loss of the
prevomers and prevomerine teeth, two
patterns are observed. In one pattern,
tooth loss occurs late; the prevomerine
bones are reduced in size until only the
dentigerous ramus and a small semicircle
of bone surrounding the inner edge of
the choana remains. The teeth and den-
tigerous ramus are then lost, in that
order. This is the pattern seen in the
Myobatrachinae. In the other pattern,
the prevomer is not greatly reduced in
size before the teeth are lost. The bone
continues to be reduced in size subse-
quent to tooth loss. This pattern is seen
in several genera of the Telmatobiinae,
Leptodactylinae, and Elosiinae. Neo-
procoela fits the intermediate condition
between Telmatobufo and Batracho-
phrymis.

Neoprocoela is intermediate between
Telmatobufo and Batrachophrymis in

122

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

the shape of the sacral diapophyses and
in shape of the cristae paroticae, as well
as the size of the prevomers. Tihen also
cited the "toad-like" body shape of Neo-
procoela as an additional bufonid char-
acter. However, Telmotobtifo is very
toad-like even in the possession of well
defined parotoid glands.

Based on the skeletal data evident in
the figures of Neoprocoela published by
Schaeffer (1949) and Tihen (1962b),
the following characteristics of my diag-
nostic definition can be stated: 2) verte-
bral shield probably absent; 4) cervical
cotylar arrangement type II; 5) cervical
and second vertebrae not fused; 6) skull
bones not exostosed, therefore derm of
head free; 8) sacral diapophyses dilated;
9) maxillary arch edentate; 10) alary
processes of premaxilla narrow, directed
posterodorsally; 11) palatal shelf of pre-
maxilla very broad, slightly indented;
12) facial lobe of maxilla deep; 13) pala-
tal shelf of maxilla broad with a well
developed pterygoid process; 14) maxil-
lary arch complete; 15) nasals large, in
broad median contact; 16) nasals in con-
tact with maxillae; 17) nasals not in con-
tact with frontoparietals; 18) frontopari-
etal fontanelle of moderate size; 19)
frontoparietals not ornamented; 21) tem-
poral arcade lacking; 23) cristae paro-
ticae moderately short, stocky; 24) zygo-
matic ramus of squamosal of moderate
length, widely separated from maxilla;
25) otic ramus of squamosal very small,
no otic plate; 28) prevomers large, eden-
tate; 29) palatines elongate; 30) sphen-
ethmoid large, entire; 32) parasphenoid
alae oriented at right angles to anterior
ramus of parasphenoid; 33) anterior
rami of pterygoids elongate, in long con-
tact with maxillae, reaching palatines;
34) occipital condyles large, not stalked,
narrowly separated medially; 36) termi-
nal phalanx of one digit knobbed; 49)
snout-coccyx length at least 66 mm. The
single terminal phalanx could be of the
thumb even if the frog had T-shaped
terminal phalanges.

Batrachophrynus Peters, 1873

(Fig. 82)

Botrachophrtjmts Peters, 1873, Mtber. k. Preuss.
Akad. Wiss., Berlin [Type-species by pres-
ent designation, Batrachophrynus macro-
stomus Peters, 1873].

Diagnostic definition. — 3) transverse
processes of posterior presacral vertebrae
oriented at right angles to sagittal line,
as long as sacral diapophyses; 5) cervical
and second vertebrae not fused; 6) cra-
nial bones not dermostosed; 7) omoster-
num present, large; 8) sacral diapoph-
yses round; 9) maxillary arch eden-
tate; 10) alary processes of premaxillae
narrow, directed posterodorsally; 11)
palatal shelf of premaxilla very broad,
slightly indented; 12) facial lobe of max-
illa deep; 13) palatal shelf of premaxilla
broad, pterygoid process relatively
small; 15) nasals large, in broad median
contact; 16) nasals in contact with max-
illae and pterygoids; 17) nasals not in
contact with frontoparietals; 18) fronto-
parietal fontanelle small; 19) frontopari-
etals not ornamented, except for a sharp
shelf immediately posterior to orbit; 21 )
temporal arcade lacking; 22) epiotic
eminences large posteriorly, obsolete an-
teriorly; 23) cristae paroticae very long
and narrow; carotid artery not enclosed
in bony canal, frontoparietals sometimes
having groove between ridge and epiotic

Figure 82. Lateral ( X 1 ) and dorsal ( X 0.7)
views of skull of Batrachophrynus macrostomiis
(KU 96127).

LYNCH: LEPTODACTYLOID FROGS

123

eminence; 24) zygomatic ramus of squa-
mosal short; 25 ) otic ramus of squamosal
very short, no otic plate; 26) squamosal-
maxillary angle 35-40°; 27) columella
present, thin; 28) prevomers small, eden-
tate, only dentigerous ramus present;
29) palatines broad, widely separated
medially, lacking odontoids; 30) sphen-
ethmoid large, entire, extending an-
teriorly to front of nasals; 31) anterior
ramus of parasphenoid broad, pointed,
keeled medially, extending anteriorly be-
tween palatines; 32) parasphenoid alae
oriented at right angles to anterior ra-
mus of parasphenoid, broadly overlapped
laterally by median rami of pterygoids;
33) pterygoids very large, anterior rami
in long contact with maxillae, reaching
palatines; 34) occipital condyles large,
not stalked, narrowly separated medial-
ly; 37) alary processes of hyoid plate on
narrow stalks; 40) m. depressor mancli-
hulae in two slips; 41) pupil horizontal;
42) males apparently lacking nuptial
asperities and vocal sac; 43) body lack-
ing glands; 44) tongue large, completely
adherent; 45) toes fully webbed, outer
metatarsal tubercle present, inner meta-
tarsal tubercle not enlarged, digital tips
narrow, first finger longer than second;
46) larvae with dextral vent, 2/3 tooth
rows, labial papillae interrupted an-
teriorly; 49) adult hrachijdactijlus are
47-58 mm. SVL and macrostomus grows
to 160 mm. SVL; 50) tympanum absent.

Composition. — Two species are rec-
ognized, hraclnjdactylus and macrosto-
mus.

Distribution. — Lago de Junin region
in central Peru.

Remarks. — Batrachophnjnus and Tel-
matohius are usually considered to be
very closely related and were separated
solely on the basis of the presence (Tel-
matobius) or absence (Batrachophnjnus)
of maxillary and prevomerine teeth.
Four populations of Telmatohius lack
maxillary teeth (brevipalmatus, edenta-
ttis, intermedins, and patagonicus). The
palatal shelf is broad in Batrachophnjnus
and narrow in Tehnatobius, the prevo-

mers are small and edentate in Batra-
chophrynus but usually are moderate-
sized to large and toothed in Tehnato-
J)ius regardless of whether there are
teeth on the maxillary arch, and the pos-
terior edge of the tongue is not free in
Batrachophrynus. The distinctions be-
tween Batrachophrynus and Tehnato-
bius are difficult to assess at present be-
cause so few species have been studied.
The aquatic Tehnatobius bear greater
resemblance in external characters to
Batrachophrynus than to the semiter-
restrial species of Tehnatobius.

Alsodini Mivart, 1869^

Alsodina Mivart, 1869:290.
Cacotina Mivart, 1869:290.
Batrachylinae Gallardo, 1965:83.

Four genera are included in this
Neotropical tribe — Batrachyla, Eupso-
plnis, Hylorina, and TJioropa. The dis-
tribution of the group is only slightly
more extensive than that of the Telma-
tobiini, in that one genus (Thoropa) is
found in the mountains of southeastern
Brasil. Eupsophus and Hylorina are
closely related as are Batrachyla and
Thoropa. However, the two generic
pairs share few significant characters.^
The cervical cotylar arrangement is type
II in Eupsophus and Hylorina, whereas
it is type I in Batrachyla and Thoropa.

^Barrio's (1970) Insuetophryniis probably be-
longs here but until specimens have been exam-
ined for all characteristics employed here
(especially osteological ) , any assignment must
be considered tenuous. Barrio considered the
genus most closely allied to Abodes (=zEup-
sophus nodosits group as used here) principally
on the basis of the secondary sex characteristics.
Further comment on Insuetophryniis is post-
poned pending completion of studies of the
skeletons of Eupsophus, Insiietophrynus, and
Tehnatobius.

" In a separate paper ( Lynch, MS ) I used
Batrachylini as the tribe name for Batrachyla
and Thoropa and assigned Eupsophus and Hy-
lorina to the Telmatobiini. This action reflects
my incomplete knowledge of Tehnatobius skele-
tons at present. 1 regard either arrangement as
defensible at present and hope to clarify the
relationships of these genera once my study of
Tehnatobius osteology is completed.

124

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

All four genera have frontoparietal fon-
tanelles, vertebral columns in which the
transverse processes of all vertebrae (ex-
cept 1 and 2) are as long as the sacral
diapoph>ses, and slightly dilated sacral
diapophyses. All four genera have free
swimming tadpoles. The tadpoles of
Batraclu/la differ from those of the other
genera in ha\ ing an uninterrupted series
of labial papillae. The tadpoles of
Tlioropa are greatly flattened and atten-
uate (Fig. 2) in an adaptation to tor-
rential stream life. Amplexus is inguinal
in Batracliyla, but is axillary in Eiipso-
phus and Hijlorina; the amplectic posi-
tion is not known for Thoropa. Eiipso-
phus and Hylorina lay numerous small
eggs in water, whereas Batraclujla lays
fewer larger eggs in terrestrial sites. The
eggs of Batraclujla hatch and the larvae
live in the jelly mass until the mass is
inundated. Tlioropa has large eggs
which are deposited on wet stones in
situations where water trickles over
stone ledges (Myers, 1946). Werner C.
A. Bokermann (pers. comm.) suggested
that the eggs are laid on the banks of
the torrential streams inhabited by
Thoropa.

The following diagnostic character-
istics are uniform among the four gen-
era of the group: 3) transverse processes
of posterior presacral vertebrae long; 5)
cervical and second vertebrae not fused;
6) cranial bones not involved in dermo-
stosis; 7) omosternum present, mod-
erately large; 8) sacral diapophyses
somewhat dilated — see Fig. 79; 9) max-
illary arch toothed, teeth blunt, pedicel-
late; 12) facial lobe of maxilla deep, not
exostosed; 17) nasals not in contact with
frontoparietals; 18) frontoparietal fon-
tanelle moderate-sized; 19) frontopari-
etals not ornamented; 20) frontoparietal
not fused with prootic; 21) temporal
arcade lacking; 37) alary processes of
hyoid plate on narrow stalks; 42) males
with nuptial asperities on thumb and
sometimes second finger; some species of
Eiipsophus have cluster of asperities on
chest; 45) outer .iietatarsal tubercle pres-

ent, inner metatarsal tubercle not en-
larged or spade-like; 46) larvae with
median vent.

The tribal name, Alsodini, is based
on Abodes Bell, 1843, which was recent-
ly shown to be a synonym of Eupsophus
(Lynch, 1968b). The heterogeneity of
the tribe and the mosaic of primitive
characteristics exhibited by the four in-
cluded genera suggest that the Alsodini
might be best regarded as a suprageneric
grade between the primitive Telmato-
biini and the advanced Eleutherodacty-
lini.

Eupsophus Fitzinger, 1843

(Fig. 83)

Eupsophus Fitzinger, 1843, Syst. Rept., p. 31
[Type-species by original designation, Cys-
tignathus roseus Dumeiil and Bibron, 1841].

HammatodactijJus Fitzinger, 1843, Syst. Rept.,
p. 31 [Type-species by original designation,
Cijstignuthus nodosus Dumeril and Bibron,
1841].

Borborocoetes Bell, 1843, Zool. Voy. Beagle,
Reptiles, 5:34 [Type-species by present
designation, Borborocoetes grayi Bell, 1843;
preoccupied by Borborocoetes Sclioenherr,
1842 (Insecta: Coleoptera)].

Abodes Bell, 1843, Ibid., .5:34 [Type-species by
monotypy. Abodes monticola Bell, 1843]. i^

'" Gallardo ( 1970) proposed recognizing Alsodes
as distinct from Eupsophus and included seven
Patagonian species in Abodes. The seven are
Abodes gargoJa Gallardo, 1970, Eupsophus il-
lotus (Barbour), 1922, £. monticola (Bell),
1843, E. nodosus (Dumeril and Bibron), 1841,
Tehnatobius montanus Lataste, 1902, T. prae-
basahicus Cei and Roig, 1968, and T. ivveiberii
Cei, 1969. Gallardo (1970) defined the genus
Alsodes as follows: atlas convex, sternum ex-
panded and notched posteriorly, tympanum not
visible externally, vocal sac absent in males,
forelimbs of reproductively active males greatly
enlarged, and nuptial spines present on fingers
and chest of reproductixely active males. Barrio
(1970) tentatively accepted Gallardo's arrange-
ment but did not include Tehnatobius pracba-
salticus and T. reverberii as species of Abodes.
Further comment on this arrangement is post-
poned pending completion of my studies of the
skeletons of several Tehnatobius (sensu Into).
The separation of Alsodes from all other lepto-
dactylids rests on the presence of pectoral plates
of nuptial spines. Some species of Tehnatobius
ijelskii group) haxe pectoral plates, as does
Insuetophrynus acat})icus (Barrio, 1970).

If Fitzinger's ( 1843 ) paper antedates Bell's

LYNCH: LEPTODACTYLOID FROCS

125

Figure 83. Lateral, dorsal, and \'entral \iews of .skull of Etipsophus roseus (AMNH 22104, X 4.3).

Eusophus Cope, 1865, Rev. Nat. Hist., 5:113
[Emendation (?) of Eupsophus Fitzinger,
1843].

Borhorocactes Cope, 1866, J. Acad. Nat. Sci.
Philadelphia, (2)6:94 [Emendation of
Borborocoetes Bell, 1843, hence taking
same type-species].

Cacotus Giinther, 1868, Proc. Zool. Soc. Lon-
don, 1868:482 [Type-species by monotypy,
Cacotus maciilatus Giinther, 1868].

Thnjnopus Peters, 1873, Mtber. k. Preuss. Akad.
Wiss., Berlin, 1873:416 [Type-species by
monotypy, Phrtjnopus peruanus Peters,
1873].

Borhorocoetea Strand, 1928, Ark. Naturgesch.,
92A:55 [Replacement name for Borboro-
coetes Bell, 1843 (preoccupied), hence
taking same type-species].

Diagnostic definition. — 4) cervical
cotylar arrangement type II; 10) alary
processes of premaxillae directed pos-
terodorsally, moderately wide at base;
11) palatal shelf of premaxilla relatively
deep, palatal process elongate; 13) pala-

(1843), as herpetologists have generally pre-
sumed, Hammatodactylus Fitzinger, 1843 (type-
species Cystignathiis nodosus Dumeril and
Bibron, 1841) is the correct name for the
generic group recognized by Gallardo (1970)
and Barrio (1970) and called Alsodes.

tal shelf of maxilla of moderate width,
pterygoid process moderately large; 14)
maxillary arch complete, quadratojugal
present; 15) nasals small, widely sep-
arated medially; 16) nasals in broad con-
tact with maxillae, not in contact with
pterygoids; 22) epiotic eminences promi-
nent; 23) cristae paroticae relatively
broad, elongate; carotid artery passes
dorsal to skull bones; 24) zygomatic ra-
mus of squamosal of moderate length,
widely separated from maxilla; 25) otic
ramus of squamosal as long as zygomatic
ramus, expanded medially into small otic
plate; 26) squamosal-maxillary angle 50-
55°; 27) columella present or absent;
28) prevomers moderate-sized, separated
medially, entire, toothed except in jun-
inensis; 29) palatines broad, widely sep-
arated medially, bearing odontoid ridges;
30) sphenethmoid entire, extending an-
teriorly to anterior edge of nasals; 31)
anterior ramus of parasphenoid broad,
short, keeled medially; 32) parasphenoid
alae oriented at right angles to anterior
ramus of parasphenoid, broadly over-
lapped laterally by median rami of ptery-

126

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

golds; 33) pterygoids moderate-sized,
anterior rami in long contact with maxil-
lae, reaching palatines; 34) occipital
condyles large, not stalked, narrowly
separated medially; 36) terminal pha-
langes knobbed; 40) m. depressor man-
dihulae in two slips except in juninensis
which has only the pars tympanicus; 41)
pupil horizontal; 42) males with median
subgular vocal sac or none; 43) body
lacking glands or having extensive, dif-
fuse glandular areas over dorsum; 44)
tongue large, round, posterior edge free;
45) toes lacking webbing or fringing to
two-thirds webbed; 46) larvae with 2/3
tooth rows, labial papillae interrupted
anteriorly; 47) amplexus axillary; 48)
eggs small and numerous, laid in gela-
tinous masses in ponds; 49) males 32-
80, females 32-60 mm. SVL; 50) tym-
panum visible externally, concealed, or
absent.

Composition. — Revisionary studies of
the Argentine and Chilean species are
available (Cei, 1962a, Grandison, 1961,
and Gallardo, 1962 ) . As a result of these
studies and my own (Lynch, 1971),
seven species of the genus are presently
recognized: illotus, juninensis, monti-
cola, nodosus, peruanus, roseus, and ver-
tehralis. The status of the genus in Peru
is poorly known.

Distribution.— The Andes of central
Peru to Argentina and Chile; between
about 10° and 50° S latitude in western
South America.

Remarks. — Boulenger (1882) com-
bined a large number of genera and
species into Borhorocoetes Bell {^Eup-
sophus) in his synopsis of living am-
phibians. While most of his generic
groupings were a vast improvement over
the previous classifications, Borhoro-
coetes was a notable exception. He in-
cluded a variety of unrelated groups in
Borhorocoetes. The Borhorocoetes of
Boulenger and Noble is best described
as a grade (in the sense of Huxley,
1958). All of the species included are
members of the Telmatobiinae, and ac-
cording to the present classification be-

long to the genera Batraclujia, Eleu-
tJierodactyhis, Eupsophus, Ischnocnema,
Niceforonia, TJioropa, and Zachaenus.

The Chilean and Argentine species of
Eupsophus were studied in detail by Cei
(1960, 1962a, and 1962b) and Grandison
( 1961 ) , but they confused one species
of BatracJu/la with Eupsophus (iaenia-
tus). Cei (i962a) and Grandison (1961)
divided Eupsophus into three species
groups — nodosus group, peruanus group,
and roseus group. These authors also
recognized a monotypic taeniatus group,
which is here included in Batraclujia.
Two species of the genus (juninensis
and monticola have lost the columella.
These species also lack tympanic annuli.
The tympanic annulus is very small in
two other species of the genus, illotus
and nodosus, and is concealed beneath
the skin. In peruanus, roseus, and ver-
tebralis, the columella is normal-sized
and the tympanic annulus is visible ex-
ternally.

Schaeffer (1949) described, but did
not name, a fossil frog from the Lower
Oligocene of Chubut, Argentina, and re-
ferred it to Eupsophus. ^^ The nasals of
the fossil are apparently in median con-
tact, unlike the condition seen in the
living species of the genus. The fossil
could equally well be a species of Tel-
matohius, were it not for the fact that
the frontoparietal fontanelle is moderate-
ly large, not small. The middle ear was
not preserved.

Hylorina Bell, 1843
(Fig. 84)

Hylorina Bell, 1843, Zool. Voy. Beagle, Rep-
tiles, 5:44 [Type-species by monotypy, Hy-
lorina sylvatica Bell, 1843].

Hylorliina Agassiz, 1846, Noniencl. Zool., in-
dex: 190 [Emendation of Hylorina Bell,
1843, hence taking same type-species].

" Bogart ( 1970) implied that the Oligocene fos-
sil of Eupsophus is not separable from living
E. roseus. However, Schaeffer (1949) cited dis-
tinguishing features of the fossil, chiefly in the
arrangement of the nasal bones. The Oligocene
fossil needs to be restudied to ascertain its dis-
tinction from Tclmatohiits.

LYNCH: LEPTODACTYLOID FROGS

127

Figure 84. Dorsal and ventral views of skull
Diagnostic definition. — 4) cervical
cotylar arrangement type II; 10) alary
processes of premaxillae directed dorsal-
ly, wide at base; 11) palatal shelf of pre-
maxilla narrow, palatal process relatively
small; 13) palatal shelf of maxilla nar-
row, pterygoid process minute; 14) max-
illary arch incomplete, quadratojugal ab-
sent, replaced by ligamentous sheath;
15) nasals moderate sized, widely sep-
arated medially; 16) nasals in tenuous
contact with maxillae, not in contact
with pterygoids; 22) epiotic eminences
moderately well defined; 23) cristae
paroticae short, stocky; carotid artery
passes dorsal to skull bones; 24) zygo-
matic ramus of squamosal of moderate
length, widely separated from maxilla;
25) otic ramus of squamosal moderately
long, shorter than zygomatic ramus, ex-
panded medially into small otic plate;
26); 27) columella present; 28) prevo-
mers moderately large, entire, narrowly
separated medially, toothed; 29) pala-
tines broad, widely separated medially,
no odontoid ridges; 30) sphenethmoid
entire, extending anteriorly to middle of
nasals; 31) anterior ramus of parasphe-
noid broad, short, not keeled medially;
32) parasphenoid alae oriented at right
angles to anterior ramus, broadly over-
lapped laterally by median rami of
pterygoids; 33) pterygoids small, an-
terior rami short, extending to middle of
orbit; 34) occipital condyles moderately

of Hijlorina stjlvatica (BMNH 91.29.17, X 3).
large, not stalked, narrowly separated
medially; 36) terminal phalanges
knobbed, elongate; 40) m. depressor
mandibtdue in two slips; 41) pupil ver-
tical; 42) males with median subgular
vocal sac; 43 ) body with glandular dorso-
lateral folds; 44) tongue large, rounded,
posterior edge free; 45 ) toes lacking web-
bing or lateral fringes, digital tips narrow,
first finger longer than second; 46) larvae
with 2/2 tooth rows, labial papillae in-
terrupted anteriorly; 47) amplexus axil-
lary; 48) eggs small, numerous, laid in
gelatinous masses at bases of plants in
water; 49) males 50-60, females 60-68
mm. SVL; 50) tympanum visible exter-
nally; 51) digits extremely long, phalan-
geal formulae not increased.

Composition. — Monotypic.

Distribution. — Central Chile.

Remarks. — Because Hylorina is un-
common, the genus was studied with the
aid of stereo-radiographs. Hylorina has
been considered generically distinct since
Bell's description of the type-species. In
part the distinction stemmed from er-
roneous data provided by Boulenger
(1882), who reported the sternum as
bony. The genus Hylorina is very dis-
tinctive even though the sternum is a
cartilaginous plate which tends to calcify
in old adults. The combination of verti-
cal pupil, free toes, greatly elongated
digits, externally visible tympanum, and
teeth on maxillary arch and prevomerine

128

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

dentigerous processes immediately dis-
tinguishes Hylorina from all other frog
genera. In spite of the distinctiveness of
Hylorina, its skeletal morphology allies
it with Eupsoplms. The data on breed-
ing biology reported by Barrio (1967b)
provide additional distinction for Hy-
lorina, but also point out the similarity
between Eupsophus and Hylorina.

Batrachyla Bell, 1843

(Fig. 85)

Batrachyla Bell, 1843, Zool. Voy. Beagle, Rep-
tiles, 5:43 [Type-species by monotypy, Ba-
traclujla leptoptis Bell, 1843].

Diagnostic definition. — 4) cervical
cotylar arrangement type I; 10) alary
processes of premaxillae directed dorsal-
ly and somewhat laterally, moderately
wide at base; 11) palatal shelf of pre-
m axilla very narrow, palatal process rela-
tively large; 13) palatal shelf of maxilla
narrow, pterygoid process lacking; 14)
maxillary arch incomplete, cjuadrato-
jugal absent; 15 ) nasals widely separated
medially, relatively small; 16) nasals

separated from maxillae and pterygoids;
22) epiotic eminences obsolete; 23)
cristae paroticae stocky, relatively long;
carotid artery passes dorsal to skull
bones; 24) zygomatic ramus of squa-
mosal elongate, widely separated from
maxilla; 25) otic ramus of squamosal
moderately long, expanding medially in-
to small otic plate; 26) squamosal-maxil-
lary angle about 60°; 27) columella
present; 28) prevomers relatively small,
entire, separated medially, toothed; 29)
palatines curved, narrow, widely sep-
arated medially; 30) sphenethmoid en-
tire, extending anteriorly to a point an-
terior to nasals; 31) anterior ramus of
parasphenoid broad, short, lacking medi-
an keel; 32) parasphenoid alae oriented
at right angles to anterior ramus of para-
sphenoid, not overlapped by median
rami of pterygoids; 33) pteiygoids small,
thin, anterior rami short, not extending
beyond middle of orbits; 34) occipital
condyles small, not stalked, widely sep-
arated medially; 36) terminal phalanges
T-shaped; 40) 7n. depressor mandilmlae

Figure 85. Lateral, dorsal, and ventral views of skull of Batrachyla leptopus (UMMZ S-2246, X 6).

LYNCH: LEPTODACTYLOID FROGS

129

in two slips; 41) pupil horizontal; 42)
males with median subgular vocal sac;
43) body lacking glands; 44) tongue
moderately large, posterior one-third
free; 45) toes lacking web or lateral
fringes, digital tips bulbous, somewhat
dilated, first finger shorter than second;

46) larvae with 2/3 tooth rows, labial
papillae not interrupted about mouth;

47) amplexus inguinal; 48) eggs rela-
tively few, large, laid in terrestrial situa-
tions, tadpoles become aquatic after nest
is inundated; 49) adults 27-40 mm. SVL;
50) tympanum visible externally.

Composition. — Barrio (1967a) recog-
nized two species (aniartandica and hp-
toptis) of Batraclnjla. Lynch (1971)
demonstrated that Eupsophiis taeniatus
belongs to the genus Batrachyla.

Distribution. — Chile and adjacent
Argentina between 32° and 50° S lati-
tude.

Remarks. — Boulenger (1882) and
Myers (1962) considered BatrachijJa
synonymous with Eleutherodactijlus
(Hylodes Fitzinger, 1843, in the case of
Boulenger). Both authors were under
the mistaken impression that the two
genera did not differ in significant char-
acters. The two differ as follows (the
condition in EleutJierodactylus is en-
closed in parentheses ) : quadratojugal
absent (present), frontoparietal fonta-
nelle present (absent), nasals small and
widely separated medially (large and in
median contact), sacral diapophyses di-
lated (rounded), males with nuptial
asperities on thumb (lacking nuptial
asperities), aquatic tadpoles (develop-
ment direct — no tadpole stage ) , and am-
plexus inguinal ( axillary ) . The breeding
biology of Batrachyla is decidedly more
primitive than that of Eleutherodactylus
but approaches the condition of the lat-
ter in that the eggs are relatively large,
few in number, and laid in moist ter-
restrial situations (Barrio, 1967a, and
Cei, 1962a). In contrast to the eleu-
therodactyline pattern, tadpoles emerge
when the egg hatches and development
proceeds in the typical anuran manner.

Thoropa Cope, 1865

(Fig. 86)

Thoropa Cope, 1865, Rev. Nat. Hist., 5:110
[Type-species by monotypy, Cysiignathus
tuissiessii Eydoux and Souleyet, 1842].

Ololygon Fitzinger (1843) is often
cited as an older generic name for
Tlioropa. The type-species of Ololygon
is Hyla strigilata Spix, 1824 (by original
designation of Fitzinger, 1843). There-
fore, Ololygon Fitzinger is a synonym of
Hyla Laurenti, 1768.

Diagnostic definition. — 4) cervical
cotylar arrangement type I; 10) alary
processes of premaxillae directed dorsal-
ly and slightly anteriorly, relatively nar-
row at base; 11) palatal shelf of premax-
illa very narrow with elongate palatal
process; 13) palatal shelf of maxilla
broad, pterygoid process present; 14)
maxillary arch complete, quadratojugal
present; 15) nasals relatively large with
moderately long maxillary processes, sep-
arated medially; 16) nasals not in con-
tact with maxillae or pterygoids; 22)
epiotic eminences relatively well defined;
23 ) cristae paroticae long and narrow in
miliaris, short and relatively stocky in
lutzi and petropolitanus; carotid artery
passes dorsal to skull bones; 24) zygo-
matic ramus of squamosal relatively
short; 25) otic ramus of squamosal mod-
erately long, no otic plate; 26) squa-
mosal-maxillary angle 50-70°; 27) colu-
mella present; 28) prevomers relative-
ly small, entire, separated medially,
toothed; 29) palatines long and narrow,
expanded laterally, separated medially;
30) sphenethmoid entire, extending an-
teriorly to posterior edge of nasals or
not reaching nasals; 31) anterior ramus
of parasphenoid broad, keeled medially,
extending anteriorly to prevomers; 32)
parasphenoid alae oriented at right
angles to anterior ramus of parasphe-
noid, relatively short, not overlapped
laterally by median rami of pterygoids;
33) pterygoids large, anterior rami in
long contact with maxillae, not reaching
palatines; 34) occipital condyles large
in miliaris, small in lutzi and petropoli-

130

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 86. Lateral, dorsal, and ventral views
tanus, not stalked, moderately to widely
separated medially; 36) terminal pha-
langes T-shaped; 40) m. depressor man-
dihulae in two sHps; 41) pupil horizon-
tal; 42) males with median subgular
vocal sac; 43) body lacking glands; 44)
tongue large, oval, posterior edge free;

45) toes lacking webbing, bearing lateral
fringes, digital tips bulbous, somewhat
dilated, first finger shorter than second;

46) larvae with 2/3 tooth rows, labial
papillae broadly interrupted anteriorly;
47 ) ; 48 ) eggs large, few in number, laid
in lotic situations; 49) males 19-78, fe-
males 24-70 mm. SVL; 50) tympanum
visible externally; 51) tadpoles with
greatly flattened and attenuate bodies
and tails.

Composition. — Three species were
recognized in the revision by Bokermann
(1965): lutzi, miUaris, and petropoli-
tanus.

Distribution. — Mountains of south-
eastern Brasil between 12° and 30° S
latitude.

Remarks. — Thoropa is least different
from Batrachijla, although most authors
have considered it inseparable from

of skull of Thoropa Jutzi (KU 92850, X 4.5).
Eupsophus. Gallardo (1965) and Lynch
( 1971 ) demonstrated the distinctiveness
of these two genera. Lutz (1954) sug-
gested that Thoropa was closely related
to Cycloramplms. Tlwropa and Cyclo-
ramphiis belong to different tribes; this
distinction is supported by osteological,
non-osteological, and behavioral and
larvae data.

Odontophrynini New Tribe

Two genera are included in this tribe
— Odontophnjnus and Proceratophnjs
(the nominal genus Macrogenioglottus
is inseparable from Odontoplinjmis) .
Proceratophnjs is the generic name used
herein for the group previously called
Stomhiis. The members of this tribe
bear considerable external resemblance
to the Ceratophryinae, especially Cera-
tophrys. The ilia of the Odontophrynini
(Fig. 38) are nearly identical to those of
the Ceratophryinae, but the two groups
differ in many ways ( see subfamily char-
acters for Ceratophryinae and Telmato-
biinae). The Odontophrynini are dis-
tributed in non-forested and some for-

LYNCH: LEPTODACTYLOID FROGS

131

ested habitats in Argentina, Paraguay,
Uruguay, and along the eastern edge
of Brasil to Estado de Ceara. Like
the other primitive Telmatobiinae, the
Odontophrynini have an aquatic stage in
the Hfe history, and amplexus is axillary.
The following diagnostic characteristics
are the same in both genera: 3) trans-
verse processes of posterior presacral
\'ertebrae short; 5) cervical and second
xertebrae free; 6) cranial bones not in-
x'olved in dermostosis; 7) omosternum
lacking; 9) maxillary arch toothed, teeth
blunt, pedicellate; 11) palatal shelf of
premaxilla broad, weakly notched, pala-
tal process large; 12) facial lobe of max-
illa deep; 14) maxillary arch complete;
16) nasals in contact with maxillae and
pterygoids; IS) frontoparietal fontanelle
lacking; 20) frontoparietals not fused
with prootics; 21 ) temporal arcade lack-
ing; 22) epiotic eminences prominent;
23) cristae paroticae long and narrow;
carotid artery passes dorsal to skull
bones; 27) columella present; 28) pre-
vomers relatively small, entire, toothed,
narrowly separated medially; 29) pala-
tines large, narrowly separated medially,
bearing odontoid ridge, expanded lat-
erally; 30) sphenethmoid large, entire,
extending anteriorly to front edge of
nasals; 31) anterior ramus of parasphe-
noid narrow, pointed, not keeled; 32)
parasphenoid alae oriented at right
angles to anterior ramus, broadly over-
lapped laterally by median rami of
pterygoids; 33) pterygoids large, an-
terior rami long, in broad sutural contact
with maxillae, maxillary process of na-
sals, and palatines; 36) terminal pha-
langes knobbed; 37) alary processes of
hyoid plate on narrow stalks; 40) m. de-
pressor mandihulae in two slips; 41 ) pu-
pil horizontal; 42) males with median,
subgular vocal sac; 44) tongue large,
round, posterior edge free; 46) larvae
with median vent, 2/3 tooth rows, and
labial papillae broadly interrupted an-
teriorly; 47) amplexus axillary; 48) eggs
small, numerous, laid in gelatinous

masses in ponds; 50) tympanum con-
cealed.

The pectoral girdle is not as massive
as in the Telmatobiini, and the omoster-
num has been lost in the Odontophry-
nini. The occipital condyles are more
widely separated in Odontophnjnus than
in Proceratophnjs (Figs. 87-88). The
cervical cotylar pattern of both genera is
type II, although the cotyles are more
widely spaced in Odontophnjnus (Fig.
79). In several character complexes, the
Odontophrynini are intermediate be-
tween the Ceratophryinae and the Tel-
matobiini, but they also bear some re-
semblance to the EleutherodactyHni.

Odontophrynus Reinhardt and Liitken,

1862

(Fig. 87)

Odontophrynus Reinhardt and Liitken, 1862,
Vid. Meddel. Naturh. Foren., 13:159 [Type-
species by monotypy, Odontophrynus cul-
tripes Reinhardt and Liitken, 1862].

Hyperoodon Philippi, 1902, Supl. Bat. Chilenas,
p. 1 [Type-species by monotypy, Hyper-
oodon asper PhiUppi, 1902; preoccupied by
Hyperoodon Lacepede, 1804 (Mammalia:
Cetacea)].

Macrogenioglottus Carvalho, 1946, Bol. Mus.
Rio de Janeiro, (new ser. ) 73:1 [Type-
species by original designation, Macrogenio-
glottus alipioi Cai-valho, 1946].

Diagnosiic definition. — 4) cervical
cotylar arrangement type II, but cotyles
well separated medially; 8) sacral dia-
pophyses slightly dilated; 10) alary
processes of premaxillae directed pos-
terodorsally, long, relatively narrow at
base; 13) palatal shelf of maxilla broad,
pterygoid process small or lacking; 14)
maxillae not expanded posteriorly; 15)
nasals relatively large, keeled, narrowly
separated anteriorly; 17) nasals not in
contact with frontoparietals; 19) fronto-
parietals not ornamented except for
ridge around posterior half of the brain-
case roof; 24) zygomatic ramus of squa-
mosal long, tapering, widely separated
from maxilla; 25) otic ramus of squa-
mosal long, expanded medially into nar-
row otic plate; 26) squamosal-maxillary
angle 50-55°; 34) occipital condyles

132

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 87. Lateral, dorsal, and ventral views of skull of Odontophrtinus carvalhoi (KU 100441, X 3.2).

large, not stalked, median separation
moderate; 42) males with nuptial as-
perities on thumb; 43) parotoid and/ or
temporal or tibial glands present; 45)
toes about one-half webbed, outer meta-
tarsal tubercle present, inner metatarsal
tubercle enlarged and spade-like, digital
tips narrow, relatively few supernu-
merary tubercles on plantar surfaces, first
finger longer than second; 49) males 30-
60, females 34-70 mm. SVL.

Composition. — Savage and Cei (1965)
recognized four species (americanus,
carvalhoi, cultripes, and occidentalis).
The type species of Macrogenioglottus,
Odontophrijmis alipioi (Carvalho) [new
combination] is here added to the genus.

Distribution. — Semi-arid and arid
non-forested habitats of northern Argen-
tina, southern Bolivia and Paraguay,
Uruguay, and along the coastal provinces
of southeastern and eastern Brasil to
Bahia.

Remarks. — Carvalho (1946) named
Macrogenioglottus alipioi on the basis of
two specimens from Bahia, Brasil. The
genus was distinguished from all others

on the basis of the greatly enlarged m.
genioglottus, slightly dilated sacral dia-
pophyses, short coccyx, and slightly dif-
ferent positions of the prevomerine den-
tigerous processes. The myological dis-
tinction between Macrogenioglottus and
Odontophrynus remains valid, but with
the description of O. carvalhoi (Savage
and Cei, 1965 ) , the other differences be-
tween the two genera were mitigated.
The architecture of the temporal region
of alipioi was figured by Limeses ( 1965)
and is like that of other species of Odon-
tophrynus. Savage and Cei (1965) rec-
ognized two groups in Odontophrynus,
one for cultripes and occidentalis, and
another for americanus and carvalhoi.
Odontophrynus alipioi belongs to the
latter group and seems closely related to
carvallioi. The two species share many
characteristics but diff^er (insofar as is
known at present ) in some body propor-
tions, color pattern, and colors in life.
Odontophrynus alipioi has been collected
recently near Sao Paulo ( W. C. A. Boker-
mann, pers. comm.).

Boulenger (1882) confused Odonto-

LYNCH: LEPTODACTYLOID FROGS

133

phnjnus with Ceratophnjs, Lepidobatra-
chus, and Proceratophnjs. The cerato-
phryine leptodactyHds are readily sep-
arated from the two tehnatobiine genera
in that the derm of the head is fused
with the skull bones and a dermostosed
vertebral shield is present in the Cerato-
phryinae. In addition, the Ceratophryi-
nae ha\'e non-pedicellate teeth whereas
all other leptodactyHds (if dentate) have
pedicellate teeth. Odontophnjnus and
Proceratophnjs, especially those of the
bigibbosa group, are somewhat difficult
to separate on external characters alone.
The thenar surfaces of Proceratoplirys
are covered with numerous conical super-
numeraiy tubercles, whereas the thenar
surfaces of Odontoplirt/nus lack super-
numerary tubercles or have relatively
few, non-conical supernumerary tuber-
cles. Some, but not all, species of Odon-
tophnjnus have body glands (parotoid,
temporal, or tibial), whereas no species
of Proceratophnjs has glands. Osteo-
logically, the two genera are readily sep-
arated. Proceratophnjs has a complete
post-orbital bridge (squamosal only),
and Odontophnjnus has a "normal"
squamosal. Proceratophnjs has extensive
exostosis of the frontoparietal bones and
Odontophnjnus has no exostosis of the
frontoparietals but does have a ridge
around the posterior half of the fronto-
parietal shelf.

Philippi (1902) named Hijperoodon
asper on the basis of two specimens from
Montevideo, Uruguay. Subsequent au-
thors credited him with naming Hij-
peroodon as a new genus although
Philippi obviously credited the genus to
Dumeril and Bibron. Hijperoodon Phil-
ippi is probably a misspelling for Upero-
don Dumeril and Bibron, a microhylid
genus. Nieden (1923) tentatively re-
ferred Hijperoodon asper to Pahidicola;
the nominal species is currently consid-
ered a possible synonym of Phijsalaemus
gracilis (Gorham, 1966).

Cei ( 1958 ) correctly pointed out
that Hijperoodon asper is an Odonto-
phnjnus and tentatively suggested that

asper was identical with O. americanus.
Inexplicably, Savage and Cei (1965) did
not consider the nominal species in their
review of Odontophnjnus. There is no
question, in view of Philippi's descrip-
tion and Cei's ( 1958 ) publication of
Philippi's long-lost plates, that Hyper-
oodon asper is an Odontophnjnus, except
for Philippi's remarks that maxillaiy and
premaxillary teeth are lacking in the
types. Philippi further recorded the
presence of prevomerine teeth, suggest-
ing that his observation on the lack of
maxillary and premaxillary teeth was er-
roneous. Hyperoodon asper cannot be
considered a synonym of any species of
Physalaemus; it is here transferred to
the synonymy of Odontophnjnus ameri-
canus.

Proceratophrys Miranda-Ribeiro, 1920

(Fig. 88)

Proceratophrys Miranda-Ribeiro, 1920, Rev.
Mus. Paulista 12:301 [Type-species by
monotypy, Ceratophnjs bigibbosa Peters,
1872].

Diagnostic definition. — 4) cervical
cotylar arrangement type II, cotyles
closely approximated; 8) sacral dia-
pophyses rounded; 10) alary processes of
premaxillae long, strongly directed pos-
terodorsally, except in bigibbosa group,
relatively narrow at base; 13) palatal
shelf of maxilla broad, pterygoid process
prominent; 14) maxillae slightly ex-
panded posteriorly; 15) nasals relatively
narrow, keeled, separated medially
(boiei group) or in contact (bigibbosa
group) medially; 17) nasals in contact
with frontoparietals; 19) frontoparietals
bear lateral crests which meet posterior-
ly; frontoparietal crests heavily exostosed
posteriorly in cristiceps and probably in
bigibbosa; 24) zygomatic ramus of squa-
mosal broad and elongate, in sutural
contact with maxilla, weakly exostosed;
25) otic ramus of squamosal large, exo-
stosed, expanded medially into relatively
large otic plate; 26) squamosal-maxillary
angle 40-50°; 34) occipital condyles
large, not stalked, closely juxtaposed;

134

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 88. (A-C) Dorsal, ventral and lateral views of skull of Proceratophnjs hoiei (KU 93076) and (13-
E) dorsal and lateral views of skull of P. cristiceps ( KU 106273). All X 2.2.

42) males lacking nuptial asperities on
thumb; 43) body lacking glands; 45)
toes free of webbing, usually with lateral
fringes, outer metatarsal tubercle pres-
ent, inner metatarsal tubercle small or
enlarged and spade-like, digital tips nar-
row, numerous conical supernumerary
thenar and plantar tubercles, first finger
longer than second; 49) adults 30-95
mm. SVL.

Composition. — Eight of the nominal
species listed as Ceratoplirys by Gorham
( 1966) belong to this genus: appendicu-
lata, Ingibbosa, boiei, cristiceps, fryi,
goyanus, renalis, and schirchi. Gorham
(1966) did not list Stombus melanopo-
gon Miranda-Ribeiro, 1926. Bokermann
(1966) considered boiei, melanopogon,
renalis, and schirchi synonymous and
used boiei, the oldest name. He also
considered goyanus a synonym of cris-
ticeps.

Distribution. — The lowland zone east
of the Brasilian Highlands from Forta-
leza (Ceara) to Santa Catarina, Brasil,
and adjacent Misiones Province, Argen-
tina.

Remarks. — Almost all of the litera-
ture pertaining to species of this genus
has accumulated under the generic name
Stombus, a synonym of Ceratoplirys (see
pp. 107-10). Reig and his students have
consistently argued that Stotnbus (auc-
torum) is generically distinct, but they
have also repeatedly questioned the ge-
neric position of cristiceps. I include
cristiceps in the genus Proceratophrys
and consider bigibbosa its closest rela-
tive; this arrangement is similar to that
used by Reig in that cristiceps is not con-
sidered especially closely related to ap-
pendiculata, boiei, and fryi. The differ-
ences between bigibbosa and cristiceps
on the one hand, and appemlicnlata,

LYNCH: LEPTODACTYLOID FROGS

135

boiei, and fnji on the other, are not of
the magnitude I would use at the ge-
neric level. The five species form a
group on the basis of the thenar and
plantar tubercle arrangement, body
shape, and the architecture of the tem-
poral region. The two species groups
differ in head shape and the correlated
and underlying cranial architecture
(snout is elongate and sloping in boiei
group, blunt and short in higihhosa
group) and in the development of cra-
nial exostosis (Fig. 88). The differences
in musculature that Limeses ( 1964,
1965) cited as suggestive of separate
genera are trivial differences; greater
ranges of variation occur within Odon-
tophnjnus and several other frog gen-
era. The species of the higihhosa group
are less unlike Odontophnjnus in exter-
nal features than are the species of the
hoiei group. In the hoiei group, the eye-
lids are provided with elongate "horns,"
whereas in the higihhosa group, the eye-
lid has only a few large tubercles.

Grypiscini Mivart, 1869

Grypiscina Mivart, 1869:295.
Cyclorhaniphinae Lutz, 1954:175.
Gycloramphiinae: Gallardo, 1965:84.

Three genera are included in this
tribe — Crossodactylodes, Cijcloramphus,
and Zachaenus. The nominal genus
Craspedoglossa is here considered to be
a synonym of Zachaenus. The tribe
name is based on Grypiscus Cope, a syn-
onym of Cycloramphus Tschudi. At
first glance, this group seems to be highly
heterogeneous, especially when one con-
"7siders the supposed relationships of the
leptodactylid genera as recognized by
herpetologists in the 1910-1930 period
when the following genera (all of this
group) were recognized: Craspedoglos-
sa, Cycloramphus, Grypiscus, lliodlscus,
Oocormus, and Zachaenus. Miranda-
Ribeiro (1926), Lutz (1954), and Coch-
ran (1955) considered Zachaenus to be
closely related to Ceratophrys. Oocor-
mus was often recognized even by Lutz
after she pointed out that it was a syn-

onym of Zachaenus (1944). Various
authors including Noble ( 1931 ) sug-
gested that Cycloramphus was closely
related to Tehnatohius. Cycloramphus,
Iliodiscus, and Grypiscus were usually
considered valid until Bokermann ( 1951 )
pointed out that they were not generi-
cally distinct. Lutz (1954) included
only Cyclorhamphus (sic) and Thoropa
in the subfamily, Lutz and Carvalho
( 1958) considered Paratelmatobius to be
a generic link between Cycloraynphus
and Batrachophrynus and Tehnatohius,
and Gallardo ( 1965 ) placed Craspedo-
glossus (sic), Cycloramphus, Holoaden,
and Zachaenus in the subfamily Gyclo-
ramphiinae. All of these authors used
labile characters (head shape and toe
webbing) to define their groupings.
Most of the genera that have been con-
sidered related to Cycloramphus are bur-
rowing frogs and have long, flat snouts.

The following diagnostic character-
istics are the same in the included gen-
era: 3) transverse processes of posterior
presacral vertebrae not shortened; 4)
cervical cotylar arrangement type I; 5)
cervical and second vertebrae free; 6)
cranial bones not involved in dermosto-
sis; 9) maxillary arch toothed, teeth
blunt, pedicellate; 12) facial lobe of
maxilla deep, not exostosed; 14) maxil-
lary arch complete, maxilla expanded
posteriorly, quadratojugal deep; 15) na-
sals relatively large, in broad median
contact; 17) nasals in contact with fron-
toparietals; 18) frontoparietal fontanelle
lacking; 20) frontoparietal not fused to
prootic; 21) temporal arcade lacking;
23) cristae paroticae very short, stocky;
carotid artery passes dorsal to skull
bones; 24) zygomatic ramus of squa-
mosal attenuate, curved, widely sepa-
rated from maxilla; 25) otic ramus of
squamosal long, curved medially and
expanded medially to form otic plate
which rests on crista parotica; 34) occip-
ital condyles relatively small, not stalked,
widely separated medially; 37) alary
processes of hyoid plate on narrow
stalks; 40) m. depressor mandihulae in

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

two slips; 41) pupil horizontal; 47) am-
plexus axillary.

The developmental pattern of Cijclo-
ramphus and Zacliaenus is intermediate
between the typical pattern of an aquatic
tadpole and the eleutherodactyline pat-
tern of direct development ( Lutz, 1944 ) .
The eggs are deposited in terrestrial
situations (usually in very wet leaves on
the forest floor) and the tadpole hatches
and lives in the moist, decomposing
gelatinous mass. The tadpoles can sur-
vive in an aquatic medium but do not
feed (Lutz, 1944). Crossodactylodes
lays only a few large eggs in bromeliads
and the tadpole develops in the moist
axillae of the bromeliads (W. C. A.
Bokermann, pers. comm.).

Holoaden, Paratelmatohius, and
Thoropa do not agree with the diagnos-
tic characteristics listed above and are
included in other groups; Holoaden is in
the Eleutherodactylini, Paratelmatohius
in the Leptodactylinae, and Tlwropa in
the Alsodini.

The three genera of the tribe occur
in forested montane areas in southeast-
ern Brasil and Uruguay. Two of the

species presently placed in Zachaenus
(roseus and sawayae) are not members
of that genus. Their status is discussed
in the account of Zachaenus.

Crossodactylodes Cochran, 1938

(Fig. 89)

Crossodactylodes Cochran, 1938, Proc. Biol.
Soc. Washington, 51:41 [Type-species by
original designation, Crossodactylodes pintoi
Cochran, 1938].

Diagnosiic definition. — 7) omoster-
num relatively small; 8) sacral dia-
pophyses dilated; 10) alary processes of
premaxillae directed posterodorsally,
broad at base; 11) palatal shelf of pre-
maxilla relatively broad, broadest lat-
erally, with long palatal process; 13)
palatal shelf of maxilla broad anteriorly,
narrow over most of its length, pterygoid
process lacking; 16) nasals in tenuous
contact with maxillae, not in contact
with pterygoids; 17) nasals in tenuous
contact with frontoparietals, narrowly
separated from frontoparietals; 19) fron-
toparietals not ornamented, lacking sa-
gittal crest; 22) epiotic eminences obso-
lete; 26) squamosal-maxillary angle

Figure 89. Lateral, dorsal, and ventral views of skull of Crossodactylodes pintoi (paratype, USNM

120611, X 8).

LYNCH: LEPTODACTYLOID FROGS

137

about 45°; 27) columella absent; 28)
prevomers small, edentate, dentigerous
rami almost completely lost, widely sep-
arated medially; 29) palatines narrow,
widely separated medially, lacking odon-
toid ridges; 30) sphenethmoid entire, ex-
tending anteriorly beneath nasals; 31)
anterior ramus of parasphenoid broad,
not keeled; 32) parasphenoid alae ori-
ented at right angles to anterior ramus,
narrowly overlapped laterally by median
rami of pterygoids; 33) pterygoids rela-
tively small, anterior rami extending to
middle of orbit, ventral pteiygoid flange
small; 36) terminal phalanges Y-shaped,
lateral processes long and slender; 37-
39); 42) males with median subgular
vocal sac; males with cluster of spines
on thumb; 43) body lacking glands; 44)
tongue oval, posterior one-third free,
non-boletoid; 45) toes not webbed, lack-
ing lateral fringes, outer metatarsal tu-
bercle present, inner metatarsal tubercle
not enlarged and spade-like, digital tips
enlarged into pads, first finger shorter
than second; 46) tadpoles semi-aquatic;
48 ) eggs large, few in number, deposited
in terrestrial bromeliads; 49) males 15-
17.5 mm. SVL; 50) tympanum absent
( not hidden, as stated by Cochran, 1938,
1955).

Composition. — Monotypic.

Distribution. — The Coastal Ranges of
Guanabara and Espirito Santo, Brasil.

Remarks.— Cochran (1938, 1955)
considered Crossodactylodes to be re-
lated to Crossodactyhis (Elosiinae).
She based her opinion on the presence
of a cluster of spines on the thumbs of
the males of both genera and the erro-
neous opinion that both genera have der-
mal glands on the top of each digital
pad. Cochran noted that when the digi-
tal tips of Crossodactylodes were dried
out, a weak furrow appeared in the cen-
ter of the digital pad; she considered this
condition a precursor of the condition
seen in elosiines (distinct dermal glan-
dular pads). The apparent glands ob-
served by Cochran are an artifact result-
ing from the presence of Y-shaped ter-

minal phalanges. Cochran pointed out
that the two genera shared the loss of
pre vomerine teeth; the pre vomerine
bones are much smaller in Crossodacty-
lodes than in Crossodactyhis, which
usually has larger prevomerine dentiger-
ous processes and rarely prevomerine
teeth.

The architecture of the temporal re-
gion and the size of the roofing bones of
Crossodactylodes are like the condition
seen in Cycloramphus and Zachaenits.
The other cranial characters of Crosso-
dactylodes are not contrary to the condi-
tions which obtain in Cycloramphus and
Zachaenus, although Crossodactylodes is
separable from these two genera by
many skull characters. The data on
breeding biology were provided by
Werner Bokermann (in litt.) and suggest
that Crossodactylodes is more closely re-
lated to Cycloramphus and Zachaenus
than to the Eleutherodactylini, which it
resembles in many osteological features.

Cycloramphus Tschudi, 1838
(Fig. 90)

Cycloramphus Tschudi, 1838, Classif. Batr., p.
81 [Type-species by monotypy, Cycloram-
phus fulginosiis Tschudi, 1838].

Cyclorhamphus Agassiz, 1846, Nomencl. Zool.,
index, p. 110 [Emendation of Cycloramphus
Tschudi, 1838, hence taking same type-
species].

Pithecopsis Giinther, 1859, Cat. Bat. Sal. British
Mus., p. 22 [Type-species by monotypy,
Pithecopsis fulifiinosus (Tschudi, 1838)].

Grypiscus Cope, 1867, J. Acad. Nat. Sci. Phila-
delphia, (2)6:206 [Type-species by mono-
typy, Grypiscus ui>ihrinus Cope, 1867].

lliocliscus Miianda-Ribeiro, 1920, Rev. Mus.
Paulista, 12:267 [Type-species by subse-
quent designation (Bokermann, 1951),
Telmatohius hrasiliensis Steindachner, 1864;
his designation is hereby rejected (see "Re-
marks" ) and the type-species is here desig-
nated as lliocliscus cluhius Miranda-Ribeiro,
1920].

Niedenia Ahl, 1923, Zool. Anz., 58:107 [Type-
species by monotypy, Niedenia spinulifer
Alil, 1923].

Diagnostic definition. — 7 ) omoster-
num moderate-sized; 8) sacral dia-
pophyses rounded to very slightly di-
lated; 10) alary processes of premaxillae

138

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 90. Lateral, dorsal, and ventral views of skull of Cycloramphus eletitherodactijlus (KU 92785,

X 3.6).

directed posterodorsally, relatively broad
at base; 11) palatal shelf of premaxilla
of moderate depth, not notched, palatal
process short; 13) palatal shelf of maxil-
la relatively narrow, pterygoid process
large; 16 ) nasals in contact with maxillae,
not with pterygoids; 19) frontoparietals
not ornamented, bearing large sagittal
crest; 22) epiotic eminences prominent
posteriorly, obsolete anteriorly; 26 ) squa-
mosal-maxillary angle less than 43°,
measurement difficult because of curva-
ture of both elements; 27) columella
present; 28) prevomers present, entire,
moderately large, toothed, separated
medially; 29) palatines broad, widely
separated medially, bearing small odon-
toid ridges; 30) sphenethmoid entire,
extending anteriorly beneath nasals, not
visible dorsally or only a small area visi-
ble between junctions of nasals and fron-
toparietals; 31) anterior ramus of para-
sphenoid broad, not keeled; 32) para-
sphenoid alae oriented at right angles to
anterior ramus, broadly overlapped by
median rami of pterygoids; 33) ptery-

goids relatively small, anterior rami ex-
tending to middle of orbit, pterygoids
bearing large ventral flange — cf. Fig. 25;
36) terminal phalanges knobbed; 42)
males with median subgular vocal sac;
males lacking nuptial asperities except
for C. ohausi which has a cluster of
spines on each thumb; 43) inguinal
glands present; 44) tongue large, round,
semi-boletoid; 45) toes free of webbing,
bearing lateral fringes, or partly to fully
webbed, outer metatarsal tubercle pres-
ent, inner metatarsal tubercle not en-
larged and spade-like, digital tips nar-
row, first finger not longer than second;
46) larvae with very brief tadpole stage,
semi-aquatic, vent median, 1/1 tooth
rows, labial papillae broadly interrupted
anteriorly; 48) eggs laid in moist ter-
restrial situations, hatch near end of lar-
val period, eggs large, few in number
(Lutz, 1929); 49) adults 30-55 mm. SVL;
50) tympanum concealed.

Composition. — Seven to nine species
are recognized depending on the author.
Gorham (1966) listed eight species —

LYNCH: LEPTODACTYLOID FROGS

139

asper, cUringshoefensi, eleutherodactylus,
ftilginosiis, gramdosus, neglectus, ohausi,
and iimhrinus, whereas Bokerniann
(1966) recognized houlengeri, diiJ)ius,
and pinderi as valid (which Gorham
considered synonyms of other names)
and placed timhrinus in the synonymy of
ftdginosus. The most recent revision of
the genus is that of Bokerniann ( 1951 ) ,
although Cochran ( 1955 ) studied a large
part of the genus. In view of the differ-
ences of opinion as to how many, and
which, species are valid, a thorough ge-
neric review is desirable.

Distribution. — Forested habitats in
southeastern Brasil.

Remarks. — Within comparatively re-
cent times ( Miranda-Ribeiro, 1926,
Noble, 1931), the genus Cycloramphus
was divided into three genera (Cyclo-
ramphus, Grypiscus, and Iliodisctis).
Bokermann (1951) and Cochran (1955)
combined the three into a single genus.
Cochran (1955) suggested that the ge-
nus is heterogeneous because C. ohausi
mitigates some of the differences be-
tween Ceratophrys, Cycloramphus, and
Crossodactyhis. Her statements reflect a
philosophy of single-character classifica-
tion and do not accurately indicate the
homogeneity of the genus Cycloram-
phus.

The generic partitioning of Cyclo-
ramphus was based on the variation in
webbing of the toes and Cope's argu-
ment that the presence of pseudoteeth
on the lower jaw of umbrinus justified
generic distinction. I have not observed
odontoids on the lower jaw of any spe-
cies of the genus, although Noble (1922)
figured a serrate lower jaw of a cotype
of umbrinus. Six species of the genus
(asper, diringshoefensi, dubius, eleu-
therodactylus, granulosus, and pinderi)
lack webbing or lateral fringes on the
toes; three species (boidengeri, fidgino-
sus, and neglectus) have one-half to
fully webbed toes. The two gi'oups are
bridged by ohausi which has basal
webbing and lateral fringes on the toes.
The basal webbing probably is best re-

garded as the broadened junction of the
fringes. I consider the lack of webbing
to be primitive, because the allied Za-
chaenus lacks webbing.

Bokermann (1951) designated Telma-
tobius hrasiliensis Steindachner (=zCy-
cloramphus fulginosus) as the type-spe-
cies of Iliodiscus Miranda-Ribeiro, 1920.
This action rendered Iliodiscus a strict
synonym of Cycloramphus. However,
Miranda-Ribeiro (1920) placed the
webless species of Cycloramphus in
Iliodiscus; Telmatobius brasiliensis was
included in Cycloramphus, not Iliodis-
cus, and therefore cannot be considered
for subsequent designation as the type-
species of Iliodiscus. Accordingly, Bok-
ermann's ( 1951 ) restriction is hereby
rejected. Miranda-Ribeiro (1920) in-
cluded four nominal species in Iliodiscus
(dubius, eleutherodactylus, pinderi, and
semipalmatus). Gorham (1966) listed
dubius as the type-species. I designate
/. dubius Miranda-Ribeiro, 1920, as the
type-species of Iliodiscus Miranda-Ri-
beiro, 1920.

Zachaenus Cope, 1866
(Figs. 91-93)

Zachaenus Cope, 1866, J. Acad. Nat. Sci. Phila-
delphia, 6:94 [Type-species by original
designation, Cystignathus parvulus Girard,
1854].

Oocormus Boulenger, 1905, Ann. Mag. Nat.
Hist., (7)16:181 [Tyi^e-species by mono-
typy, Oocormus microps Boulenger, 1905].

Craspedoglossa L. Miiller, 1922, Blatter Aquar.
Terr., 33:167 [Type-species by monotypy,
Craspedoglossa sanctaecatharinae L. Miiller,
1922].

Diagnostic definition. — 7) omoster-
num relatively large; 8) sacral dia-
pophyses rounded; 10) alaiy processes of
premaxillae directed sharply posterodor-
sally, relatively broad at base; 11) pala-
tal shelf of maxilla relatively deep, not
notched, palatal process moderate-sized;
13) palatal shelf of maxilla of moderate
width, pterygoid process lacking; 16)
nasals in contact with maxillae, not with
pterygoids; 19) frontoparietals not orna-
mented except for prominent sagittal
crest and supraorbital processes; 22)

140

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 91. Dorsal, lateral, and \'entra] views of
skuW oiZachaemisparvulusiKU 107090, X 7.5).

epiotic eminences obsolete; 26) sqiia-
mosal-maxillary angle about 45°; 27)
columella present; 28) prevomers rela-
tively large, entire, toothed, narrowly
separated medially; 29) palatines slen-
der, widely separated medially, no odon-
toid ridges; 30) sphenethmoid entire,
usually not visible dorsally; 31) anterior
ramus of parasphenoid narrow, not
keeled; 32) parasphenoid alae oriented
at right angles to anterior ramus, broadly
overlapped laterally by median rami of
pterygoids; 33) pterygoids relatively
small, anterior rami extending to middle
of orbit, large ventral flange; 36) termi-

nal phalanges knobbed; 42) males with
median subgular vocal sac; males lack-
ing nuptial asperities; 43) body lacking
glands; 44) tongue round, semi-boletoid;
45) toes lacking webbing and lateral
fringes, outer metatarsal tubercle pres-
ent, inner metatarsal tubercle enlarged,
not spade-like, digital tips narrow, first
finger as long as second; 46) develop-
ment abbreviated, tadpole semi-aquatic,
vent median, 1/1 tooth rows, labial pa-
pillae broadly interrupted anteriorly
(Lutz, 1944); 48) eggs large, few in
number, deposited in moist, terrestrial
situation, larvae hatch and remain in

Figure 92. Dorsal, lateral, and ventral xiews of
skull of Zachaenus stejnegeri (KU 92742, X 7.5).

LYNCH: LEPTODACTYLOID FROGS

141

Figure 93. Body outlines and sides of heads of (A-B) Zachaeniis stejnegeii (KU 92744) and (C-D)

Z. parvtilus (KU 93078). All X 2.

gelatinous mass until metamorphosis;
49) adults less than 30 mm. SVL; 50)
tympanum concealed.

Composition. — The nominal species
of Craspedoglossa (holitoglossa, sanctae-
cathorinae, and stejnegeri) and of Za-
chaeniis (parvidus, roseus, and sawaijae).
Bokermann (1966) considered sanctae-
catharinae to be a synonym of bolitoglos-
sus. I do not consider roseus or sawayae
members of this genus ( see Remarks ) .

Distribution. — Southern and south-
eastern Brasil.

Remarks. — Lutz (1944) demonstrated
that Oocormus microps is a synonym of
Zachaenus parvidus. Parker (1926)
noted the striking similarities in color
pattern and proportions between Oocor-
mus microps (^Zachaenus parvulus)
and SmintluUus hrasiUensis (=Eupark-
erella), which occur sympatrically in
southeastern Brasil. Boulenger ( 1905 )
confused them and included Euparker-
ella in the syntypic series of Oocormus
microps as juvenile specimens. The two

genera belong to different ti-ibes and
can be distinguished externally only by
the shape and lengths of the fingers and
toes (Fig. 94).

Craspedoglossa is here placed in the
synonymy of Zachaenus for the first
time. Cochran (1955) pointed out that
the two nominal genera might be best
combined but separated them on the
basis of the axillary patagium of Zachae-

FiGURE 94. Hands of three frogs of the Telmato-
biinae. (A) Zachaenus parvulus (KU 93078, X
4), (B) Euparkerella hrasiUensis ( KU 112370,
X 6.6), and (C) Scydirophnjs sawayae (USNM
125530, X 6.6).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

nus (see Fig. 93). Tehnatohufo is the
only other leptodactyHd with an axillary
patagium, although the loose, "baggy"
skin of Batrachophnjmis and the sti-ictly
aquatic Tehnotohius produces a poorly
defined patagium. The snout is more
sloping in Zachaenus parvitlus than in
Craspedoglossa (sensu strictu). The axil-
lary patagium is used as a species-group
character in other groups of frogs (for
example, the Hijla marmorata and H.
godmani groups), and is best regarded
as a species-group character here as
well. In all other characters used in the
generic diagnoses, Craspedoglossa and
Zachaenus are identical.

Cope (1890) named a second species
of the genus (roseiis) based on a single
specimen from Port Otway, Patagonia;
the unifiue holotype is now a macerated
heap of fragments (Cochran, 1955,
1961b). The original description in-
cludes several points that clearly disasso-
ciate roseiis from Zachaenus (tympanum
visible, tongue not boletoid, prevomerine
dentigerous processes small and round,
outer metatarsal tubercle lacking, and
toes fringed). The osteological data
provided by Cope (nasals small, widely
separated medially, and frontoparietals
complete, i.e., no fontanelle) and those
observed by me (pterygoid lacking
ventral flange, zygomatic ramus of squa-
mosal short and straight otic ramus of
squamosal not curved medially but ex-
panded medially into small otic plate)
clearly disassociate roseus from Zachae-
nus. Unfortunately, the present data are
insufficient for generic assignment. Za-
chaenus roseus Cope is tentatively con-
sidered a species inquirenda in the fam-
ily Leptodactylidae, probably in the Tel-
matobiinae.

Cochran (1953) named a single spec-
imen of a frog from Parana, Brasil, as a
third species of Zachaenus (saivayae).
The species is clearly not a Zachaenus,
although its relationships are not appar-
ent; only the holotype is known, and no
osteological observations can be made.
Zachaenus sawayae is considered by me

to be the type-species of a new genus of
the Telmatobiinae; the tribal relation-
ships are not apparent. The new genus
is named at the end of the account of the
Telmatobiinae.

Eleutherodactylini Lutz, 1954
Eleutherodactylinae Lutz, 1954:175.

Lutz (1954) proposed this subfamily
for the inclusion of Eleutlierodactyhis
alone; as was the case with the Cyclo-
rhamphinae, she did not provide any
diagnostic statements for the new group.
Gallardo (1965) included Basanitia,
Ctenocranius, Eleutherodactyhis, Micro-
hairachyhis, and Syrrliophus in the Eleu-
therodactylinae. I include the following
genera in the tribe Eleutherodactylini:
Amhhjphryniis, Eleutherodactyhis, Eu-
parkereUa, Holoaden, Hylactophryne,
Ischnocnema, Niceforonia, Sminthillus,
Syrrhophus, and Tomoductylus. The
nominal genera Basanitia, Nohlella,
Phrynanodus, and Trachyphrynus are
considered to be synonyms of Eleuthero-
dactyhis (Lynch, 1968c, 1968d). The
nominal genera Nohlella Barbour and
Pseudohyla Andersson are considered to
be synonyms of Eleutherodactyhis (see
below ) .

The following diagnostic characteris-
tics are the same in all of the included
genera: 3) tranverse processes of pos-
terior presacral vertebrae not shortened;

4) cervical cotylar arrangement type I;

5) cervical and second vertebrae not
fused; 8) sacral diapophyses rounded;
9) maxillary arch usually toothed, if
toothed, teeth blunt, pedicellate; 14)
maxillary arch complete, maxillae taper-
ing posteriorly, quadratojugal shallow;
17 ) nasals not in contact with frontopari-
etals; 18) frontoparietal fontanelle usual-
ly absent; 21) temporal arcade lacking;
23) carotid artery passes dorsal to skull
bones; 41) pupil horizontal; 42) males
lacking nuptial asperities; 45) outer
metatarsal tubercle present, inner meta-
tarsal tubercle not spade-like; 46) de-
velopment direct in known species; 47)
amplexus axillary in known species; 48)

LYNCH: LEPTODACTYLOID FROGS

143

eggs relatively large, few in number, de-
posited in terrestrial situations, brome-
liads, etc., in all known species.

Except for the nature of the T-
shaped terminal phalanges, the skeleton
of the eleutherodactylines is relatively
generalized. The sternum is cartilagi-
nous, an omosternum is usually present
and relatively large, the cervical cotyles
are widely separated medially, the trans-
verse processes of the presacral vertebrae
are neither greatly expanded nor short-
ened, the sacral diapophyses are rounded
or only sHghtly dilated, the ilia are of the
leptodactyline type, all cranial bones
are present although a few species have
lost the prevomerine bones or the colu-
mellae. The cranial bones are not der-
mostosed but exostosis is developed in
several groups. The nasals are large and
usually in median contact, and the fron-
toparietal fontanelle is rarely developed.
There is considerable variation in the
size and shape of the zygomatic ramus
of the squamosal among the genera and
species of this tribe, and most of the
variation is observed within Eleuthero-
dactyhis.

Morphologically, the tribe Eleuthero-
dactylini is difficult to define. The rela-
tionships of ten genera I include in this
tribe are not entirely obvious. The defi-
nition of the group rests solely on the
mode of reproduction, and the included
genera are judged to be related because
the various sections of the tribe can be
tied together through the use of several
different character complexes. Species of
the following genera are known to lay
terrestrial eggs and to lack a free tadpole
stage: Eleutherodactylus, Holoaden, Hy-
lactophryne, Sminthillus, Syrrhophus,
and Tomodactylus. Euparkerella and
Niceforonia probably lay large terrestrial
eggs, judging from the size of mature
eggs and the oviducts. Reproductive
data are not available for Amhlyphrynus
and Ischnocnema. Females of the latter
genus have moderately large, unpig-
mented eggs, but the eggs are not so
large as to be suggestive of direct devel-

opment. However, the eggs of Ischno-
cnema are no smaller than those of Eleu-
therodactyJus maussi, which exhibits di-
rect development (Heatwole, 1962).
Eleutherodactylus, Sminthillus, Syrrho-
phus, and Tomodactylus have broadly
T-shaped terminal phalanges. The ter-
minal phalanges of Euparkerella are
broad and small, but could not be accu-
rately described as T-shaped. The ter-
minal phalanges of the other five genera
(Amhlyphrynus, Holoaden, Hylacto-
phryne, Ischnocnema, and Niceforonia)
of the tribe are knobbed. Amhlyphrynus
bears considerable resemblance to the
Eleutherodactylus cornutus group and
differs only in the nature of the terminal
phalanges. Hylactophryne and Ischno-
cnema bear considerable resemblance to
one another and to the Eleutherodacty-
lus hinotatus and guentheri groups; these
genera also differ only in the nature of
the terminal phalanges. Niceforonia is
superficially similar to Eupsophus and to
the Eleutherodactylus unistrigatus com-
plex (several species groups are in-
volved). Niceforonia differs from Eup-
sophus in several osteological characters
as well as in reproductive pattern, but
differs from the Eleutherodactylus uni-
strigatus complex only in the nature of
the terminal phalanges. Holoaden is not
obviously related to any group of Eleti-
therodactylus, but is osteologically simi-
lar to Niceforonia.

Amhlyphrynus Cochran and Coin, 1961

Amhhjphnjnus Cochran and Coin, 1961, Fieldi-
ana, Zool., 39:543 [Type-species by orig-
inal designation, Amhhjphnjnus ingeri Coch-
ran and Coin, 1961].

Diagnostic definition. — 7 ) omoster-
num present; 9) maxillary arch toothed;
10) alary processes of premaxillae di-
rected posterodorsally; 11); 12) facial
lobe of maxilla deep, not exostosed; 13);
15) nasals large, in broad median con-
tact; 16) nasals in tenuous contact with
maxillae, not in contact with pterygoids;
19) frontoparietal bears large, well de-
fined, exostosed, lateral crests; 20); 22)

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

epiotic eminences obsolete; 23) cristae
paroticae long, narrow; 24) zygomatic
ramus of squamosal of moderate length,
widely separated from maxilla; 25) otic
ramus of squamosal short, not expanded
medially into otic plate; 26); 27) colu-
mella present; 28) prevomers large, en-
tire, toothed, in median contact; 29)
palatines large, narrowly separated me-
dially; 30) sphenethmoid entire, extend-
ing anteriorly beneath nasals; 31) an-
terior ramus of parasphenoid broad, not
keeled, pointed anteriorly; 32) para-
sphenoid alae oriented at right angles to
anterior ramus, broadly overlapped lat-
erally by median rami of pterygoids; 33 )
pterygoids of moderate size, lacking ven-
tral flange, anterior rami not reaching
palatines, median rami long; 34) occip-
ital condyles large, not stalked, widely
separated medially; 36) terminal pha-
langes knobbed; 37-40); 42); 43) body
lacking glands; 44) tongue large, round,
posterior edge free; 45) toes free of
webbing, digital tips narrow; 46-48);
49) the two known specimens are 51.5
and 83.0 mm. SVL; 50) tympanum visi-
ble externally.

Composition. — Monotypic.

Distrihiition. — Known from two lo-
calities in the Andes of central Colombia
(850-2350 meters). Peracca's (1914)
record of an Eleutlierodacti/ltis cornutiis
from the highlands in Departamento
Antioquia, Colombia, may refer to this
species.

Remarks. — Cochran and Coin ( 1961 )
suggested that AmhJijphnjnus is a mem-
ber of the "broad-headed leptodactylid"
group which includes the ceratophryine
genera, Proceratophrys, and Zachaenus.
They suggested that Amhiijphnjnus was
probably most closely related to Zaclme-
nus. The resemblance between these
two genera is spurious. This association
can only be made by comparison of de-
scription of characters and will not bear
up against specimen comparison or the
additional osteological characteristics.
The suggestion that this species is allied
with the ceratophryine leptodactylids is

contradicted by the lack of morphologi-
cal agreement between the two groups.
The ceratophryines have large, casqued
skulls with extensive dermostosis and
exostosis, a distinctly different type of
cervical-occipital articulation, non-pedi-
cellate teeth, expanded transverse proc-
esses of the anterior presacral vertebrae
and shortened transverse processes of
the posterior presacral vertebrae, and a
dermostosed vertebral shield.

In many respects, Amblyplirynus re-
sembles the large-headed frogs of the
Eleiitlwrodactyhis cornutus group. The
holotype of Amhlyphrynus ingeri was
first reported by Dunn ( 1944 ) as an
EleutJierodactylus cornutus. The only
characteristic separating these two
groups is the nature of the terminal pha-
langes. When additional specimens of
A. ingeri become available an effort must
be made to determine if the terminal
phalanges of the hind feet are knobbed
or T-shaped. As pointed out previously,
the terminal phalanges of the fingers
may be knobbed and those of the toes
T-shaped. This pattern is observed in
several groups of Eleutherodactylus.

The only specimens of this species
available were studied through the use
of stereo-radiographs.

Eleutherodactylus Dumeril and Bibron,

1841

(Figs. 95-99)

Cornufcr Tschudi, 1838, Classif. Batr., p. 28
[Type-species by monotypy, Cornufer uni-
color Tschudi, 1838 {=Eleutherodactijlus
inoptatus) . Suppression of Tschudi's names
was requested by Zvveifel ( 1966)].

Elentherodactijlus Dumeril and Bibron, 1841,
Erp. gen., 8:620 [Type-species by mono-
typy, Hylodes rnaiiinicensis Tschudi, 1838.
Myers ( 1962 ) Hsted the type-species desig-
nation as l)y original designation. The name
Elciitherodactyhis was included in the syn-
onymy of Hylodes martinicensis by Dume-
ril and Bibron (1841). Apparently they
had planned to use the generic name in
their £rpetoIogie generate until Tschudi
(1838) named their martinicensis in the
genus Hylodes].

Hylodes Fitzinger (non Hylodes Fitzinger, ,
1826), 1843, Syst. Rep., p. 31 [Type-species i

LYNCH: LEPTODACTYLOID FROGS

145

by original designation, Hijlodes martinicen-
sis Tschudi, 1838].

Euhyas Fitzinger, 1843, Ibid., p. 31 [Type-
species by original designation, Hylodes
ricordii Dumeril and Bibron, 1841].

Crau'^astor Cope, 1862, Proc. Acad. Nat. Sci.
Philadelphia, p. 153 [Type-species by sub-
sequent designation ( Dunn and Dunn,
1940), Hijlodes fltzingeri O. Schmidt,
1858].

Strahomantis W. Peters, 1863, Mtber. k. Preuss.
Akad. Wiss., Berlin, p. 405 [Type-species
by monotypy, Strahomantis biporcatus W.
Peters, 1863].

Leitjla Keferstein, 1868, Ark. Naturges., 34:296
[Type-species by monotypy, Leiyla guen-
therii Keferstein, 1868].

Liijla Cope, 1870, Proc. Amer. Philos. Soc, 11:
160 [Emendation of Leiyla Keferstein,
1868].

himnophys Jimenez de la Espada, 1870, J. Sci.
Math., Phys. Nat. Lisboa, 3:59 [Type-
species by subsequent designation ( Myers,
1962), himnophys cornutus Jimenez de la
Espada, 1870].

Pristimantis Jimenez de la Espada, 1870, Ibid.,
3:61 [Type-species by monotypy, Pristi-
mantis galdi Jimenez de la Espada, 1870].

Cyclocephalus Jimenez de la Espada, 1875,
Vert. Viaje Pacif., Batr., pi. 3 [Type-species
by monotypy, Cyclocephalus lacrimosus Ji-
menez de la Espada, 1875].

Hypodictyon Cope, 1885, Proc. Amer. Philos.
Soc, 22:383 [Type-species by original des-
ignation, Phyllobates ridens Cope, 1866].

Liohyla Cope, 1894, Ibid., 31:335 [Emendation
of Leiyla Keferstein, 1868].

Basanitia Miranda-Ribeiro, 1923, Rev. Mus.
Paulista, 13:851 [Type-species by mono-
typy, Basanitia lactea Miranda-Ribeiro,
1923].

Noblella Barbour, 1930, Zoologica, 11:81
[Type-species by original designation,
Sminthillus peruvianus Noble, 1921].

Phrynanodus Ahl, 1933, Zool. Anz., 104:29
[Type-species by monotypy, Phrynanodus
nanus Ahl, 1933].

Teletrema Miranda-Ribeiro, 1937, O Campo
(Rio de Janeiro), 8 (89): 67 [Type-species
by monotypy, Teletrema heterodactylum
Miranda-Ribeiro, 1937].

Microbatrachylus Taylor, 1940, Univ. Kansas
Sci. Bull., 26:499 [Type-species by original
designation, Eleutherodactylus hobartsmithi
Taylor, 1936].

Ctenocranius Melin, 1941, Medd. Goteborgs
Mus. Zool. Avd., 88:49 [Type-species by
subsequent designation (Myers, 1962),
Lirmwphys cornutus Jimenez de la Espada,
1870].

Pseudohyla Andersson, 1945, Ark. Zool., 37A:
86 [Type-species by monotypy, Pseudohyla
nigrogrisea Andersson, 1945].

Trachyphrynus Coin and Cochran, 1963, Proc.
California Acad. Sci., 31:502 [Type-species
by original designation, Trachyphrynus
myersi Coin and Cochran, 1963].

Diagnostic definitioti. — 7) omoster-
num usually present, small, medium-
sized, or large, long and narrow or rela-

FiGURE 95. Lateral and dorsal views of skull of
Eleutherodactylus sulcatus (KU 100355, X 2.4).

tively broad, absent in at least one spe-
cies— E. ruthae; 9) maxillary arch
toothed; 10) alary processes of premaxil-
lae relatively broad at base, direction
variable — dorsal, dorsolateral, or pos-
terodorsal; 11) palatal shelf of premaxil-

FiGURE 96. Lateral and dorsal views of skull of
Eleutherodactylus diastema (KU 68263, X 6).

146

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

la deep, usually deeply dissected; 12) fa-
cial lobe of maxilla deep, usually not
exostosed; 13) palatal shelf of maxilla
broad, usually with prominent pterygoid
process; 15) nasals large, in broad me-
dian contact, narrowly separated in some
species; 16) nasals not in contact with
pterygoids, sometimes in contact with
maxillae; 18) frontoparietal fontanelle
absent in adults except in E. pahneri and
E. lohymperi; 19) frontoparietals not
ornamented in most species groups, but
bearing lateral crests in biporcattis, cor-
mitus, galdi, and iinlstrigatus complexes;
20) frontoparietals fused with prootics
or not. The bones are fused in most
species in the West Indies and northern
Andes, free in other groups; 22) epiotic
eminences prominent to obsolete — group
variable; 23) cristae paroticae short and
stocky to long and narrow; 24) zygo-
matic ramus of squamosal short to long,
sometimes knobbed, in contact with
maxilla in one species (rutliae); 25) otic
ramus of squamosal short to long, usual-
ly forming a small otic plate, orna-
mented in a few species groups — notably
the galdi group; 26) squamosal-maxil-
lary angle 44-67°, most 50-60°; 27) colu-
mella present; 28) prevomers nearly al-

ways toothed, entire, narrowly separated
medially to broadly separated medially;
29) palatines long, usually expanded lat-
erally, relatively widely separated me-
dially, no odontoid ridges; 30) spheneth-
moid entire, extending anteriorly be-
neath nasals variable distance; 31) an-
terior ramus of parasphenoid narrow to
broad, relatively long, nearly reaching
prevomers, not keeled medially; 32)
parasphenoid alae in two patterns: 1.
alae deflected posteriorly, short, not
overlapped laterally by median rami of
pterygoids, and 2. alae oriented at right
angles to anterior ramus, rarely deflected
posteriorly, long, broadly overlapped by
median rami of pterygoids. The first pat-
tern is seen in West Indian species and
some Andean species, whereas the sec-
ond is seen in Central American and
lowland South American species; 33)
pterygoids slender to relatively massive,
lacking ventral flange, anterior rami
relatively short, not reaching palatines,
median rami short to long, straight or
bent; 34) occipital condyles relatively
small, stalked or not, widely separated
medially; 36) terminal phalanges nearly
always clearly T-shaped, inner phalan-
ges usually knobbed, terminal phalanges

Figure 97. Dorsal views of the skulls of Eleutherodactylus conspicillattis (KU 108988, X 4.5) and E.
planirostris (KU 92656, X 4.5). Arrows point to prevomerine teeth which are visible in dorsal view.

LYNCH: LEPTODACTYLOID FROGS

147

Figure 98. Lateral views of skulls of Eletithew-

dacitihis. (A) platuwstm (KU 92656, X 6), (B)

^aldi (USNM field GOV 8944, X 6), and (C)

ruthae ( AS 4237, X 2).

of toes more T-shaped than those of
fingers; presence of T-shaped terminal
phalanx is expressed externally by the
presence of a terminal transverse groove
across the tip of the digital pad; 37)
alary processes of hyoid plate on narrow
stalks; 40) m. depressor mandibukie in
one slip in E. galdi, in one slip with sep-
aration into two slips dorsally in species
of hiporcatus and cornutus groups, or in
two large slips in most species; 42) males
with single subgular vocal sac or none,
internal or external; 43) glands on body
usually absent, those present are usually
loosely organized inguinal glands; 44)
tongue long and narrow to large and
round, posterior edge usually free; 45)
toes free of webbing to nearly fully
webbed; when webbing is present, it is
indicative of a close association with
streams, webbing is greatest in anoma-
lus, karhchmidti, punctariohis, and rani-
formis, although several species of the
rugulosiis group in Central America have
the toes one-half webbed; digits usually
bear large pads; 49) adults range from
12-100-|- mm. SVL; 50) tympanum ab-

sent in anotis, concealed in many species,
visible externally in most species.

Composition. — Gorham (1966) listed
nearly 300 species in the most recent
compilation of names in the genus. Al-
bert Schwartz informed me that there
are 100 species in the West Indies (in-
cluding Trinidad). My own estimate is
that the genus contains about 400 spe-
cies, many yet unnamed.

Distribution. — Sinaloa and Tamau-
lipas, Mexico (but not on the Mexican
Plateau) southward and eastward
throughout Middle America to northern
Argentina and southern Brasil; all West
Indian islands; introduced into Florida.

Remarks. — Myers (1962) discussed
the generic synonymy of this genus and
included most of the generic synonyms
listed above as well as Sijrrhophus and
Lithodytes in the synonymy of Eleti-

FiGURE 99. Dorsal views of skulls of (top)

Eleiithewdactyhts btifoniformis ( KU 80621, X

2) and E. palmcri (KU 110923, X 3).

148

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

thewdactijhis. Microhatmchijlus was
synonymized by Lynch (1965), and he
later (1968c, 1968d) considered Bamni-
tia, Phnjnanodus; and Traclujphnjmis
inseparable from Eleutlierodactijlus.

Two generic synonyms are added at
this time: NohJello Barbour and Pseu-
dohyla Andersson. Noble (1921) named
Smint]}iUiis penwiami.s on the basis of
several specimens of a minute frog with
an anterior epicoracoidal fusion (the
distinguishing character of the genus).
At the time of the description of peru-
vianus, the genus SminthiUus was known
only from Cuba, but with the description
of a species from southeastern Brasil by
Parker (1926), it began to appear that
SminthiUus was a widespread, Neotropi-
cal genus of small frogs. Noble ( 1926b,
1931) suggested that Sminthilhis was de-
rived from Eleiitherodactyliis or Sijrrho-
phus but placed it in another family
( Brachycephalidae ) . Griffiths ( 1959 )
placed each of the three species of
Sminthilhis in a separate genus, utilizing
Barbour's (1930) NohleUa ior peruviana.
Barbour (1930) proposed Nohlella for
peruviana because the Cuban species
was geographically remote from the two
South American species and because he
believed SmintluUus to be a PJujUobates
(Dendrobatidae). Griffiths offered a
new generic name, Euparkerelh, for the
Brasilian species. Etiparkerella, while
here retained as an eleutherodactyline,
is very distinct from all other genera of
the tribe and subfamily. SminthiUus is
retained as a genus of eleutherodacty-
lines on less secure grounds — the maxil-
lary teeth are absent. NohleUa peruvi-
ana, on the other hand, has maxillary
teeth, although it apparently has no pre-
vomers. The single specimen of this
species available to me cannot be gener-
ically separated from EJeutherodactylus;
an epicoracoidal bridge is not, in my
opinion, sufficient basis for the mainte-
nance of an otherwise undefinable ge-
neric group ( see discussion on page 59 ) .
NohleUa therefore is placed in the syn-
onymy of Eleutherodactylus. This action

creates one minor problem — Noble's
peruvianus was proposed in 1921, but
Melin ( 1941 ) proposed a Hylodes peru-
vianus which becomes a secondary hom-
onym of Noble's name. Rather than
propose a replacement name for a prob-
ably invalid species, I suggest that Me-
lin's name be kept in mind by the person
who eventually studies the conspicillatus
group in the Amazon Basin of Peru.

Andersson (1945) proposed Pseudo-
hyla as a hylid genus, but having studied
the holotype of P. nigrogrisea, I do not
consider the genus separable from Eleu-
therodactylus (Lynch, 1969a). Eleu-
therodactylus nigrogriseus is a small spe-
cies of the genus and has been found in
the valley of the Rio Pastaza and on the
slopes of the Cordillera Due in eastern
Ecuador.

The genus Eleutherodactylus is a
large and unwieldy one, although Hyla
and possibly Rana are more unwieldy at
present. In contrast to the latter two
genera (and perhaps to the opinions of
several herpetologists) the genus Eleu-
therodactylus is marked by considerable
homogeneity. There is a large range of
sizes among the species; most species
lack webbing between the toes but some
have it, including one species with fully
webbed feet (E. karlschmidti). Exostosis
of the cranial bones is developed in sev-
eral species groups but in general the
phenomenon is an uncommon trait in the
genus. Some species have minute digi-
tal pads and only small lateral projec-
tions of the terminal phalanges on the
hands, but all species have moderate to
large T-shaped terminal phalanges on
the toes (see pp. 65-66 for further dis-
cussion ) .

Three characters exhibit especially
interesting variation in Eleutherodacty-
lus— 1) fusion of the frontoparietal and
otoccipital bones; 2) degree of overlap
of the parasphenoid alae and median
rami of the pterygoids; and 3) median
separation of the prevomers. On the
basis of the variation in these characters,
the species of the West Indies and parts

LYNCH: LEPTODACTYLOID FROGS

149

of the Andean system form one group
and the species of Mexico, Central
America, and lowland South America
form a second group. These characters
are discussed below.

Baldauf and Tanzer ( 1965 ) im-
proved our knowledge of leptodactylid
skulls with the description of the cra-
nium of SyrrJiopJms maniockii. In this
work they pointed out the fusion of the
frontoparietal and prootic in this species.
These bones are fused in all species of
Syrrlioplnis and Tomodactyltis, whereas
they are not fused in the Eleutlwrodoc-
tylus of Mexico and Middle America.
Although fewer than one-third of the
species of Eleutherodactylus have been
studied for this character, I feel that I
have checked a sufficiently representa-
tive sample in that I have examined spe-
cies from all parts of the range of the
genus. Fusion of the frontoparietal and
prootic ( otoccipital, since the prootic is
usually fused with the exoccipital) oc-
curs in the species of the genus found in
the West Indies from Bermuda and the
Bahamas (and Florida) to Trinidad.
The species of the Hispaniolan inoptatiis
group {inoptatiis and nifhae examined)
as well as the Puerto Rican karhchmidti
apparently have the two bones free. At
least some (perhaps all) of the species
of the Andean groups exhibit fusion of
these bones. No species was examined
from the Andes south of Ecuador, All
species of the genera Syrrhophus and
Tomodoctyhis as well as SmintJiiUiis ex-
hibit the fusion. No species of Eleu-
therodactylus in Mexico or Central
America normally exhibits frontoparietal-
prootic fusion (see below), nor do spe-
cies of the genus found in Chocoan Co-
lombia and Ecuador. Insofar as I am
aware, no species found in the Amazon
Basin exhibits the fusion nor do the rep-
resentatives of the genus in southeastern
Brasil.

Some variability was noted. One
specimen of E. fitzingeri (JDL S-407)
exhibited fusion of the frontoparietal
and otoccioital bones whereas the other

nine specimens examined did not. One
of three specimens of E. cldoronotus
examined exhibited the fusion. Two of
the 19 specimens of E. curtipes examined
did not exhibit fusion of these bones; both
were small specimens, suggesting that the
fusion is an ontogenetic phenomenon.
In the cases of the first two species, I
regard the fusion as aberrant. In each
instance, the frontoparietals, nasals, pre-
vomers, and parasphenoid were bound
to the sphenethmoid and otoccipitals
with no apparent sutures. This suggests
that the fusion in these cases resulted
from extensive calcification rather than
osteological fusion.

The degree of overlap between the
median rami of the pterygoids and para-
sphenoid alae follows the same pattern
as the frontoparietal-prootic fusion with
some departure. The median pterygoid
rami of the rhodopis group of Eleuthero-
dactylus are short and bent so that there
is no actual contact between the ptery-
goid and parasphenoid, but the median
ramus of the pterygoid abuts against the
otic capsule just above the parasphenoid
ala. In the majority of species of the
genus, the median ramus is broadly in
contact with the anterior edge of the
parasphenoid ala or the median ramus is
shortened and may or may not reach the
otic capsule and does not reach the
shortened parasphenoid ala. In those
species with a pterygoid-parasphenoid
overlap, the frontoparietals and prootic
bones are free. The members of the
rhodopis complex of Eleutherodactylus
have the bent pterygoid and hence do
not have a typical pterygoid-parasphe-
noid overlap but because the median
rami of the pterygoids are proportion-
ately long, these species are included in
the same complex as those species with
a broad overlap of the parasphenoid alae
and median pterygoid rami. In the
rhodopis complex the frontoparietal and
prootic bones are not fused. Those spe-
cies with a very short median ramus on
the pterygoid and no pterygoid-para-
sphenoid overlap also exhibit the fused

150

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

frontoparietal-prootic. For purposes of
further discussion, these two major
groups are here termed the Alpha and
Beta groups of Eleutherodactylus. Mem-
bers of the Alpha group (fused fronto-
parietal-prootic, non-overlap between
pteiygoids and parasphenoid alae)
usually have relatively widely separated
prevomers, whereas the Beta group
frogs (frontoparietal and prootic not
fused, pterygoids overlap parasphenoid
alae ) usually have the prevomers in con-
tact or only narrowly separated.

I submit that the two divisions, Alpha
and Beta, are natural divisions within
the genus Eleutherodactylus and not
simply a chaotic array of species ex-
hibiting two osteological patterns. I bor-
rowed the terminology of Etheridge
( 1960 ) for the major divisions since at
this time I am not willing to afford the
two divisions nomenclatorial recognition
and prefer the informal divisional names.
This course is taken because only a rela-
tively small part of the genus has been
surveyed, and many species could not
be assigned to subgenus were nomen-
clatorial assignments made. If the two
divisions were accorded nomenclatorial
status, the Alpha group would be the
subgenus Eleiithewdactyhis and the Beta
group would take the name Cratigastor.

Several osteological features seem to
lend themselves well to the possible use
of skeletal morphology in the assessment
of species group relationships within the
genus Eleutherodactylus. The degree of
median separation of the prevomers has
potential in that it varies concordantly
with several other osteological traits ( the
development of cranial crests, shape of
the rami of the squamosal, and size and
shape of the nasal bones). Among the
West Indian species of the genus, the spe-
cies groups have long been based at least
in part on the length of the prevomerine
dentigerous processes, and I would ex-
pect this character complex to be of at
least some value, although its use is
greatly hampered by the occasional loss
of dentigerous processes in several

groups of the genus. I have not attempted
to divide the species of the genus into spe-
cies groups, because I have not studied all
species of the genus and I cannot rely
upon the literature for many characters
that I regard as of potential importance.
A study of the osteology of the genus
Eleutherodactylus is envisioned, but
prior to its initiation considerable re-
search must be done in straightening out
many nomenclatorial entanglements, the
description of many more species, and
synonymizing of many names.

Euparkerella Griffiths, 1959
(Fig. 100)

Euparkerella Griffiths, 1959, Proc. Zool. Sue.
London, 132:477 [Type-species by original
designation, Sminthillus brasiliensis Parker,
1926].

Diapiostic definition. — 7) omoster-
num present, small; 9) maxillary arch
toothed; 10) alary processes of premaxil-
lae directed posterodorsally, broad at
base; 11) palatal shelf of premaxilla very
broad, not notched, palatal process mi-
nute; 12) facial lobe of maxilla shallow;
13) palatal shelf of maxilla broad, taper-
ing posteriorly, no pterygoid process;
15) nasals small, moderate median sep-
aration; 16) nasals in contact with maxil-
lae, separated from pterygoids; 19) fron-
toparietals not ornamented; 20) fronto- i
parietal fused to prootic; 22) epiotic I
eminences small; 23) cristae paroticae
very broad, stocky; 24) zygomatic ramus
of squamosal of moderate length, widely
separated from maxilla; 25) otic ramus
of squamosal elongate, not expanded
medially into otic plate; 26) squamosal-
maxillary angle about 60°; 27) columel-
la absent; 28) prevomers reduced to
minute slivers, widely separated medial-
ly, edentate, dentigerous rami lost; 29)
palatines very slender, reduced in size,
widely separated medially; 30) sphen-
ethmoid divided, extending anteriorly
under posterior edge of nasals; 31) an-
terior ramus of parasphenoid very broad,
not keeled medially; 32) parasphenoid
alae oriented at right angles to anterior

LYNCH: LEPTODACTYLOID FROGS

151

Figure 100. Lateral, dorsal, and ventral views of skull of Euparkeivlla brasiiieims (KU 93192,

X5).

ramus, short, not overlapped by median
rami of pterygoids; 33) pterygoids rela-
tively small, anterior rami long, not
reaching palatines; 34) occipital con-
dyles small, stalked, widely separated
medially; 36) terminal phalanges short
and broad, bearing small hook-like lat-
eral processes; 37) alary processes lack-
ing on hyoid plate; 40) in. depressor
mandihtilae in two slips; 42) males with
median subgular vocal sac; 43) body
lacking glands; 44) tongue large, not
notched, posterior edge free; 45) toes
lacking webbing, digital tips pointed, not
dilated, fingers and toes short; 46-48);

49) adults small, less than 20 mm. SVL;

50) tympanum absent.
Composition. — Monotypic.
Distribution. — Known only from the

Serra dos Orgaos, state of Guanabara,
Brasil.

Remarks. — Parker (1926) named
Sminthillus hrasiliensis on the basis of

the "juvenile" cotypes of Boulenger's
Oocormus microps (=zZachaenus parvu-
lits). Noble examined Parker's drawings
of the pectoral girdle of hrasiliensis and
agreed with Parker that the species fit
the characteristics of Sminthillus. At
that time, SmintluUiis comprised tliree
species — one Cuban, one Peruvian, and
one Brasilian. Griffiths (1959) argued
that if all three species were independent
derivatives of Eleutherodactijlus, then
each belongs to a separate genus. His
solution was to place each in a mono-
typic genus. Griffiths proposed Etipark-
erelJa for the Brasilian species but did
not provide diagnostic statements for the
genus. Euparkerella is very distinctive
in its osteology. In external morphology,
Euparkerella is superficially very similar
to Zachaenus parvulus. These two spe-
cies live in the same habitat ( leaf litter )
in southeastern Brasil and are frequently
collected syntopically. The coloration of

152

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

the two species is nearly identical. Eu-
parkerella brasiliensis and Zachaenus
parvuJus differ in adult size and the
length of the fingers ( Fig. 94 ) .

The skeleton of Eiiparkerelh does
not bear any close resemblance to that
of any other leptodactylid genus, al-
though the squamosal architecture, lack
of columellae, and shape of the hyoid
plate of Euparkerella suggest a relation-
ship to the genus Holoaden. The termi-
nal phalanges of Euparkerella are unique
in the shape of the lateral expansions
(Figs. 43-44). The digits are not pad-
like and lack the terminal transverse
groove that is found on the digital tips of
Eleiitherodactyhis, Sijrrhophus, Smin-
thiUus, and Tomodactylus. The fronto-
parietal and prootic fusion of Euparker-
ella is suggestive of a relationship with
the Alpha division of Eleutherodactylus.
At present, I regard the relationships of
Euparkerella as obscure but feel that the
genus is probably more closely related to
Holoaden than to either division of Eleu-
therodactylus.

Holoaden Miranda-Ribeiro, 1920

(Fig. 101)

Holoaden Miranda-Ribeiro, 1920, Rev. Mus.
Paulista, 12:319 [Type-species by mono-
typy, Holoaden hiederwaldti Miranda-
Ribeiro, 1920].

Diagnostic definition. — 7) omoster-
num moderate-sized; 9) maxillary arch
toothed; 10) alary processes of premaxil-
lae directed dorsally, moderately wide at
base; 11) palatal shelf of premaxilla
broad, slightly indented; 12) facial lobe
of maxilla relatively shallow, not ex-
ostosed; 13) palatal shelf of maxilla of
moderate width, no distinct pterygoid
process; 15) nasals moderate-sized, nar-
rowly separated medially; 16) nasals
separated from maxillae and pterygoids;
18) frontoparietal fontanelle moderate-
sized; 19) frontoparietals not orna-
mented; 20) frontoparietals not fused
with prootics; 22) epiotic eminences
small; 23) cristae paroticae short, stocky;
24 ) zygomatic ramus of squamosal mod-
erately long, widely separated from
maxilla; 25) otic ramus of squamosal

Figum: 101. Lateral (X 8), dorsal (X 4) and ventral (x 4) views of skull of Holoaden biadei

(KU 107087).

LYNCH: LEPTODACTYLOID FROGS

153

long, not expanded medially into otic
plate; 26) squamosal-maxillary angle
about 65°; 27) columella absent; 28)
prevomers entire, toothed, separated me-
dially; 29) palatines narrow, elongate,
separated medially; 30) sphenethmoid
entire, extending anteriorly beneath pos-
terior edge of nasals; 31) anterior ramus
of parasphenoid moderately broad, not
keeled medially; 32) parasphenoid alae
oriented at right angles to anterior
ramus, widely separated from median
rami of pterygoids; 33) pterygoids slen-
der, anterior rami long, nearly reaching
palatines; 34) occipital condyles small,
not stalked, widely separated medially;
36) terminal phalanges knobbed; 37)
hyoid plate lacking alary processes; 40)
m. depressor mandihulae in two slips;
42) males lacking vocal sac; 43) entire
skin glandular forming large, indefinite
parotoid, Hank, and inguinal glands and
a large glandular mass on the thighs;
44) tongue oval, not notched, posterior
one-half free; 45) toes free of webbing,
digital tips narrow; 49) adults small, 20-
48 mm. SVL; 50) tympanum absent.

Composition. — Two species are pres-
ently known (hradei and Itiederwoldti).

Distribution. — The coastal Serras of
southeastern Brasil.

Remarks. — Miranda-Ribeiro ( 1920,
1926) included Holoaden in the Telma-
tobiidae with Acris, Iliodlscus, and Tel-
matohius. He considered the Telmato-
biidae to be intermediate between the
Hylidae and Leptodactylidae. Lutz
(1958) considered Holoaden a member
of a generic cline (Cijcloramphus-Cras-
pedoglossa-HoIoaden) but did not place
the genus in the Cycloramphinae. Holo-
aden is superficially similar to Zachaenus
stejnegeri (Craspedoglossa auctorum) but
differs in several osteological characters.
Holoaden and Eiiparkerella lack alary
processes on the hyoid plate and differ
from all other leptodactylids except
Limnomedusa and Sminthillus in this
character. Holoaden is osteologically
similar to the Andean Niceforonia. This
similarity may reflect a faunal relation-

ship between the Brasilian highlands
and the Andes or may reflect conver-
gence in the arrangement of the skull
bones resulting from adaptation to bur-
rowing. In osteological and external
characters, Holoaden does not seem
especially closely related to any other
genus included in the Eleutherodactylini
except Eiiparkerella.

Hylactopliryne Lynch, 1968
(Figs. 102-03)

Hylactophrijne Lynch, 1968, Univ. Kansas
Pubis. Mus. Nat. Hist., 17:511 [Type-
species jjy original designation, HyJodes
august i Duges, 1879].

Diagnostic definition. — 7) omoster-
num large; 9) maxillary arch toothed;
10) alary processes of premaxillae di-
rected posterodorsally, moderately wide
at base; 11) palatal shelf of premaxilla
broad, deeply dissected; 12) facial lobe
of maxilla relatively deep, not exostosed;
13) palatal shelf of maxilla of moderate
width, pterygoid process large; 15) na-
sals very large, in broad median contact;
16) nasals in contact with maxillae, not
with pterygoids; 19) frontoparietals not
ornamented; 20) frontoparietals not
fused to prootics; 22) epiotic eminences
large; 23) cristae paroticae long, narrow;

24) zygomatic ramus of squamosal of
moderate length, not reaching maxilla;

25) otic ramus of squamosal long, as
long as zygomatic ramus, expanded me-
dially into small otic plate; 26) squa-
mosal-maxillary angle about 55°; 27)
columella present; 28) prevomers large,
entire, narrowly separated medially; 29)
palatines large, broad, separated medial-
ly; 30) sphenethmoid entire, large, ex-
tending anteriorly beneath posterior
edge of nasals; 31) anterior ramus of
parasphenoid broad, not keeled; 32)
parasphenoid alae oriented at right
angles to anterior ramus, broadly over-
lapped by median rami of pterygoids;
33) pterygoids moderate-sized, anterior
rami long, reaching palatines; 34) oc-
cipital condyles moderate-sized, not
stalked, widely separated medially; 36)

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 102. Dorsal views of skulls of Ilylactophryue aiigiisti (KU 56187, X 2.3, left) and hchnocnema

quixensis (KU 104,388, X 4.7, right).

terminal phalanges knobbed; 37) alary
processes of hyoid plate on narrow stalks;
40) ill. depressor matidibuhe in two
slips; 42) males with median subgular
vocal sac; 43) body free of glands; 44)
tongue large, oval, posterior edge free;
45) toes free of webbing or lateral
fringes, digital tips narrow, first finger
longer than second; 49) males 37-77, fe-

males 40-95 mm. SVL; 50) tympanum
visible externally.

Composition. — Two species are cur-
rently recognized (augusti and torahti-
maraeyisis). The fonuer is composed of
four subspecies. The group was revised
by Zweifel (1956b).

Distribution. — Mexican Plateau from
Arizona, New Mexico, and Texas to cen-

FiGURE 103. Lateral and ventral views of skulls of Hylactophnjne augusti (KU 56187, X 3, left) and
hchnocnema quixensis (KU 104388, X 3 and X 4.7, right).

LYNCH: LEPTODACTYLOID FROGS

155

tral Mexico (Cordillera Volcanica and
western Sierra Madre del Sur). An iso-
lated population is known from the
mountains in the Isthmus of Tehuan-
tepec.

Remarks. — Hylactopliryne is very dis-
tinctive when compared with the Central
American and Mexican leptodactylids
but is less distinctive when compared
with some of the South American eleu-
therodactylines. When I named Hijlac-
tophryne (Lynch, 1968a), I suggested
that the genus was allied to Eupsoplms.
At that time I was under the mistaken
impression that Oreohaies qiiixensls was
an Eupsophus. Subsequently, I have
examined all of the species of frogs re-
ferred to the genus Eupsophus by vari-
ous authors and concluded that the
genus Eupsophus in the broad sense
(that of Noble, 1931, and Gorham, 1966)
includes representatives of five genera.
One of these genera is Ischnocnema (see
following account), which contains two
species in South America. Hyhcto-
phryne and Ischnocnema are very simi-
lar. The skulls of the two genera differ
in proportions but are otherwise the
same (Figs. 102-03). These genera are
tentatively separated on the basis of the
presence (HyJactophryne) or absence
(Ischnocnema) of a discoidal fold. The
two genera may prove to be independent
derivatives of Eleutherodacfylus rather
than primitive as I previously suggested
(Lynch, 1968a). The Eleutherodactyhis
guentheri group of frogs bear consider-
able resemblance to Ischnocnema quix-
ensis and /. verrucosa. Eleutherodacty-
his carrioni of the southern Ecuadorian
Andes is very similar to HyJactophryne
augiisti. In both cases, the species of
EleutJierodactyhis differ from the two
genera in having T-shaped terminal pha-
langes (rather than knobbed phalanges)
and in having the terminal transverse
groove on the digital pad (rather than
no groove or pad ) . For the present, I do
not advocate combining HyJactopliryne
and Ischnocnema with one another or
with Eleutlierodactylus.

Ischnocnema Reinhardt and Liitken,
1862

(Figs. 102-03)

Ischnocnema Reinhardt and Liitken, 1862, Vid.
Meddel. Natnrh., Foren., 1861:239 [Type-
species by monotypy, Leiupenis verrucosus
Reinhardt and Liitken, 1862].

Oreohates Jimenez de la Espada, 1872, An. Soc.
Espaiiola Hist. Nat., 1:87 [Type-species by
monotypy, Oreohates cfuixcnsis Jimenez de
la Espada, 1872].

Diagnostic definition. — 7 ) omoster-
num moderate-sized; 9) maxillary arch
toothed; 10) alary processes of premaxil-
lae directed dorsally, moderately wide at
base; 11) palatal shelf of premaxilla rela-
tively deep, notched; 12) facial lobe of
maxilla deep, not exostosed; 13) palatal
shelf of maxilla relatively narrow, ptery-
goid process large; 15) nasals very large,
in broad median contact; 16) nasals
separated from maxillae and pterygoids;
19) frontoparietals only sHghtly orna-
mented; 20) frontoparietals not fused to
prootics; 22) epiotic eminences poorly
defined; 23) cristae paroticae relatively
long and narrow; 24) zygomatic ramus
of squamosal relatively long, widely sep-
arated from maxilla; 25) otic ramus of
squamosal slightly shorter than zygo-
matic ramus, expanded medially into
small otic plate; 26) squamosal-maxillary
angle about 50°; 27) columella present;
28) prevomers large, entire, toothed,
narrowly separated medially; 29) pala-
tines large, broad, separated medially,
bearing odontoid ridge in quixerisis; 30)
sphenethmoid entire, extending anterior-
ly beneath posterior edge of nasals; 31)
anterior ramus of parasphenoid relative-
ly narrow, not keeled medially; 32) para-
sphenoid alae oriented at right angles to
anterior ramus, slightly overlapped lat-
erally by median rami of pterygoids; 33)
pterygoids moderate-sized, anterior rami
long, reaching palatines; 34) occipital
condyles small, on small stalks, widely
separated medially; 36) terminal pha-
langes knobbed; 37) alary processes of
hyoid plate on narrow stalks; 40) m. de-
pressor mandiJ)ukie in two slips; 42)
males with median subgular vocal sac;

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

43) body lacking glands; 44) tongue
large, oval, posterior edge free; 45) toes
lacking webbing, digital tips narrow, first
finger longer than second; 46-48); 49)
adults to 55 mm. SVL; 50) tympanum
visible externally.

Composition.— Two species are pres-
ently known (quixensis and verrucosa).
Distribution. — Western edge of the
Amazon basin in Ecuador, northeastern
Peru, and adjacent Brasil (quixensis),
and in the coastal Serras of southeastern
Brasil (verruscosa). Both species are
found in forested habitats.

Remarks. — As mentioned before (Hy-
lactophryne account), Ischnocnema is
very similar to Hylactophryne; the two
genera are here separated on the basis of
trivial external characters, geography,
and a lack of knowledge concerning
breeding behavior and biology. The
similarities in morphology between these
two geographically isolated groups of
eleutherodactylinc frogs is suggestive of
an independent origin of each from an
EleutherodactyJus stock through a de-
parture from the arboreal adaptive zone.
Typical Eleutherodactylus have toe pads
(and are frequently mistaken for hylids
by the uninitiated herpetologist ) and
are usually semi-arboreal or arboreal in
habits. Both Hylactophryne and Isch-
nocnema are terrestrial frogs; the former
lives in arid, non-forested regions and
the latter lives in moist, forested en-
vironments.

Hylactophryne and Ischnocnema may
represent relicts of a formerly wide-
spread eleutherodactylinc stock from
which more successful genera (Eleu-
therodactylus) evolved. At present, too
little is known of the osteology of Eleu-
therodactylus to determine the evolu-
tionary directions.

Niceforonia Coin and Cochran, 1963

(Fig. 104)

Niceforonia Coin and Cochran, 1963, Proc.
California Acad. Sci., 31:499 [Type-species
by original designation, Niceforonia nana
Coin and Cochran, 1963].

Diagnostic definition. — 7) omoster-
num moderate-sized; 9) maxillary arch
toothed; 10) alary processes of premaxil-
lae directed dorsally, relatively broad at
base; 11) palatal shelf of premaxilla
broad, deeply notched; 12) facial lobe
of maxilla deep anteriorly, tapering pos-
teriorly; 13) palatal shelf of maxilla
broad, pterygoid process moderate-sized;

15 ) nasals small, narrowly separated me-
dially; nasals in contact in fiavomacidata;

16) nasals in contact with maxillae, not
with pterygoids; 19) frontoparietals not
ornamented, except in flavomaculata; 20)
frontoparietals not fused with prootics;
22) epiotic eminences obsolete; 23) cris-
tae paroticae broad, stocky; 24) zygo-
matic ramus of squamosal of moderate
length, widely separated from maxilla;

25) otic ramus of squamosal long, ex-
panded medially into small otic plate;

26) squamosal-maxillary angle about
55°; 27) columella present in most spe-
cies, absent in montia, probably absent
in simon.sii; 28) prevomers toothed or
not, entire, relatively large, separated
medially; 29) palatines slender, sep-
arated medially; 30) sphenethmoid en-
tire, extending anteriorly beyond an-
terior edge of nasals; 31 ) anterior ramus
of parasphenoid broad, long, not keeled
medially; 32) parasphenoid alae oriented
at right angles to anterior ramus, short,
not overlapped laterally by median rami
of pterygoids; 33) pterygoids small, me-
dian rami short, anterior rami relatively
long, not reaching palatines; 34) occipi-
tal condyles relatively small, stalked,
widely separated medially; 36) terminal
phalanges knobbed; 37) alary processes
on hyoid plate on narrow stalks; 40) m.
depressor mamlihulue in two slips; 42)
males with large median subgular vocal
sac; 43) body lacking glands; 44) tongue
large, round, free at posterior edge; 45)
toes lacking webbing and lateral fringes,
digital tips narrow; 46-48); 49) adults
small, less than 30 mm. SVL; 50) tympa-
num visible externally, concealed, absent
(montia), or possibly absent (simonsii).

Composition.— Lynch (1968d) in-

LYNCH: LEPTODACTYLOID FROGS

157

Figure 104. Lateral, dorsal, and ventral views of skull of Niceforonia montia (MCZ 24352, X 5).

eluded N. nana, N. festae, and N. montia
as definite members of the genus and
referred N. colinnbiana and N. simonsii
to the genus. Based on examination of
paratypes of Eupsophiis tvettsteini by
stereo-radiographs, I now include that
species in Niceforonia. EleutJierodacty-
lus flavomaculattis Parker is a Nicefo-
ronia (Lynch, 1969b) and Cei's (1968b)
Syrrlwplnis hplacai is probably a mem-
ber of Niceforonia as well.

Distribution. — High elevations in the
Andes of Colombia, Ecuador, Peru, and
Bolivia.

Remarks. — Coin and Cochran (1963)
considered Niceforonia to be most
closely allied to Eupsophus. In external
characters, Eupsophus and Niceforonia
cannot be separated. The cervical coty-
lar arrangement and the median separa-
tion of the occipital condyles suggests
that Niceforonia is more closely related
to the Eleutherodactylini than to the
Alsodini. The breeding biology of Nice-
foronia is unknown except that N. flavo-
maculata lays large terrestrial eggs, but

the large eggs are suggestive of direct
development in all of the species of the
genus. The very distinctive spheneth-
moid of Niceforonia is duplicated by at
least one species of Eleutlierodactijlus
(bogotensis). Niceforonia is separated
from Eleutlierodactyhis because the
digital tips are narrow, there is no ter-
minal transverse groove on the digital
tips, and the terminal phalanges are
knobbed. The slight median separation
of the nasal bones in Niceforonia occurs
in several groups of Eleutlierodactylus,
although the trait is uncommon in Eleu-
tlierodactylus. The other eleuthero-
dactylines with knobbed terminal pha-
langes (Amblyphrynus, Holoaden, Hy-
lactop]}ryne, and Ischnocnema) are dis-
tinctive when compared with Nicefo-
ronia, although Holoaden resembles
Niceforonia in the arrangement of the
cranial bones. This osteological simi-
larity possibly reflects convergence in
view of the dissimilar morphology of the
hyoid plates of these two genera.

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Sminthillus Barbour and Noble, 1920

(Fig. 105)

SminthiUus Barbour and Noble, 1920, Bull.
Mus. Conip. Zool., 63:402 [Type-species liy
original designation, Plii/Uobates limhatus
Cope, 1862].

Diagnostic dejinition. — 7) omoster-
num small, elongate; 9) maxillary arch
edentate; 10) alary processes of pre-
maxillae directed dorsolaterally, short,
broad at base; 11) palatal shelf broad,
deeply dissected; 12) facial lobe of max-
illa shallow; 13) palatal shelf of maxilla
of moderate width, no pterygoid process;
15) nasals small, narrowly separated me-
dially; 16) nasals in tenuous contact
with maxillae, separated from pteiygoids;
19) frontoparietals not ornamented; 20)
frontoparietals fused to prootics; 22)
epiotic eminences present, small; 23)
cristae paroticae very broad, stocky; 24)
zygomatic ramus of squamosal very
short, knobbed; 25) otic ramus of squa-
mosal very long, no otic plate; 26) squa-

mosal-maxillary angle about 60°; 27)
columella present; 28) prevomers mi-
nute, greatly reduced in size, entire,
widely separated medially; 29) palatines
slender, widely separated medially, lat-
eral to prevomers; 30) sphenethmoid en-
tire, extending anteriorly to middle of
nasals; 31) anterior ramus of parasphe-
noid narrow, not keeled medially; 32)
parasphenoid alae slightly deflected pos-
teriorly, very short, not overlapped lat-
erally by median rami of pterygoids; 33)
pterygoids very small, median and pos-
terior rami short, anterior rami relatively
long, extending to middle of orbit; 34)
occipital condyles small, stalkc^d, widely
separated medially; 36) terminal pha-
langes T-shaped; 37 ) hyoid plate lacking
alary processes; 40) m. depressor man-
dihulae in two slips; 42) males with me-
dian subgular vocal sac; 43) body lack-
ing glands; 44) tongue narrow, posterior
one-third free; 45) toes lacking webbing
and lateral fringes, digital tips bear pads;

Figure 105. Lateral (x 7.5), dorsal, and ventral views of skull of Sminthilhis limhatus ( KU 68684,

X23).

LYNCH: LEPTODACTYLOID FROGS

159

49) adults small, less than 15 mm. SVL;

50) tympanum visible externally.

Composition. — Monotypic (S. limha-
tusj with two subspecies.

Distribution. — Cuba.

Remarks. — SmintliiUus is closely re-
lated to the West Indian species of Eleu-
therodactylus and is most similar to the
E. auriculatus group or to the E. dimi-
diatus group (as defined by Shreve and
Williams, 1963). SmintliiUus diiiers horn
the Alpha division of EleutJierodactylus
in two characters — the loss of teeth and
the loss of the alary processes of the
hyoid plate. Barbour and Noble ( 1920 )
considered SmintliiUus a dendrobatid or
ranoid derivative, but Griffiths (1959)
demonstrated that the genus was closely
related to Eleutherodactyhis, an opinion
often voiced by Noble (e.g., 1931).

SfninthiUus was named by Barbour
and Noble ( 1920 ) on the basis of a par-
tial epicoracoidal fusion. However, the
fusion is not as great as maintained by
Noble ( 1926a, 1931 ) and occurs in many
more frogs than he believed. I consider
the presence of the fusion to reflect more
accurately the care of a dissection than
morphological divergence. The other
two species named in SmintliiUus are
now placed in other genera — the Peru-
vian species (peruvianus) is placed in the
Beta division of Eleutherodactyhis (see
pp. 148-149) and the Brasilian species
(hrasiliensis) is the only species of the
genus Euparkerella.

Syrrhophus Cope, 1878
(Fig. 106)

Epirhexis Cope, 1866, J. Acad. Nat. Sci. Phila-
delphia, 6:96 [Type-species by original
designation, Batrachyla longipes Baird,
1859; suppression of the generic name re-
quested by Lynch, 1967].

Syrrhophus Cope, 1878, Amer. Nat., 12:253
[Type-species by monotypy, Syrrhophus
marnockii Cope, 1878].

Malachylodes Cope, 1879, Proc. Amer. Philos.
Soc, 18:264 [Type-species by monotypy,
Malachylodes guttilatus Cope, 1879].

Syrrhopus Boulenger, 1888, Proc. Zool. Soc.
London, p. 206 [Emendation of Syrrhophus
Cope, 1878].

Syirhaphus Giinther, 1901, Biol. Cent.-Amer.,
Kept, and Batr., p. 215 [Emendation of
SyirhopJius Cope, 1878; hence taking same
type-species (marnockii) and not verrucu-
lattis as claimed by Gorham (1966)].

Diagnostic definition. — 7) omoster-
num moderate-sized; 9) maxillary arch
toothed; 10) alary processes of premaxil-
lae directed dorsally, relatively narrow
at base; 11) palatal shelf of premaxilla
broad, deeply dissected; 12) facial lobe
of maxilla shallow; 13) palatal shelf of
maxilla broad anteriorly, narrowing pos-
teriorly, no pterygoid process; 15) nasals
large, in broad median contact; 16) na-
sals not in contact with maxillae or
pterygoids; 19) frontoparietal not orna-
mented; 20) frontoparietal fused to
prootic; 22) epiotic eminences obsolete;
23) cristae paroticae short, broad; 24)
zygomatic ramus of squamosal very
slender, relatively short; 25) otic ramus
of squamosal elongated, not forming otic
plate; 26) squamosal-maxillary angle
about 65°; 27) columella present; 28)
prevomers reduced in size, dentigerous
ramus lost, widely separated medially, or
dentigerous rami present, bearing a few
teeth; 29) palatines narrow, separated
medially, in contact with maxillae; 30)

Figure 106. Lateral and dorsal views of skull
of a male Syrrhophus pipilans (KU 59950, X 4).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

sphenethmoid entire, extending anterior-
ly beneath posterior edge of nasals; 31)
anterior ramus of parasphenoid broad,
not keeled medially; 32) parasphenoid
alae deflected posteriorly, short, widely
separated from median rami of ptery-
goids; 33) pterygoids slender, all rami
short; 34) occipital condyles small, not
stalked, widely separated medially; 36)
terminal phalanges T-shaped; 37) alary
processes of hyoid plate on narrow stalks;
40) m. depressor mandibulae in two
slips; 42) males with or without large
median subgular vocal sac; 43) axillary
and /or inguinal glands present; 44)
tongue narrow to relatively broad and
rounded, posterior edge free; 45) toes
lacking webbing, bearing lateral fringes
or not, digital tips very slightly to broad-
ly dilated into pads; 49) adults 16-40 mm.
SVL; 50) tympanum concealed or visible
externally.

Composition. — Lynch (1970a) recog-
nized 12 species, two of which are poly-
typic: cystignatlwides, dennisi, gtittila-
tus, interorJ)itaUs, leprus, longipes, mar-
nockii, modestus, nivocolimae, pipilans,
nd)rimacidatus, and verrucipes.

Distribution. — Discontinuous in the
Pacific lowlands from Sinaloa, Mexico,
to El Salvador, also in the eastern low-
lands of Mexico from the Edwards Pla-
teau of Texas to British Honduras. High-
land species occur along the Sierra
Madre Oriental up to 2000 meters.

Remarks.— l^ynch (1968a, 1970a)
discussed the generic separation of Eleu-
tlierodactyhts, Sijrrhophus, and Tomo-
dactylus. In external characters, Syr-
rhophus is not separable from all Eleii-
therodactylns. The osteological peculi-
arities of Syrrlwphus are duplicated by
Tomodactylus and the Alpha division of
Eleutlierodactyhis. Syrrhophus and To-
mod actyJus are distinguished in some
external characters (lumbar gland, ar-
rangement of the supernumerary plantar
tubercles) and by one paedomorphic
skeletal character — the sphenethmoid is
usually divided in Tomodactylus and is
entire in Syrrhophus. The division of

the sphenethmoid is a paedomorphic
feature and therefore should not be
given undue weight in any classification.
The separation of Syrrhophus and To-
modactylus as distinct genera is a de-
batable point, and I retain them as
generically distinct only as a matter of
convenience. The character of the
glands used by Lynch ( 1968a ) to sep-
arate the two genera tends to be less
diagnostic when the species of the two
genera from northwestern Mexico are
compared (S. interorhitalis, S. modestus,
T. saxatilis).

Until a comprehensive revision of the
genus Eleutlierodactyhis is completed
and the skeletons of the majority of
species studied, it will not be possible to
argue definitively whether the genera
Syrrhophus and Tomodactylus are deriv-
atives of the Alpha or the Beta divisions
of Eleutlierodactyhis. In an attempt to
clarify this point, I studied representa-
tives of all species groups of Central
American Eleiitherodactylu^ and found
no group which exhibits the osteological
characteristics of the Alpha division. If
Syrrhophus and Tomodactylus were de-
rivatives of the South American groups
of the Alpha division, one might expect
some relict species to be distributed in
parts of Lower Central America. How-
ever, all Central American species of
Eleutlierodactyhis examined by me are
Beta division Eleutherodactylus, as are
those species of the genus found in the
Choco of Colombia and Ecuador. There-
fore I suggest that the Mexican Syrrho-
phus and Tomodactylus are more closely
related to the West Indian Eleuthero-
dactylus (Alpha division) than to any
other groups of the genus. Within the
Alpha division, the auriculatus group
most closely approaches the morphology 1
of the endemic Mexican eleutherodacty-
line genera Syrrhophus and Tomodac-
tylus.

Tomodactylus Giinther, 1901

(Fig. 107)

Tomodactylus Giinther, 1901, Biol. Centr.-
Amer., Rept. and Batr., p. 219 [Type-

LYNCH: LEPTODACTYLOID FROGS

161

species by monotypy, Tomodactyhis amuJae
Giinther, 1901].

Diagnostic definition. — 7) omoster-
num moderate-sized; 9) maxillary arch
toothed; 10) alary processes of premaxil-
lae directed dorsally, broad at base; 11)
palatal shelf of premaxilla narrow, pala-
tal process elongate; 12) facial lobe of
maxilla shallow; 13) palatal shelf of
maxilla narrow, pterygoid process mod-
erate-sized; 15) nasals large, in broad
median contact; 16) nasals not in con-
tact with maxillae or pterygoids; 18)
frontoparietal fontanelle absent in adults,
often present in young males; 19) fron-
toparietals not ornamented; 20) fronto-
parietal fused to prootic; 22) epiotic
eminences obsolete; 23) cristae paroti-
cae short and stocky; 24) zygomatic
ramus of squamosal sliver-like, very
short; 25) otic ramus of squamosal very
long, no otic plate; 26) squamosal-maxil-
lary angle 50-60°; 27) columella pres-
ent; 28) prevomers reduced in size,
edentate, widely separated medially; 29)
palatines slender, widely separated me-

dially; 30) sphenethmoid usually di-
vided, not extending anteriorly to nasals;
31 ) anterior ramus of parasphenoid rela-
tively broad, not keeled medially; 32)
parasphenoid alae deflected posteriorly,
short, not overlapped by median rami of
pterygoids; 33) pterygoids slender, all
rami short; 34) occipital condyles small,
not or but slightly stalked, widely sep-
arated medially; 36) terminal phalanges
T-shaped; 37) alary processes of hyoid
plate on narrow stalks; 40) m. depressor
mandibulne in two slips; 42) males with
large, median, subgular vocal sac; 43)
lumbar gland usually well defined, axil-
lary glands sometimes present; 44)
tongue relatively small, narrow, posterior
edge free; 45) toes lacking webbing,
digital tips sHghtly to broadly dilated;
49) adults 21-31 mm. SVL; 50) tympa-
num visible externally.

Composition. — The genus was revised
by Dixon ( 1957 ) and two species were
subsequently named. Nine species are
presently recognized: albolabris, angus-
tidigitorum, dilatiis, ftiscus, grandis, niti-

FiGURE 107. Lateral, dorsal, and ventral views of skull of a female Tomodactyhis nitidus (KU 102649,

X 4.7).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

dus (3 subspecies), rufescens, saxatilis,
and syristes. Dixon and Webb (1966)
briefly commented on an unnamed spe-
cies from Nevado de Colima, Jalisco,
Mexico.

Distribution. — The Cordillera Vol-
canica of Mexico from western Veracruz
to Colima; the Oaxacan Plateau, the
Sierra Madre del Sur of Oaxaca and
Guerrero, and the Pacific lowlands from
Sinaloa to Michoacan. The genus is
largely allopatric to Sijrrhophus.

Remarks. — Gallardo (1965) placed
Tonwclactyliis in the Leptodactylinae
and Syrrliophus and Eleutherodactylus
in the Eleutherodactylinae. This asso-
ciation was based on erroneous data
concerning the breeding biology of
Tomodactylus.

Tomodactylus is primarily a lower
montane genus, whereas the closely re-
lated Syrrhophus is primarily a lowland
genus ( Lynch, 1970a ) , but the two gen-
era are sympatric in the lowlands of
western Mexico. The differences between
them are expressed to the greatest de-
gree in eastern and southern Mexico and
expressed to a lesser degree in western
Mexico, suggesting that the generic di-
chotomy occurred in western Mexico.

Tribe incertae .sedis
Scythrophrys new genus

Type-species. — Zachaenus sawayae
Cochran, 1953.

The following characteristics of the
diagnostic definition are observable:
1) sternum cartilaginous; 2) vertebral
shield lacking; 6) cranial bones not der-
mostosed; 7) omosternum small; 8)
sacral diapophyses rounded; 9) maxillary
arch toothed, teeth blunt, pedicellate;
14) maxillary arch complete; 18) fronto-
parietal fontanelle lacking; 21) temporal
arcade lacking; 24) zygomatic ramus of
squamosal relatively long, widely sep-
arated from maxilla; 28) prevomers
toothed, dentigerous processes large,
transversely elongate, situated posterior
to choanae; 36) terminal phalanges ap-

parently knobbed; 40) vi. depressor
mandilndae in two slips, pars tympan-
icus \e\y large, p. scapidaris minute;
41) pupil horizontal; 43) body lacking
glands; 44) tongue relatively large, pos-
terior edge free; 45) toes lacking web-
bing but have lateral fringes, outer meta-
tarsal tubercle present, inner metatarsal
tubercle not enlarged or spade-like, digi-
tal tips narrow on fingers, those of toes
slightly dilated, thumb very short; 49)
single adult female known is 16.9 mm.
SVL; 50) tympanum concealed.

Etymology. — G reek, scythros -{-
phryne, meaning "sullen toad."

Remarks. — A single specimen of this
southeastern Brasilian frog is known; it
was named as a member of Zachaenus
by Cochran (1953). In external appear-
ance, the frog resembles Pliysaluemus
{maculiventris group) or Paratelmato-
bius and, in some respects, Zachaenus.
Cochran (1953) considered the small
tubercles on the upper eyelid as indica-
tive of some relationship with Zachaenus
but noted the many points of disagree-
ment between parvuhis and sawayae.
The most striking difference between the
two species is seen in the length and
shape of the fingers ( Fig. 94 ) . The very
short thumb of Scythrophrys is sugges-
tive of a reduced phalangeal formula for
the hand. The tympanum is recessed
and smaller than indicated by Cochran
(1953).

Scythrophrys is placed with confi-
dence in the Telmatobiinae but is not
assigned to any tribe, because many
characters are not known. Based on the
available information, I consider the ge-
nus to belong either to the Crypiscini or
Eleutherodactylini, but until the osteol-
ogy and breeding biology of Scyth-
rophrys are known, definite assignment
to a tribe would be presumptuous.

Cochran's (1953) description of the
holotype (USNM 125530) is relatively
accurate. My measurements of the holo-
type differ somewhat from hers, reflect-
ing either differences in techniques or
possibly shrinkage. I recorded the fol-

LYNCH: LEPTODACTYLOID FROGS

163

lowing measurements (in millimeters):
snout-vent length 16.9, shank length 8.4,
head width 6.3, head length 5.6, eye
length 1.9, eyelid width 1.7, and inter-
orbital distance 2.6. A few cranial char-
acters are visible through a small tear in
the skin of the head. The frontoparietals
are broad and a fontanelle is lacking.
The nasals appear to be relatively large
and in median contact. Two characters
of the foot were not mentioned by Coch-
ran. There is a small calcar on the heel
and a narrow outer tarsal fold extending
for the length of the tarsus onto the fifth
toe.

Elosiinae Miranda-Ribeiro, 1926

Elosiidae Miranda-Ribeiro, 1926:27.
Elosiinae: Noble, 1931:504. Gallardo, 1965:
84.

Miranda-Ribeiro (1926) proposed the
Elosiidae for three genera; the concept
and content of the group has remained
unchanged since its proposal. Three
genera are presently included in the sub-
family: Crossodactijlus, Hijlodes (^EIo-
sia aucfomm), and Megaelosia. Cochran
(1938) named Crossodactijlodes, which
she considered possibly related to the
elosiines. She thought that the digital
morphology of Crossodactijlodes indi-
cated that the genus exhibited primitive
elosiine characters. The apparent dorsal
dermal glands on the digital pads of
Crossodactijlodes are artifacts reflecting
the Y-shaped terminal phalanges. Coin
and Cochran ( 1963 ) suggested that
Tracliijplirynus was related to Crosso-
dactijlus. I previously discussed this
point and placed Traclujphrijnus in the
synonymy of Eleutherodactijlus (Lynch,
1968d).

Noble (1922, 1931) considered the
Elosiinae to be a bufonid group. The
subfamily was associated with the Lep-
todactylidae by Davis ( 1936 ) , who
pointed out that the Bufonidae and Lep-
todactylidae could be familially distin-
guished. The type-genus of the subfam-
ily, Elosia Tschudi, 1838, is a synonym of
Hylodes Fitzinger, 1826 ( which is not to

be confused with Hylodes Fitzinger,
1843, which is a synonym of Eleuthero-
dactijlus Dumeril and Bibron, 1841).
The family-group name need not be
changed simply because the type-genus
is a synonym (see Article 40, Interna-
tional Code of Zoological Nomenclature,
1961). At any rate, the family-group
name could not be changed so as to be
based on Hylodes Fitzinger, 1826, be-
cause Giinther ( 1859a ) proposed a Hy-
lodidae based on Hylodes Fitzinger,
1843.

Until recently, the subfamily was
known only from southeastern Brasil and
Misiones Province, Argentina, but it is
now known to occur also in Venezuela
(Cerro Duida, Guiana Massif). The sub-
family is homogeneous morphologically
and is readily distinguished from all
other leptodactylid groups. The follow-
ing diagnostic characters are common to
the three genera of the Elosiinae: 1)
sternum cartilaginous, tending to cal-
cify in old adults; 2) vertebral shield
lacking; 3) transverse processes of an-
terior presacral vertebrae short, those of
posterior presacral vertebrae also short-
ened; 4) cervical cotylar arrangement
type I; 5) cervical and second vertebrae
not fused; 6) cranial bones not involved
in dermostosis; 7) omosternum present,
moderate-sized; 8) sacral diapophyses
rounded; 9) maxillary arch toothed,
teeth pointed, pedicellate; 12) facial
lobe of maxilla shallow; 13) palatal shelf
of maxilla narrow, no pterygoid process;
15) nasals small, widely separated me-
dially; 17) nasals not in contact with
frontoparietals; 18) frontoparietal fon-
tanelle lacking; 19) frontoparietals not
ornamented; 21) temporal arcade lack-
ing; 23) carotid artery passes dorsal to
skull bones; 27) columella present; 30)
sphenethmoid very large, entire, extend-
ing anteriorly to anterior edge of nasals;
36) terminal phalanges T-shaped; 37)
alary processes of hyoid plate on narrow
stalks; 38) cricoid cartilage not divided
ventrally; 39) m. petrohyoidetis anterior
and m. sternohyoideiis insert on lateral

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

edge of hyoid plate; 40) m. depressor
mandibuhe in two slips; 41) pupil hori-
zontal; 43) body lacking glands; 44)
tongue large, not notched, posterior edge
free; 45) toes bearing large lateral
fringes, large flap-like tarsal fold present,
outer metatarsal tubercle present, inner
metatarsal tubercle not spade-like, digi-
tal tips broad; 46) larvae with 2/3 tooth
rows, labial papillae broadly interrupted
anteriorly; 47) amplexus axillaiy in
known species; 48) eggs small, numer-
ous, laid in moist terrestrial situations or
in ponds or streams; 50) tympanum
visible externally; 51) each digital pad
has a pair of dermal, scute-like glandu-
lar pads on dorsal surface. The verte-
bral arches of all elosiines are very non-
imbricate and the vertebrae and coccyx
are poorly ossified.

The Elosiinae are of particular in-
terest in that the poison-arrow frogs
( Dendrobatidae ) are apparently derived
from this leptodactylid subfamily. The
two groups agree in cranial morphology,
vertebral columns, the T-shaped termi-
nal phalanges, the dermal glandular
pads on top of the digital pads, and in
the presence, in at least some species of
each group, of toxic skin secretions (the
secretions of the elosiines have not been
chemically analyzed). The two groups
differ in breeding biology and in the
architecture of the pectoral girdle.

Crossodactylus best fits my concept
of the primitive elosiine but has diverged
in at least one character — the loss of the
quadratojugal. The genus is distinctive
in its ranoid pattern of the attachment of
the distal tendons of the thigh muscula-
ture. The ranoid pattern of the thigh
musculature of Crossodactylus is exactly
like that seen in the dendrobatids and
mitigates the importance of one of the
criteria used by Griffiths ( 1963 ) to asso-
ciate the dendrobatids as a Neotropical
subfamily of the Ranidae. Crossodacty-
lus has a median subgular vocal sac and
nuptial asperities (cluster of spines) in
the males. This condition is like that
seen in most Telmatobiinae (excepting

the Eleutherodactylini which lack nup-
tial asperities); I regard the presence of
vocal sac and nuptial asperities as primi-
tive. The tadpoles of Crossodactylus
have median vents in contrast to the
dextral vents of the tadpoles of Hylodes
and Megaelosia. Hylodes and Megae-
losia have the bufonid pattern of the
arrangement of the distal tendons of the
thigh musculature, derived conditions of
the vocal apparatus, and quadratojugal
bones. I envision the dendrobatids as
having diverged from the Crossodactylus
stock prior to the loss of the quadrato-
jugal, but after Hylodes and Megaelosia
had evolved.

Crossodactylus Dumeril and Bibron,
1841

(Fig. 108)

Crossodactylus Dumeril and Bibron, 1841,
Erpetologie generale, 8:635 [Type-species
by monotypy, Crossodactylus gaudichaudii
Dumeril and Bibron, 1841].

Limnocharis Bell, in Darwin, 1843, Zool. Voy-
age Beagle, Reptiles, 5:33 [Type-species by
monotypy, Limnocharis fuscus Bell, 1843].

Tarsopterus Reinhardt and Liitken, 1862, Vid.
Meddel. Naturh., Foren., 1861:177 [Type-
species by monotypy, Tarsopterus tracluj-
stomus Reinhardt and Liitken, 1862].

Calamohates DeWitte, 1930, Miss. Biol. Beige
Bresil, 2:219 [Type-species by monotypy,
Calamohates boidengeri DeWitte, 1930].

Diagnostic definition. — 10) alary proc-
esses of premaxillae directed anterodor-
sally and laterally, broad at base; 11)
palatal shelf of premaxilla narrow, pala-
tal process elongate; 14) maxillary arch
incomplete, maxilla tapering posteriorly,
quadratojugal absent; 16) nasals sep-
arated from both maxillae and ptery-
goids; 20) frontoparietal fused to pro-
otic; 22) epiotic eminences moderately
distinct; 23) cristae paroticae short,
stocky; 24) zygomatic ramus of squa-
mosal of moderate length, pointed,
widely separated from maxilla; 25) otic
ramus of squamosal slightly shorter than
zygomatic ramus, no otic plate; 26)
squamosal-maxillary angle about 50°;
28) prevomers moderately small, sep-
arated medially, dentigerous ramus lost

LYNCH: LEPTODACTYLOID FROGS

165

or not, rarely toothed; 29) palatines
small, narrow, widely separated medial-
ly; 31) anterior ramus of parasphenoid
short, broad, not keeled medially; 32)
parasphenoid alae oriented at right
angles to anterior ramus, short, widely
separated from median rami of ptery-
goids; 33) pterygoids with slender rami,
anterior rami not reaching palatines; 34)
occipital condyles small, stalked, widely
separated medially; 35) mandible lack-
ing odontoids; 42) males with median,
subgular vocal sac, nuptial spines in a
cluster in all species; 46) larvae with
median vent; 49) adults relatively small,
less than 40 mm. SVL.

Composition. — Six species are pres-
ently recognized (aeneus, dispar, gaudi-
chaudii, grandis, schmidti, and tracluj-
stoma), although some authors favor re-
garding dispar and grandis as conspe-
cific. The most recent revision of the
genus (Cochran, 1955) did not treat C.
grandis B. Lutz, 1951, or C. schmidti
Gallardo, 1961.

Distribution. — Southeastern Brasil in

the lowlands from southern Minas Gerais
to Misiones Province, Argentina.

Remarks. — The generic synonymy of
Crossodactylus has been stable for many
years. Cochran (1955) and Gallardo
( 1961 ) presented studies of intrageneric
variation and relationships. Crossodac-
tylus is primitive to the other elosiines in
tadpole morphology (median vent) and
in the secondary sex characteristics (me-
dian subgular vocal sac and nuptial as-
perities ) , but is specialized in the loss of
the quadratojugal. Hylodes is similar to
Crossodactylus in having a poorly ossi-
fied quadratojugal, but differs from
Crossodactylus in the other characteris-
tics mentioned.

The ranoid pattern of the attachment
of the distal tendons of the thigh muscu-
latme of Crossodactylus distinguishes
the genus from the other elosiines, as
well as from all other Neotropical lepto-
dactylids. The thigh musculature of
Crossodactylus adds yet another charac-
ter to the impressive list of characters
shared by the dendrobatids and Crosso-

FiGiTRE 108. Lateral, dorsal, and ventral views of skull of Crossodactylus gaudichaudii ( KU 92759,

X 4.7).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

dactijlus. Noble ( 1931 ) argued that the
Dendrobatinae were derived from the
bufonid genus Crossodactyhis, because
he found the digital pad structure of the
two groups to be identical and to exist
in no other Neotropical bufonoid frogs.
Noble argued that the condition of the
epicoracoidal cartilages of Crossodacty-
his is precursorial to the condition seen
in developing PlnjUobates (:=Coloste-
thus) sidjpunctatus. Noble considered
his data adequate to demonstrate that
the firmisternal dendrobatids passed
through an arciferal condition in ontog-
eny. Griffiths (1959, 1963) sought to
ally the dendrobatids with ranids and
argued that the dendrobatids do not pass
through an arciferal stage in develop-
ment. The developmental pattern of the
pectoral girdle exhibited by Colostethus
sidjpunctatiis is clearly ranoid. My own
study of this subject completely supports
that of Griffiths.

Griffiths (1963) rejected Noble's
argument that the thigh musculature was
of primary importance in anuran classifi-
cation, but cited the thigh musculature
as additional evidence supporting his
contention that dendrobatids are a ranid
subfamily. Griffiths cited the similar de-
velopment of the digital pads of the
Petropedetinae (Ranidae) as supportive
evidence for the close relationship be-
tween the dendrobatids and ranids. His
argument requires that we ignore the
many similarities between elosiines and
dendrobatids and regard several char-
acter complexes as evolving in a parallel
fashion in leptodactylids and ranids.
Associating the dendrobatids with the
bufonoids requires that we regard the
firmisternal pectoral girdles of dendro-
batids and ranids to have been inde-
pendently evolved. This position is made
less objectionable by the occurrence of a
nearly complete transition from arcifery
to firmisterny within the Neotropical
Bufonidae. Noble's (1922) position that
the firmisternal pectoral girdle has ap-
peared more than once is regarded as
correct.

Griffiths ( 1963 ) cited several other
characters as exclusively ranoid. The
bursa angularis oris is not restricted to
dendrobatids and ranids as Griffiths
stated but occurs in most, if not all,
groups of advanced frogs (Baldauf and
Tanzer, 1965, Trueb, 1966, 1968, 1970,
and Starrett, 1968).

The dendrobatids are amply distinct
from the elosiine leptodactylids. I do
not intend to present an argument that
the two families ought to be combined,
because I think that there is value in
recognizing the small, brightly colored,
poisonous Neotropical dendrobatid frogs
as familially distinct. The dendrobatids
have lost the palatines, which are re-
tained, although they are small, in elo-
siine leptodactylids. The firmisternal
architecture of the pectoral girdle of
dendrobatids is markedly different from
the arciferal architecture exhibited by all
leptodactylids. The breeding behavior
and biology of the dendrobatids is not
unique among frogs but is very different
from that of the elosiine leptodactylids.
Dendrobatid tadpoles usually have 2/3
tooth rows, although several species have
reduced numbers (Starrett, 1960). The
tadpoles of all dendrobatids have a broad
anterior interruption of the labial papil-
lae as do most leptodactylids. Dendro-
bates has either a median or dextral
vent, whereas all known tadpoles of
Colostethus and Plujlhbates have dex-
tral vents.

Hylodes Fitzinger, 1826

(Fig. 109)

Hylodes Fitzinger, 1826, Neue Class. Kept., p.

38 [Type-species by monotypy, Hylodes

ranoides (=HyIa ranoides Spix, 1824)].
Enydwbius Wagler, 1830, Nat. Syst. Ampli., p.

202 [Substitute name for Hylodes Fitzinger,

1826; hence taking same type-species].
Elosia Tschudi, 1838, Classif. Batr., p. 77

[Type-species by monotypy, Elosia nasiita

Tschudi, 1838].
Scimicodes Fitzinger, 1843, Syst. Kept., p. 32

[Type-species by original designation, Hyla

nastis Lichtenstein, 1823].

Diagnostic definition. — 10) alary proc-
esses of premaxillae directed anterodor-

LYNCH: LEPTODACTYLOID FROGS

167

sally and laterally, broad at base; 11)
palatal shelf of premaxilla relatively
narrow, palatal process relatively small;
14) maxillary arch complete, quadrato-
jugal poorly ossified; 16) nasals not in
contact with maxillae or pterygoids; 20)
frontoparietals fused to prootics; 22)
epiotic eminences moderately distinct;
23) cristae paroticae short, stocky; 24)
zygomatic ramus of squamosal short,
truncate, widely separated from maxilla;
25) otic ramus of squamosal about as
long as zygomatic ramus, not expanded
into otic plate; 26) squamosal-maxillary
angle about 40°; 28) prevomers entire,
toothed, separated medially; 29) pala-
tines long, narrow, widely separated me-
dially; 31) anterior ramus of parasphe-
noid short, broad, keeled medially in at
least some species; 32) parasphenoid
alae oriented at right angles to anterior
ramus, narrowly overlapped laterally by
median rami of pterygoids; 33) ptery-
goids relatively small, rami slender, an-
terior rami elongate, reaching palatines;
34) occipital condyles small, not stalked,
widely separated medially; 35) mandible
lacking odontoids; 42) males with
paired, lateral, membranous vocal sacs,

absent in one species, and with nuptial
asperities in a pad on thumb; 46) larvae
with dextral vent; 49) adults 30-45 mm.
SVL.

Composition. — Bokermann ( 1966)
listed nine species (aspera, glahrus, lat-
eristrigatus, magalhaesi, mertensi, merid-
ionaUs, nasus, perplicatus, and piilcher)
in the genus, then known only from
southeastern Brasil. Gorham (1966)
listed gla])nis as a synonym of lateri-
strigatiis and meridionalis as a subspecies
of nasus. Bokermann ( 1967 ) named an
additional species from Rio de Janeiro
(ornata), and Rivero (1968) named dui-
densis from Venezuela. All of these
authors used the generic name Elosia, as
did Cochran ( 1955 ) in her study of the
species of southeastern Brasil.

Distribution. — Coastal southern and
southeastern Brasil from Rio Grande do
Sul north to Minas Gerais. One species
occurs on Cerro Duida in Amazonian
Venezuela.

Remarks. — Frogs of this genus ex-
hibit relatively little intrageneric varia-
tion and have been recognized as a dis-
tinctive generic unit for many decades.

Figure 109. Lateral, dorsal, and ventral views of skull of Hylodes asper ( KU 92870, X 4).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

The nomenclatorial problems of the
genus are by no means minor. Myers
(1962) pointed out that the proper ge-
neric name for these frogs is Hylodes
Fitzinger, 1826, and not Elosia Tschudi,
1838, the name which had been appHed
more or less universally for 60 years.
The usage of one generic name in pref-
erence to another in a significant work-
must be taken into account when dealing
with any question of priority and /or
nomenclatorial stability. It is therefore
significant that Cochran (1955) used
Ehsia in her important study of the
frogs of southeastern Brasil. However,
the argument against usage of Hylodes
as the proper generic name is based on
the fact that Fitzinger proposed Hylodes
twice, each time with a different type-
species. Fitzinger (1826) proposed Hy-
lodes for Hyla ranoides Spix, 1824, a
member of the group later named Elosia
by Tschudi (1838), and in 1843 pro-
posed Hylodes for Hylodes martinicensis
Tschudi, 1838, the type-species of Eleu-
therodactyhis Dumeril and Bibron, 1841.
I am in complete agreement with Myers
( 1962 ) , because there is no longer any
confusion of what Hylodes is — the last
author to use it in the sense of Fitzinger
(1843) was Melin (1941). We have
used Eleutherodactylus as the proper
generic name since the early part of this
centuiy when Stejneger (1904) pointed
out the synonymies of Fitzinger's names.
The genus Hylodes (or Ehsia) is a small
one; even the least conservative author
would not recognize a dozen species.
The genus is restricted in distribution,
and the species of the genus are rela-
tively rare and therefore have not been
frequently mentioned in the literature.
A nomenclatorial change at the generic
level creates relatively little instability
even when the generic name used is one
that has a junior homonym that is far
better known. I do not regard the use of
Elosia defensible while there are two
older generic names (Hylodes and Eny-
drobius) for the group.

Megaelosia Miranda-Ribeiro, 1923
(Fig. 110)

Megaelosia Miranda-Ribeiro, 1923, Rev. Mus.
Paulista, 13:819 [Type-species by moiio-
typy, Megaelosia bufonia Miranda-Ribeiro,
1923].

Diagnostic definition. — 9) maxillary
teeth very long; 10) alary processes of
premaxillae directed sharply posterodor-
sally; 11) palatal shelf of premaxilla of
moderate depth, palatal process elon-
gate; 14) maxillary arch complete, pos-
terior end of maxilla expanded, quad-
ratojugal present, deep; 16) nasals in
contact with maxillae, separated from
pterygoids; 20) frontoparietals not fused
to prootics; 22) epiotic eminences obso-
lete; 23) cristae paroticae broad, stocky;

24) zygomatic ramus of squamosal long,
expanded, in broad contact with maxilla;

25) otic ramus of squamosal relatively
long, expanded medially into small otic
plate; 26) squamosal-maxillary angle
about 15°; 28) prevomers moderate-
sized, entire, toothed; 29) palatines elon-
gate, relatively broad, widely separated
medially; 31) anterior ramus of para-
sphenoid elongate, narrow, not keeled
medially; 32) parasphenoid alae de-
flected posteriorly, broadly overlapped
laterally by median rami of pterygoids;
33) pterygoids large, anterior rami not
reaching middle of orbits; 34) occipital
condyles moderately large, not stalked,
narrowly separated medially; 35) man-
dible bearing a serrated odontoid ridge;
42) males lacking vocal sac and nuptial
asperities; 46) larvae with dextral vent;
49) males reach 70 mm., females reach
120 mm. SVL.

Composition. — M o n o ty p i c (goeldi);
the type-species of the genus is a junior
synonym.

Disiribution.— The Coastal Ranges of
southeastern Brasil (Estados Rio de
Janeiro and adjacent Minas Gerais and
Sao Paulo).

Remarks. — Ever since its separation
from Hylodes (Ehsia aiictorum), Megae-
losia has been a poorly defined genus.
Noble (1931) and Cochran (1955) re-

LYNCH: LEPTODACTYLOID FROGS

169

marked that Megaelosia was merely a
giant Elosia with somewhat enlarged
maxillary teeth. Cochran ( 1955 ) recog-
nized the genus because of its greater
adult size.

Megaelosia has diverged markedly
from the other elosiines. The external
morphology of M. goeldi compels me to
retain it in the Elosiinae. The skull of
this monotypic genus is strikingly dif-
ferent from those of the other genera of
the subfamily (see Figs. 108-10). In
contrast to the rather delicate maxillary
arch in the other genera of the subfam-
ily, Megaelosia has a large, massive max-
illa and quadratojugal. The teeth of
Megaelosia are fang-like and much
larger than those of the other genera of
the subfamily. The squamosal architec-
ture of Megaelosia is very different from
that seen in the other elosiines; the zygo-
matic ramus is enlarged and in broad
contact with the maxilla, and the otic
ramus of Megaelosia is more like that
seen in the Grypiscini ( Telmatobiinae )
than that in Hylodes or Crossodactijlus.
The very large (compared to those of
the other elosiines) occipital condyles of

Megaelosia are suggestive that the genus
is primitive. In external morphology,
Megaelosia goeldi is very similar to Hy-
lodes. The tadpoles of the two genera
are very difficult to separate. These data
suggest that the two genera are indeed
related although the skull morphology
suggests that the external similarities are
convergent or parallel.

Leptodactylinae Berg, 1896 ( 1838 )

Cystignathi Tschudi, 1838:78.

Cystignathidae: Giindier, 1859a: 26.

Pleurodemae Cope, 1866:95.

Paludicolina Mivart, 1869:290.

Plectromantidae Mivart, 1869:291.

Cystignathina: Mivart, 1869:293-94.

Leptodactylidae Berg, 1896:161 [A replace-
ment name for the Cystignathidae, the type-
genus of which is a synonym of Leptodac-
tylus].

Cystignathinae: Gadow, 1901:211.

PaludicoHdae: Miranda-Ribeiro, 1926:153.

LeptodactyHnae: Noble, 1931:504.

Pseudopaludicolinae Gallardo, 1965:84.

The unifying characteristic of this
subfamily is the bony style or osseous
plate in the sternum as compared with
the cartilaginous sterna of the other lep-
todactylids. The group is strictly Neo-

FiGURE 110. Lateral, dorsal, and ventral views of skull of Megaelosia goeldi (KU 106271, X 2.3).

170

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

tropical and ranges south to southern
Chile and north to the southern United
States. For the most part, the group is a
lowland component. The widespread
genus Leptodactylus rarely reaches ele-
\'ations abo\'e 1000 meters. The genus
Pleurodema occurs in the Andean system
in Chile, Bolivia and southern Peru, and
therefore reaches elevations exceeding
4000 meters. Even at these elevations,
the group breeds in ponds. The pond-
breeding habits of the Leptodactylinae
have restricted the dispersal of the group
to lower elevations. Many of the species
of the subfamily are the wide-spread,
common, lowland frogs encountered in
most tropical situations in the Americas.

The following characteristics of the
diagnostic definitions are common to all
ten of the included genera : 1 ) sternum
containing an osseous element; 2) ver-
tebral shield lacking; 3) transverse proc-
esses of anterior presacral vertebrae not
expanded or shortened; 5) cervical and
second vertebrae not fused; 6) cranial
bones not involved in dermostosis; 7)
omosternum present; manubrium ex-
panded in all genera except Paratelma-
tobius and PJu/salaemtis; 9) maxillary
arch usually toothed, if present, teeth
blunt, pedicellate; 11) palatal process of
premaxilla long; 20) frontoparietal not
fused to prootic; 21) temporal arcade
lacking; 23) carotid artery passes dorsal
to skull bones; 24) zygomatic ramus of
squamosal widely separated from maxil-
la; 30) sphenethmoid entire; 35) mandi-
ble lacking odontoids; 38) cricoid carti-
lage not divided ventrally; 40) m. de-
pressor mandihuJae in two slips; 44) pos-
terior edge of tongue free; 45) outer
metatarsal tubercle present; 47) amplex-
us axillary in all observed species.

Most herpetologists familiar with the
Neotropical fauna have recognized two
informal groups of aquatic breeding lep-
todactyline frogs — those associated with
Leptodactylus (Hydrolaetare, Leptodac-
tyhis, Limnomedusa, and Litlwdytes)
and those long called "PahidicoUr (En-
gystomops, Eupemphix, Paratelmato-

hius, PhysaJaemus, Pleurodema, and
Pseiidopaludicola). The rare genus Eda-
lorhina is usually associated with the lat-
ter group but bears considerable resem-
blance to Lithodytes.

Boulenger (1882) was familiar with
most of the generic groups here included
in the Leptodactylinae. Hydrolaetare
and Paratelmaiohius were described
subsequent to his studies. Boulenger in-
cluded Enfi,ystomops and Eupempliix in
the Bufonidae and incorrectly associated
Hylorina with the Leptodactylinae, be-
cause he believed that H. sylvatica had
a bony sternum. One of the genera rec-
ognized by Boulenger was Paludicola,
which he considered to be wide-spread
and generalized. Mehely (1904) and
Parker ( 1927 ) pointed out the hetero-
geneity of Boulenger's Paludicola and
each proposed a partitioning based on
osteological characters. Mehely divided
Paludicola into two genera — Paludicola
and Pleurodema. Parker (1927) divided
it into three genera — Physalaemus, Pleu-
rodema, and Pseudopaludicola. Mehely
separated Boulenger's Paludicola into
one group with prevomerine teeth and a
simple (non-bifurcate) sternal style
(Pleurodema) and into another group
without prevomerine teeth and with a
bifurcate sternal style (Paludicola). Nie-
den (1923) uncritically followed Me-
hely's system and included Edalorhina
in Pleurodema. Parker (1927) criticized
Mehely's arrangement because relatively
few species had been studied; he pro-
posed another classification of the palu-
dicoline frogs based on loss of the pre-
vomerine teeth, loss of the quadratojugal,
shape of the sternal style, and the pres-
ence of an antebrachial tubercle. Parker
(1927) characterized Pseudopaludicola
as having an elongate, cartilaginous, or
calcified sternum. Virtually all subse-
ciuent authors have repeated Parker's
characterization of the sternum of Pseu-
dopaludicola ( Barrio, 1954, Cochran,
1955, Rivero, 1961, and Gallardo, 1965).
Principally on the basis of the cartilagi-
nous sternum and breeding habits, Gal-

LYNCH: LEPTODACTYLOID FROGS

171

lardo ( 1965 ) proposed a new subfamily
for Pseudopahidicola. In the present
study, at least one specimen of each of
five currently recognized species of Pseu-
dopahidicola was cleared and stained.
The sternal style is narrow and elongate
and is much more dense than the epi-
coracoidal cartilages, but is not more
dense than the coracoid bones. I con-
sider the sternal style of Pseudopahidi-
cola to be osseous. Parker (1927) re-
ported simple and T-shaped terminal
phalanges in frogs of the genus Pseudo-
pahidicola. All specimens which I exam-
ined have knobbed terminal phalanges.

The foam-nesting habits of leptodac-
tylines have been known for some time
and have been used in the classification
of the group ( Noble, 1927, Breder, 1946,
Bokermann, 1962). Barrio (1954) first
reported the ethological differences be-
tween PJn/salaemus and Pseudopahidi-
cola. P.seudopaludicola lays its eggs
singly or in small clumps in water with-
out the benefit of a foam-nest. The spe-
cies of Physalaemus (including Engy-
stomops and Eupemphix) lay their eggs
in a foam-nest floating on water (Fig.
2). Frogs of the genus Leptodactyhis
also make foam-nests but there is con-
siderable variation within the genus. The
species of the melanonotus and ocellatus
groups lay their eggs in a foam-nest
floating on water, as do Physalaemus and
Pleurodema. The species of the pen-
tadactylus group differ only slightly in
that the foam-nest is deposited in pot
holes filled with water along the edges
of streams or ponds. The species in the
fuscus or sibilatrix group deposit their
eggs in a foam-nest in a burrow, and the
tadpoles emerge after the nest is inun-
dated. This recalls the situation seen in
Heleioponis. The frogs of the nmnnora-
tus group deposit their few, large eggs
in a terrestrial underground incubating
chamber in a foam-nest. There are no
aquatic larvae. Edalorhina has aquatic
larvae (R. Etheridge, in litt.). Repro-
ductive data are unavailable for Bary-

cholos, Hydrolaetare, Limnomedusa,
Lithodytes, and Paratehnatobius.

The subfamily Leptodactylinae was
defined by Noble ( 1931 ) on the basis of
the presence of an osseous style in the
sternum. Other authors suggested that
the foam-nest habits are characteristic of
the subfamily, but these authors mis-
takenly believed that Pseudopaludicola
is not related to the Leptodactylinae. In
view of the appearance and variability
of sternal styles and osseous plates else-
where among the Salientia, it can be
argued that the subfamily Leptodactyli-
nae is poorly defined and possibly poly-
phyletic (see generic account for Para-
telmatohiiis, pp. 182-4). Progressively
graduated vicinal similarities of several
characters within this group of genera
were used to infer relationship through
the whole set of genera.

The sternum is an osseous plate in
Paratelmatohius with a large cartilagi-
nous xiphisternum. In the other nine
genera of the subfamily, the sternum is a
style. In Pleurodema the style is rela-
tively broad, and in Limnomedusa, Bary-
cholos, Edalorhina, and Physalaemus,
the style is only slightly narrower than
that of Pleurodema. The sternal style is
elongate and narrow in Hydrolaetare,
Leptodactyhis, Lithodytes, and Pseudo-
paludicola. I consider the presence of a
discrete style in the sternum as sound
evidence of close relationship. The rela-
tionships of Paratelmatohius are obscure
— it does not have a bony style in the
sternum. The transverse processes of the
posterior presacral vertebrae are some-
what shortened in Lithodytes, Paratel-
matohius, Physalaemus, Pleurodema, and
Pseudopaludicola, but are not shortened
in Barycholos, Edalorhina, Hydrolaetare,
Leptodactyhis, or Limnomedusa. The
occipital condyles are relatively large
and narrowly separated in Limnomedusa
and Pleurodema. Limnomedusa has a
type II cervical cotylar arrangement
whereas, all other genera of the subfam-
ily have a type I cervical cotylar ar-
rangement.

172

MISCELLANEOUS PUBLICATIOiX MUSEUM OF NATURAL HISTORY

I consider Pleurodema most like the
primitive leptodactyline stock because it
has a generahzed body form, pectoral
architecture, skeleton, and tadpole.
Pleurodema is specialized in one inter-
esting character — the loss of the quad-
ratojugal. Pleurodema is externally sim-
ilar to Eupsophus, and the external simi-
larity reflects the similarity in the osteol-
ogy of these two genera. The two gen-
era are readily distinguished by the loss
of the quadratojugal in Pleurodema and
the presence of a bony style in the ster-
num of Pleurodema. The tadpole of
Pleurodema has a median vent, 2/3 tooth
rows, and a broad anterior interruption
of the labial papillae, as do the tadpoles
of Eupsophus and Leptodactylus. The
tadpoles of Phijsalaemus and Pseudo-
paludicola have a dextral vent, 2/3 tooth
rows, and a broad anterior interruption
of the labial papillae. Noble ( 1931 ) sug-
gested that Physalaemiis was the stem
genus of the Leptodactylinae, but my
study suggests that Pleurodema more ac-
curately fills this role. Physalaemus has
departed from other leptodactylines in

several morphological features, the most
striking of which is the hyolaryngeal ap-
paratus.

Pleurodema Tschudi,

(Fig. Ill)

1838

Pleurodema Tschudi, 1838, Class. Batr., p. 47
[Type-species by monotypy, Pleurodenm
hibroni Tschudi, 1838].

Leiiipenis Dumeiil and Bibron, 1841, Erpetol-
ogie generale, 8:420 [Type-species by
monotypy, Leiii penis marmorattis Dumeril
and Bibron, 1841].

Fhijsalaemus Fitzinger {non Fhijsalaemus Fitz-
inger, 1826), 1843, Syst. Rept., p. 31
[Type-species by original designation, Cijs-
tignathus bihroni of Dumeril and Bibron,
1841 { =:Pleurodema bibroni Tschudi,
1838)].

Lystiis Cope, 1868, Proc. Acad. Nat. Sci. Phila-
delphia, 20:312 [Type-species by monotypy,
Lystiis biachyops Cope, 1868].

Diagnostic definition. — 1 ) sternum
bearing a broad, osseous style which
tends to bifurcate in large adult females;
3) transverse processes of posterior pre-
sacral vertebrae somewhat shortened; 4)
cervical cotylar arrangement type I; 7)
omosternum cartilaginous, usually not

Figure 111. Lateral, dorsal, and ventral views of skull of Pleurodema cinerea (KU 80836, X 4).

LYNCH: LEPTODACTYLOID FROGS

173

elongated, manubrium large; 8) sacral
diapophyses slightly dilated; 9) maxil-
lary arch toothed; 10) alary processes of
premaxillae directed posterodorsally,
broad at base; 11) palatal shelf of pre-
maxilla broad; 12) facial lobe of maxilla
deep; 13) palatal shelf of maxilla nar-
row, pterygoid process lacking; 14) max-
illary arch incomplete, quadratojugal
lacking; 15) nasals in median contact,
relatively small; 16) nasals not in contact
with maxillae or pterygoids; 17) nasals
not in contact with frontoparietals; 18)
frontoparietal fontanelle large; 19) fron-
toparietals not ornamented; 22) epiotic
eminences small; 23) cristae paroticae
relatively short, narrow; 24) zygomatic
ramus of squamosal relatively short; 25)
otic ramus of squamosal slightly longer
than zygomatic ramus, no otic plate; 26)
squamosal-maxillary angle 40-45°; 27)
columellae present; 28) prevomers rela-
tively large, toothed, narrowly separated
medially; 29) palatines relatively nar-
row, arched, narrowly separated medial-
ly; 30) sphenethmoid large, extending
anteriorly to posterior edge of nasals;
31 ) anterior ramus of parasphenoid long,
narrow, not keeled medially; 32) para-
sphenoid alae oriented at right angles to
anterior ramus, broadly separated from
median rami of pterygoids; 33) ptery-
goids relatively slender, anterior rami
long, reaching palatines; 34) occipital
condyles large, not stalked, widely sep-
arated medially; 36) terminal phalanges
knobbed; 37) alary processes of hyoid
plate on narrow stalks; 39) m. petro-
htjoideiis anterior and m. sternohijoideus
insert on lateral edge of hyoid plate; 41)
pupil horizontal; 42) males with median
subgular vocal sac, nuptial asperities on
thumb; 43) lumbar glands present or
not; 44) tongue large, round; 45) toes
lacking webbing, with lateral fringes or
not, metatarsal tubercles spade-like or
not, digital tips narrow; 46) larvae with
median vent, 2/3 tooth rows, labial
papillae broadly interrupted anteriorly;
48) eggs laid in foam-nest in water,
small and numerous; 49) adults 35-65

mm. SVL; 50) tympanum visible exter-
nally or concealed.

Composition. — Ten species are pres-
ently recognized: hihroni, brachyops,
hufonina, cinerea, darwinii, diplolistris,
guayapae, marmorata, nebulosa, and tu-
cumanay- The widespread P. hihroni is
probably a composite superspecies.

Distrihution. — Central Andean Peru
south to southern Chile and Argentina
and northeastward to Uruguay, along
the coastal lowlands of extreme eastern
Brasil in non-forested habitats; the arid
and semiarid coastal belt from the Guay-
anas through Venezuela to the Maricaibo
Basin, the islands north of Venezuela,
and in the savanas of central Panama.

Remarks. — With the exception of P.
hrachyops and P. diplolistris, frogs of the
genus Pleiirodema are restricted to
southern South America and temperate
climates; P. diplolisiris occurs in the sub-
tropical areas of eastern Brasil, and P.
marmorata ranges northward in the high
Andes to centi'al Peru. Leiupertis ver-
rucosus Reinhardt and Liitken was
placed in Pleiirodema by Parker (1927),
who was uncritically followed by all
subsequent authors. The species is a
member of the genus Ischnocnema (Tel-
matobiinae, Eleutherodactylini) .

Parker (1927) included 18 nominal
species in Pleurodema. Since his revi-
sion of the genus one new species (guay-
apae) has been named; Pleurodema illota
is now placed in Eupsophus, and P. mex-
icanus is an Eleutlierodactylus. Parker
recognized P. pseudopliryne Philippi and
P. montevidense Philippi, which are now
considered synonyms of P. hihroni and
P. dartvinii, respectively. Parker in-
cluded Pleurodema coquimhensis in

^^ Donoso-Barros (1969) pointed out that Pleu-
rodema bibrotiii Tschudi, 1838, is the same as
Pleurodema darwinii Bell, 1843, and that the
taxon Dumeril and Bibron ( 1841 ) called Pleu-
rodema bihronii is Bufo tliatil Lesson, 1826.
Thus Pleurodema bibronii Tschudi applies to
the Uruguayan species called P. darwinii, and
the correct name for the Chilean frogs called
P. bibronii by most authors is P. thaul (Lesson),
1826.

174

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Physalaemus, but Cei (1962a) pointed
out that it is a synonym of Pleurodema
hibroni.

At present, the most pressing sys-
tematic problem in the genus Pleuro-
dema is the status of the species pres-
ently called P. hibroni and P. cinerea.
Schmidt (1954b) recognized P. plebeya
Philippi for the southern populations
now called P. bihroni. Cei (1962a) com-
bined the two nominal species but rea-
lized that the complex is seriously in
need of further study. Pleurodema cin-
erea is closely related to P. hibroni; the
two species are distinguished only by
the external expression of the tympanum
(concealed in bihroni, evident in cin-
erea). The hibroni-cinerea complex is
much in need of a detailed review.

Of the ten species of Pleurodema
recognized here, five (bihroni, hrachij-
ops, hufonina, cinerea, and dancinii)
have lumbar or lumbo-inguinal glands.
These glands are well-defined, often
brightly patterned, and present in both
sexes. At least some of these species use
the glands in a defense posture. The
frog arches its back and tucks its head
down thus presenting the large, pat-
terned glands to a predator or aggressor.
In this position, the glands appear to be
large eyes (Cei, 1962a). The lumbar
glands are moderate-sized in hibroni,
hrachyops, cinerea, and darwinii, and
very large in hufonina.

Parker ( 1927 ) suggested that the spe-
cies of Pleurodema with prevomerine
teeth were more primitive than those
lacking prevomerine teeth. Like so
many other herpetologists of the 19th
and early 20th centuries, Parker (1927)
regarded the presence or absence of
prevomerine teeth as a major character
in leptodactylid classification. His pri-
mary "key character" in subdividing the
genus was the presence or absence of
prevomerine (vomerine) teeth. As has
been repeatedly pointed out in recent
years, the prevomerine teeth are readily
lost in many groups of frogs. The teeth
may be present but concealed beneath

the tissue of the palate ( as in Eleuthero-
dactylus myersi) or they may be lost
entirely. In most of the cases where the
presence of prevomerine teeth has been
reported to be variable within a species,
I have found that the teeth are present
but concealed beneath the tissue of the
palate. Main (1957) criticized the use
of the presence of prevomerine teeth as
the primary character in the division of
the species of Crinia into two groups.

In four species of Pleurodema (hi-
broni, hufonina, cinerea, and marmo-
rata), the metatarsal tubercles are not
enlarged. In these species, the outer
metatarsal tubercle is small and conical
and the inner metatarsal tubercle is an
elongate oval. In the other six species of
the genus, the inner metatarsal tubercle
is enlarged, laterally compressed, and
spade-like. In these six species, the outer
metatarsal tubercle is enlarged and
either compressed (hrachyops, diplolis-
tris, '^uayapae, and nebulosa) or not
(dancinii and tucumana).

Pleurodema hibroni and cinerea have
a short inner tarsal fold, and hufonina
has a long inner tarsal fold; all other
species of the genus lack tarsal folds.
Pleurodema diplolistris has a prominent
tarsal tubercle, much like that seen in
many species of Physalaemus. Pleuro-
dema nebulosa has a fimbriated anal
flap, whereas no other species of the
genus has more than a low transverse
ridge above the anal opening. The sig-
nificance of these characters is not ap-
parent at this time, although the pres-
ence of a tarsal fold in bihroni, cinerea,
and hufonina is suggestive that they are
closely related. These three species also
agree in having lumbar glands and in
having small, non-compressed metatar-
sal tubercles.

Of the eleven species listed in the
genus Pleurodema by Gorham (1966),
two are not included in this genus by
me: sa<^ittifer O. Schmidt, 1857, is here
treated as a species inquierenda, and
verrucosus Reinhardt and Liitken, 1862,

LYNCH: LEPTODACTYLOID FROGS

175

is placed in the genus Ischnocnema (Tel-
matobiinae, Eleutherodactylini ) .

Limnomedusa Fitzinger, 1843

(Fig. 112)

Limnomedusa Fitzinger, 1843, Syst. Rept., p.
31 [Type-species by original designation,
Ciistifinatliu.s inacwglossits Dumeril and
Bibron, 1841].

Litopleiira Jimenez de la Espada, 1875, Vert.
Viaje Pacif., Batr., p. 82 [Type-species by
monotypy, Litopleiira maritimum Jimenez
de la Espada, 1875].

Diagnostic definition. — 1 ) sternum
bearing a broad osseous style; 3) trans-
verse processes of posterior presacral
vertebrae not shortened; 4) cervical co-
tylar arrangement type II; 7) omoster-
num cartilaginous, somewhat elongated,
manubrium large; 8) sacral diapophyses
round; 9) maxillary arch toothed; 10)
alary processes of premaxillae directed
dorsally, broad at base; 11) palatal shelf
of premaxilla relatively narrow; 12) fa-
cial lobe of maxilla moderately deep;
13) palatal shelf of maxilla narrow,
pterygoid process small; 14) maxillary

arch complete; 15) nasals relatively
small, narrowly separated medially; 16)
nasals not in contact with maxillae or
pterygoids; 17) nasals not in contact
with frontoparietals; 18) frontoparietal
fontanelle large; 19) frontoparietals not
ornamented; 22) epiotic eminences mod-
erately large; 23) cristae paroticae short,
stocky; 24) zygomatic ramus of squa-
mosal short; 25) otic ramus of squamosal
long, no otic plate; 26) squamosal-maxil-
lary angle about 40°; 27) columellae
present; 28) prevomers small, entire,
toothed, widely separated medially; 29)
palatines slender, widely separated me-
dially; 30) sphenethmoid not reaching
nasals; 31) anterior ramus of parasphe-
noid narrow, not keeled; 32) parasphe-
noid alae oriented at right angles to
anterior ramus, narrowly overlapped lat-
erally by median rami of pterygoids; 33)
pterygoids small, all rami slender, an-
terior rami long, reaching palatines; 34)
occipital condyles large, not stalked, nar-
rowly separated medially; 36) terminal
phalanges knobbed; 37) hyoid plate

Figure 112. Skull of Limnomedusa macroglossa. Dorsal, ventral, and lateral views. (KU 92960,

X 3.5).

176

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

lacking alary processes; 39) m. petro-
hyoideus anterior and m. sternohijoideus
insert on lateral edge of hyoid plate; 41 )
pupil vertical; 42) males with median
subgular vocal sac, nuptial asperities on
thumb; 43) body free of glands; 44)
tongue large, round; 45) toes fringed,
basally webbed, metatarsal tubercles not
enlarged, digital tips narrow; 46-48);
49) adults less than 60 mm. SVL; 50)
tympanum visible externally.

Composition. — Two species are rec-
ognized: macroglossa and misiones.

Distrihution. — Coastal lowlands of
southern Brasil, Uruguay, and adjacent
Misiones Province, Argentina. In Brasil,
the genus is found in Parana, Santa Cata-
rina, and Rio Grande do Sul.

Remarks. — Frogs of this genus bear
considerable external resemblance to
Leptodactylus but differ from it in hav-
ing vertical pupils, a nuptial pad on the
thumb of the male, and a broad sternal
style. Limnomedusa differs from all
other leptodactylines in having a type II
cervical cotylar arrangement.

Nothing is known of the breeding
biology of the frogs of this genus. The
presence of a nuptial pad on the thumb
suggests that clasping takes place in
water.

Hydrolaetare Gallardo, 1963

(Figs. 113-14)

Hydrolaetare Gallardo, 1963, Neotropica, 9:42
[Type-species by original designation, Lim-
nomedusa schmidti Cochran and Coin,
1959].

Diagnostic definition. — 1 ) sternum
bearing an elongate osseous style; 3)
transverse processes of posterior pre-
sacral vertebrae not shortened; 4) cer-
vical cotylar arrangement type I: 7)
omosternum large, elongate, cartilagi-
nous; 8) sacral diapophyses rounded; 9)
maxillary arch toothed; 10) alary proc-
esses of premaxillae directed posterodor-
sally, broad at base; 11) palatal shelf of
premaxilla broad; 12) facial lobe of max-
illa deep; 13) palatal shelf of maxilla
moderately wide, no pterygoid process;

Figure 113. Dorsal and lateral views of skull of
Hydrolaetare schmidti (KU 110613, X 3).

14) maxillary arch complete; 15) nasals
relatively large, in broad median contact;
16) nasals in broad contact with maxil-
lae, separated from pterygoids; 17) na-
sals in broad contact with frontopari-
etals; 18) frontoparietal fontanelle lack-
ing; 19) frontoparietal bearing sagittal
crest, slight exostosis; 22) epiotic emi-
nences large posteriorly; 23) cristae

Figure 114. Ventral view of skull of Hydro-
laetare schmidti (KU 110613, X 3).

LYNCH: LEPTODACTYLOID FROGS

177

paioticae narrow, relatively short; 24)
zygomatic ramus of squamosal elongate;
25) otic ramus of squamosal relatively
short, expanded medially into small otic
plate; 26) squamosal-maxillary angle
about 30°; 27) columella present; 28)
prevomers large, toothed, in median con-
tact; 29) palatines broad, separated me-
dially by anterior ramus of parasphe-
noid, bearing odontoid ridge; 30) sphen-
ethmoid extending anteriorly to middle
of nasals; 31) anterior ramus of para-
sphenoid narrow anteriorly, extending to
between palatines, not keeled medially;
32) parasphenoid alae deflected pos-
teriorly, broadly overlapped by median
rami of pterygoids; 33) pterygoids large,
anterior rami extending to middle of
orbit, prominent ventral flange present;
34) occipital condyles of moderate size,
not stalked, widely separated medially;
36) terminal phalanges knobbed; 37)
alary processes of hyoid plate wing-like;
39) m. petrolujoicleiis anterior and m.
sternohijoideus insert on lateral edge of
hyoid plate; 41) pupil vertical; 42) males
with median subgular vocal sac, no nup-
tial asperities; 43) body lacking glands;
44) tongue large, deeply notched pos-
teriorly; 45) toes fully webbed, metatar-
sal tubercles not enlarged, digital tips
narrow; 46-48); 49) adults large, known
specimens 80-105 mm. SVL; 50) tympa-
num visible externally.

Composition. — Monotypic.

Distribution. — A m a z o n i a n South
America.

Remarks. — Cochran and Coin (1959)
named Limnomedusa schmidti on the
basis of a single specimen collected near
Leticia, Colombia. They pointed out
that the new species was strikingly dif-
ferent from the other two species of
Limnomedusa. Gallardo (1963) reported
additional specimens of the species and
named a new genus for it (Hydrolaetare).
Hydrolaetare and Liimiotnedusa share
only two significant characters — vertical
pupils and an osseous element in the
sternum.

Several osteological characters of

Hydrolaetare suggest that the genus is
allied to the Grypiscini ( Telmatobiinae ) .
As in CyclorampJnis and Zachaenus, Hy-
drolaetare has a relatively deep maxilla
and quadratojugal, the nasals are rela-
tively large, in broad median contact,
and in broad contact with the maxillae,
the nasals are in broad contact with the
frontoparietals, the skull lacks a fonta-
nelle and has a sagittal crest, the zygo-
matic ramus of the squamosal is elon-
gate, and the pterygoid bears a ventral
flange. Unfortunately, reproductive data
are lacking for Hydrolaetare, but the
species lacks nuptial pads on the thumb,
suggesting that amplexus occurs in ter-
restrial situations (as in the Grypiscini).
The three genera now included in the
Grypiscini have large, cartilaginous ster-
na, in contrast to the leptodactyline ster-
num of Hydrolaetare. On the basis of
the skull alone, I would place Hydro-
laetare in the Grypiscini, but the sternal
apparatus and striking similarity be-
tween Hydrolaetare and Leptodactylus
{ocellattis and pentadactylus groups)
suggest that the genus is a leptodacty-
line. Hydrolaetare lacks one important
character of the Grypiscini; the otic
ramus of the squamosal of Hydrolaetare
is not medially curved to form a broad
otic plate. The otic plate of Hydrolae-
tare is small and like that seen in the
melanonotiis, ocellattis, and pentadacty-
lus groups of Leptodactylus.

Edalorhina Jimenez de la Espada, 1870
(Fig. 115)

Edalorhina Jimenez de la Espada, 1870, J. Sci.
Math. Phys. Nat., Lisboa, 3:58 [Type-
species by monotypy, Edalorhina perezi
Jimenez de la Espada, 1870].

Btihonias Cope, 1874, Proc. Acad. Nat. Sci.
Philadelphia, 26:124 [Type-species by
monotypy, Btibonias plicifrons Cope, 1874].

Diagnostic definition. — 1 ) sternum
bearing broad osseous style; 3) trans-
verse processes of posterior presacral
vertebrae not shortened; 4) cervical co-
tylar arrangement type I; 7) omoster-
num elongate, cartilaginous, manubrium
large; 8) sacral diapophyses rounded;

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 115. Dorsal and lateral views of skull
of Edalorhina perezi (KU 124225, X 6.4).

9) maxillary arch toothed; 10) alary
processes of premaxillae directed dorsal-
ly, moderately wide at base; 11) palatal
shelf of premaxilla of moderate width;
12) facial lobe of maxilla deep; 13) pala-
tal shelf of maxilla relatively narrow,
pterygoid process small; 14) maxillary
arch complete; 15) nasals large, in broad
median contact; 16) nasals not in con-
tact with maxillae or pterygoids; 17)
nasals not in contact with frontopari-
etals; 18) frontoparietal fontanelle lack-
ing; 19) frontoparietals bear large, ex-
ostosed, lateral crests; 22) epiotic emi-
nences obsolete; 23) cristae paroticae
short, stocky; 24) zygomatic ramus of
squamosal of moderate length; 25) otic
ramus of squamosal as long as zygomatic
ramus, no otic plate; 26) squamosal-
maxillary angle about 50°; 27) columella
present; 28) prevomers relatively small,
entire, toothed, separated medially, den-
tigerous processes lie posterior to choa-
nae; 29) palatines slender, separated me-
dially; 30) sphenethmoid short, extend-
ing anteriorly to posterior edge of na-
sals; 31) anterior ramus of parasphenoid
broad, relatively short, not keeled me-
dially; 32) parasphenoid alae oriented at

right angles to anterior ramus, narrowly
overlapped laterally by median rami of
pterygoids; 33) pterygoids slender, an-
terior rami long, nearly reaching pala-
tines; 34) occipital condyles small, not
stalked, widely separated medially; 36)
terminal phalanges knobbed; 37) alary
processes of hyoid plate on narrow
stalks; 39) in. petrolnjoideus anterior
and 7n. sternohijoideus insert on lateral
edge of hyoid plate; 41) pupil horizon-
tal; 42) males with median subgular
vocal sac, nuptial asperities on thumb;
43) inguinal glands present; 44) tongue
large; 45) toes lacking webbing, meta-
tarsal tubercles not enlarged, digital tips
narrow; 46-48) larvae aquatic; 49) adults
small, less than 45 mm. SVL; 50) tym-
panum visible externally.

Composition. — Two species are cur-
rently recognized: nasuta and perezi.

Distribution. — Amazonian lowlands
of Ecuador and northern and central
Peru, and in extreme western Brasil.

Remarks. — The genus was reviewed
by Dunn (1949), who combined the
nominal species huckleiji, perezi, and
plicifrons, but noted that the Peruvian
population of perezi usually lacks a snout
projection, whereas the Ecuadorian pop-
ulation has one. Dunn also pointed out
that Shreve's (1941) Edalorhina piistu-
lata (Pacific lowlands of Ecuador) was
not an Edalorhina but is closely related
to the Middle American Engijstomops
pustulosus. I concur with Dunn but in-
clude Engijstomops in Physalaemiis.

Parker (1927) and Noble (1931)
considered Edalorliina to be merely a
Plnjsalaemiis with cranial crests and
elongate papillae on the eyelids. Dunn
(1949) disagreed and suggested that
Edalorhina was more closely related to
Pletirodema. I consider the genus to be
intermediate between Litlwdytes and
Physalaemus. The breeding biology of
Edalorliina is unknown and could pro-
vide useful clues to the relationships of
the genus to the paludicoline leptodac-
tylids.

LYNCH: LEPTODACTYLOID FROGS

179

Lithodytes Fitzinger, 1843
(Figs. 116-17)

Lithodytes Fitzinger, 1843, Syst. Kept., p. 31
[Type-species by original designation, Rana
lineata Schneider, 1799].

Diagnostic definition. — 1 ) sternum
bearing an elongate osseous style; 3)
transverse processes of posterior pre-
sacral vertebrae somewhat shortened; 4)
cervical cotylar arrangement type I; 7)
omosternum bearing an elongate osseous
style and large cartilaginous manubrium;
8) sacral diapophyses rounded; 9) max-
illary arch toothed; 10) alary processes
of premaxillae directed dorsally, broad
at base; 11) palatal shelf of premaxilla
relatively broad; 12) facial lobe of max-
illa relatively deep, not exostosed; 13)
palatal shelf of maxilla relatively broad,
no pterygoid process; 14) maxillary arch
complete; 15) nasals large, in tenuous
median contact; 16) nasals not in contact
with maxillae or pterygoids; 17) nasals
not in contact with frontoparietals; 18)
frontoparietal fontanelle lacking; 19)
frontoparietals not ornamented; 22) epi-
otic eminences small; 23) cristae paroti-
cae broad, stocky; 24) zygomatic ramus
of squamosal relatively long; 25) otic
ramus of squamosal short, expanded me-

FiGURE 116. Dorsal and lateral views of skull
of Lithodytes lineatus (KU 104340, X 2.6).

Figure 117. Ventral view of skull of Lithodytes
lineatus (KU 104.340, x 2.6).

dially into small otic plate; 26) squa-
mosal-maxillary angle about 50°; 27)
columella present; 28) prevomers large,
entire, toothed, narrowly separated me-
dially; 29) palatines relatively narrow,
widely separated medially; 30) sphen-
ethmoid extending anteriorly to middle
of nasals; 31) anterior ramus of para-
sphenoid broad, relatively short, not
keeled medially; 32) parasphenoid alae
deflected posteriorly, short, not over-
lapped laterally by median rami of
pterygoids; 33) pterygoids slender, an-
terior rami long, not reaching palatines;
34) occipital condyles small, not stalked,
widely separated medially; 36) terminal
phalanges T-shaped; 37) alary processes
of hyoid plate on narrow stalks; 39) m.
petroJujoideus anterior and m. sterno-
hyoideus insert on lateral edge of hyoid
plate; 41 ) pupil horizontal; 42) male with
median subgular vocal sac, no nuptial
asperities; 43) body lacking glands; 44)
tongue large, rounded; 45) toes lacking
webbing, metatarsal tubercles not en-
larged, digital tips dilated, each pad
bearing terminal transverse groove; 46-
48); 49) adults medium-sized, to 50 mm.
SVL; 50) tympanum visible externally.

Composition. — Monotypic.

Distrihution. — Edge of the Amazon
Basin from Guayana to Bolivia.

Remarks. — S e v e r a 1 authors have
placed Lithodytes lineatus in Eleuthero-
dactijlus. Some did so following Noble
(1917), who ignored the presence of

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

osseous styles in the oniostenuim and
sternum and placed extra weight on the
presence of T-shaped terminal phalan-
ges. Ruthven (1919) effectively rejected
Noble's arguments.

Rana lineata Schneider has been fre-
quently confused with Hylodes lineatus
Brocchi. The latter is a Guatemalan spe-
cies of Eleiitherodactylus and bears no
resemblance to the Amazonian Litho-
chjtes lineatus. Litlwdytes lineatus bears
considerable superficial resemblance to
some species of the Eleutlierodactylus
jitzingeri group. Dunn ( 1931 ) named
Litlwdytes gaigeae (erroneously spelled
gaigei), a species found in Costa Rica
and Panama, and Piatt (1934) correctly
pointed out that gaigeae was a species of
Eleutherodactylus. The two species are
strikingly similar in color pattern.

The skull of Lithodytes bears con-
siderable resemblance to those of the
paludicoline genera, but the sternal style
is elongate, like that of Leptodactylus.
The hyolaryngeal apparatus of Litlw-
dytes is like that seen in Edalorkina,
Leptodactylus, and Pleurodema. Nothing
is known of the breeding biology of
Lithodytes, but the lack of nuptial asper-
ities suggests that the genus clasps on
land and may exhibit direct develop-
ment like the species of the Leptodac-
tylus marmoratus group.

Physalaemus Fitzinger, 1826

(Figs. 118-19)

Physalaemus Fitzinger, 1826, Neue Class. Rept.,
p. 39 [Type-species by monotypy, Physa-
laemus cuvieri Fitzinger, 1826].

Paludicola Wagler, 1830, Syst. Amph., p. 206
[Type-species by monotypy, Bujo albifrons
Spiz, 1824].

Gomphobates Reinhardt and Liitken, 1862, Vid.
Meddel. Naturh. Foren., 1861:172 [Type-
species by monotypy, Gomphobates notatus
Reinhardt and Liit]<en, 1862].

Eupemphix Steindachner, 1863, Sber. Akad.
Wiss. Wien, 48:188 [Type-species by mono-
typy, Eupemphix nattereri Steindachner,
1863].

Nattereria Steindachner, 1864, Verb. Zoob-Bot.
Gas. Wien, 1864:279 [Type-species by
monotypy, Nattereria lateristrigata Stein-
dachner, 1864].

lUobates Fitzinger in Steindachner, 1867, Reise
Novara, p. 12 [Listed as a generic synon>TO
(manuscript name) of Gomphobates fusco-
maculatus Steindachner; this generic name
is invahd and not available].

Engystomops Jimenez de la Espada, 1872, An.
Soc. Espanola Hist. Nat., 1:86 [Type-
species by monotypy, Engystomops petersi
Jimenez de la Espada, 1872].

Microphryne W. Peters, 1873, Mtber. k. Preuss.
Akad." Wiss. Berlin, 1873:616 [Type-species
by monotypy, Paludicola (Microphryne)
pustulosa Cope, 1864].

Peralaimos Jimenez de la Espada, 1875, Vert.
Viaje Pacif. Batr., p. 163 [Type-species by
monotypy, Bujo stentor Jimenez de la Es-
pada, 1872].

Diagnostic definition. — 1 ) sternum
bearing relatively broad osseous sternal
style; 3) transverse processes of posterior
presacral vertebrae somewhat shortened;
4) cervical cotylar arrangement type I;
7) omosternum elongate, cartilaginous,
manubrium small to large; 8) sacral
diapophyses slightly dilated; 9) maxil-
lary arch toothed or not; 10) alary proc-
esses of premaxillae directed dorsally or
slightly anterodorsally, relatively narrow
at base; 11) palatal shelf of premaxilla
relatively broad; 12) facial lobe of maxil-
la short, of moderate depth; 13) palatal
shelf of maxilla narrow, no pterygoid
process; 14) maxillary arch complete; 15)
nasals relatively large, in broad median
contact; 16) nasals not in contact with
maxillae or pterygoids; 17) nasals not in
contact with frontoparietals; 18) fronto-
parietal fontanelle lacking; 19) fronto-
parietals not ornamented; 22) epiotic
eminences small; 23) cristae paroticae
short, stocky; 24) zygomatic ramus of
squamosal short; 25) otic ramus of squa-
mosal short, no otic plate; 26) squa-
mosal-maxillary angle about 60°; 27)
columella present; 28) prevomers entire,
small, usually toothless, widely separated
medially; 29) palatines long, slender,
widely separated medially; 30) sphen-
ethmoid extending anteriorly beneath
posterior part of nasals; 31) anterior ra-
mus of parasphenoid narrow, not keeled
medially; 32) parasphenoid alae oriented
at right angles to anterior ramus, nar-
rowly separated from or narrowly over-

LYNCH: LEPTODACTYLOID FROGS

181

Figure 118. Lateral, dorsal, and ventral views of skull of Physalaemus pustulosus (KU 68271, X 4.7).

Figure 119. Lateral, dorsal, and ventral views of skull of Physalaemus ephippifer (KU 93005, X 4.7).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

lapped laterally by median rami of ptery-
goids; 33) pterygoids slender, rami long,
anterior rami reaching palatines; 34) oc-
cipital condyles small, on short stalks,
widely separated medially; 36) terminal
phalanges knobbed; 37) alary processes
of hyoid plate broad and wing-like; 39)
m. petrohyoideus anterior and m. sterno-
hijoideus insert on hyoid plate near mid-
line; 41) pupil horizontal; 42) males
with large external subgular vocal sacs,
tending to bilobe, males with nuptial
pads on thumb; 43) parotoid glands
present or absent, inguinal glands pres-
ent or absent, flank glands present or
absent; 44) tongue relatively narrow;
45) toes free of webbing and lateral
fringes, tarsus bearing tubercle on inner
edge or not, metatarsal tubercles en-
larged and spade-like or not, digital tips
narrow; 46) larvae with dextral vent,
2/3 tooth rows, labial papillae broadly
interrupted anteriorly; 48) eggs small,
numerous, laid in foam nest floating on
water; 49) adults range in size from 17-
60 mm. SVL; 50) tympanum usually
concealed, visible externally in pustu-
lattis.

Composition. — With the combination
of the nominal genera Engystomops, Eu-
pemplux, and Physahemiis into a single
genus, Physahemiis is one of the larger
genera of leptodactylids. I recognize 34
nominal species (aguirrei, albifrons, albo-
notatiis, harrioi, biligonigerus, centralis,
cicada, cuvieri, enesefae, ephippifer,
evangelistai, fernandezae, ftiscomacula-
tus, gracilis, henseli, kroeyeri, jordanen-
sis, macidiventris, moreirae, nanus, nat-
tereri, ohtectus, olfersi, paraenHs, peter-
si, pustulattis, pustulosus, riograndemis,
santafecinus, schereri, signiferus, soaresi,
stentor, and ternetzi).

Distribution. — Southern Mexico to
Argentina in lowland non-forested re-
gions (and through second growth and
occasionally primary forest) except for
the very arid Pacific lowlands south of
Ecuador and over most of central and
southern Argentina and Chile.

Remarks. — In a separate paper, I

(Lynch, 1970b) justified the combination
of Engystomops, Eupemphix, and PJiy-
salaemus. In that paper I suggested the
recognition of at least four species
groups — the petersi group, the maculi-
ventris group, the curvieri group, and
the fuscomaculatus group.

Physalaemus has the criniine pattern
of insertion of the hyoid musculature on
the hyoid plate. The only other Neo-
tropical leptodactyhds with this pattern
are the species of the Leptodactylus
marmoratus group and the genus Pseu-
dopaludicola, although the hyoid plate
of Hydrolaetare is like that seen in Phy-
salaemus and Pseudopaludicola. For the
present I consider Physalaemus and
Pseudopaludicola to be relatively closely
related but realize that the two genera
differ in many respects. In some char-
acteristics Physalaemus is closest to Lep-
todactylus and Pleurodema, but in others
it is closer to Edalorhina, Lithodytes,
and Paratelmatobius.

Paratelmatobius B. Lutz and Carvalho,

1958

(Fig. 120)

Paratelmatobius B. Lutz and Carvalho, 1958,
Mem. Inst. Oswaldo Cmz, .56:241 [Type-
species by original designation, Paratclrnato-
hius lutzi B. Lutz and Car\'alho, 1958].

Diagnostic definition. — 1 ) sternum
bearing a broad osseous plate; 3) trans-
verse processes of posterior presacral
vertebrae shortened; 4) cervical cotylar
arrangement type I; 7) omosternum
present, small, cartilaginous, manubrium
minute; 8) sacral diapophyses dilated;
9) maxillary arch toothed; 10) alary
processes of premaxillae directed dorsal-
ly, narrow at base; 11) palatal shelf of
premaxilla broad; 12) facial lobe of max-
illa shallow, expanded in snout region;
13) palatal shelf of maxilla broad, ptery-
goid process moderate-sized; 14) maxil-
lary arch complete; 15) nasals small,
narrow, separated medially; 16) nasals
not contacting maxillae or pterygoids,
nasal with elongate maxillary process
which nearly reaches maxilla; 17) nasals

LYNCH: LEPTODACTYLOID FROGS

183

not in contact with frontoparietals; 18)
frontoparietal fontanelle moderate-sized;
19) frontoparietals not ornamented; 22)
epiotic eminences well defined; 2-3) cris-
tae paroticae short, stocky; 24) zygo-
matic ramus of squamosal relatively
long, slender; 25) otic ramus of squa-
mosal short, curved medially to form
small otic plate; 26) squamosal-maxil-
lary angle about 55°; 27) columella
absent; 28) prevomers small, entire,
toothed, widely separated medially; 29)
palatines thin, elongate, broadly sep-
arated medially; 30) sphenethmoid ex-
tending anteriorly to middle of nasals;
31) anterior ramus of parasphenoid
broad, short, lacking median keel; 32)
parasphenoid alae short, narrow, de-
flected posteriorly, not overlapped later-
ally by median rami of pterygoids; 33)
pterygoids small, median rami short,
anterior rami not reaching palatines; 34 )
occipital condyles large, not stalked,
widely separated medially; 36) terminal
phalanges knobbed; 37) alary processes
of hyoid plate on narrow stalks; 39) m.
petrohyoideus anterior and m. sterno-

hyoideiis insert on lateral edge of hyoid
plate; 41) pupil horizontal; 42) males
lacking vocal sac, nuptial pads on first
two fingers; 43) body lacking glands;
44) tongue large, round; 45) toes fully
webbed, metatarsal tubercles not en-
larged, digital tips narrow; 46-48); 49)
adults small, less than 30 mm. SVL; 50)
tympanum concealed.

Composition. — Two species are pres-
ently known (Ititzi and gaigeae). The
latter was named P. pictiventris A. Lutz
in B. Lutz and Carvalho (1958) but is a
nomen nudum and an obligate synonym
of Leptodactylus gaigeae Cochran, 1938.

Distribution. — The coastal ranges in
Estado Rio de Janeiro, Brasil.

Remarks. — Aldopho Lutz collected
the first specimens of this genus in De-
cember 1931. He made water color
sketches of the two specimens and noted
that they represented a new species of
Faludicola or Eupemphix. However, he
never described the specimens or other-
wise published on them. Cochran
(1938) who received these two speci-
mens, named and described them as

Figure 120. Lateral and dorsal (KU 107089) and ventral (KU 92981) views of skulls of Paratelma-

obius littzi. All X 8.

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Leptodactylus gaigeae. She suggested
that the species was related to L. mar-
moratus and served as a generic link
between Leptodactylus and Zachaenus.
Lutz and Carvalho ( 1958 ) discovered a
new species allied to the frogs collected
by A. Lutz nearly 30 years before and
named it as a new genus and new spe-
cies, Paratehnatohius lutzi; at the same
time they published A. Lutz's figures of
the other species and used his manu-
script name, P. pictiventris, for them.
They suggested that Paratehnatohius
was intermediate between the endemic
southeastern Brasilian genus Cycloram-
phus and the Andean Tehnatohius.

The architecture of the otic ramus of
the squamosal of Paratehnatohius is
identical to that seen in the Grypiscini,
and the four genera bear considerable
external resemblance to one another.
The sternal plate in Paratehnatohius is
not like the sternal style seen in the lep-
todactylines but appears to be an ossifi-
cation of the sternal plate — an advance-
ment over the calcification of the same
element seen in old individuals of a
variety of leptodactylid genera and spe-
cies. Nevertheless, Paratelmatohius dif-
fers in several osteological characters
from the Grypiscini — the presence of a
frontoparietal fontanelle, the wide me-
dian separation of the nasals, the absence
of a ventral flange on the pterygoid.

In summary, Paratehnatohius is os-
teologically intermediate between the
Leptodactylinae and the Grypiscini
(Telniatobiinae). This might be re-
garded as some evidence for polyphyly
of the Telmatobiinae since one group of
Telmatobiinae (i.e., Alsodini) undoubt-
edly gave rise to the Leptodactylinae
and I am here pointing out the possibil-
ity of genetic relationship between Para-
tehnatohius and the Grypiscini. The
squamosal architecture of Paratehnato-
hius and the Grypiscini may be a parallel
(or convergent) development rather
than a result of relationship. The ap-
pearance of a very similar otic plate in
Megaelosia (Elosiinae) is very sugges-

tive that the appearance of this sort of
otic ramus is a labile feature and should
not be used in primary inferences of
relationships. In the Grypiscini, several
other osteological characters combine to
render this character confirmatory and
therefore it is used in the diagnosis of
that group (p. 135). I am tentatively
assigning Paratehnatohius to the Lepto-
dactylinae. In several respects the genus
bears some similarity to Physalaemus,
although I do not regard the relationship
(if any) to be close.

The presence of nuptial asperities
and a frontoparietal fontanelle, although
small, in Paratehnatohius suggest that
the genus is not allied with the Eleu-
therodactylini. The nature of the oc-
cipital condylar-cervical articulation as
well as a variety of other osteological
and external characters does not permit
its association with the Ceratophryinae,
Alsodini, Odontophrynini or Telmato-
biini. The Elosiinae is a compact group,
and the external and many internal fea-
tures serve to illustrate the lack of cor-
respondence between Paratehnatohius
and the Elosiinae. If a new family group
is not proposed for this small genus, then
the genus must belong to the Giypiscini
or the Leptodactylinae. I have fewer
difficulties associating it with the latter,
perhaps because the latter is a more
heterogeneous group. The presence of
an osseous plate in the sternum, al-
though it is rather unlike the sternal
style seen in the other genera of the
subfamily, is not contrary to the diag-
nostic feature of the subfamily. No other
leptodactylid known to me normally pos-
sesses an osseous post-zonal element.
Although the presence of an osseous
post-zonal sternal element is the only
uniform character in the subfamily, I
consider the subfamily to be monophy-
letic (see the generic account of Hydro-
laetare for further comment).

Pseudopaludicola Miranda-Ribeiro, 1926

(Fig. 121)

Pseudopahidicola Miranda-Ribeiro, 1926, Arch.
Mus. Nac. Rio de Janeiro, 27:152 [Type-

LYNCH: LEPTODACTYLOID FROGS

185

Figure 121. Lateral, dorsal, and ventral views of skull of Pseudopahidicola falcipes (KU 93068, X 8).

species by monotypy, Liuperus (sic) falcipes
Hensel, 1867].

Diagnostic definition. — 1) sternum
bearing an elongate, osseous or calcified
style; 3) transverse processes of posterior
presacral vertebrae shortened; 4) cervi-
cal cotylar arrangement type I; 7) omo-
sternum elongate, cartilaginous, manu-
brium small; 8) sacral diapophyses
rounded; 9) maxillary arch toothed; 10)
alary processes of premaxillae directed
dorsally, broad at base; 11) palatal shelf
of premaxilla broad; 12) facial lobe of
maxilla shallow; 13) palatal shelf of
maxilla narrow, no pterygoid process;
14) maxillary arch incomplete, quadrato-
jugal absent; 15) nasals small, widely
separated medially; 16) nasals not in
contact with maxillae or pterygoids; 17)
nasals not in contact with frontoparietals;
18) frontoparietal fontanelle absent,
frontoparietals usually narrowly sepa-
rated for entire length; 19) frontopari-
etals not ornamented; 22) epiotic emi-
nences obsolete; 23) cristae paroticae

very short, broad; 24) zygomatic ramus
of squamosal relatively short; 25) otic
ramus of squamosal long, expanded me-
dially into narrow otic plate; 26) squa-
mosal-maxillary angle 50-60°; 27) colu-
mella present; 28) prevomers small, re-
duced to sliver-like elements, dentiger-
ous rami lost, widely separated medially;
29) palatines narrow, sliver-like, in con-
tact with maxillae, widely separated me-
dially; 30) sphenethmoid very short, ex-
tending anteriorly to posterior edge of
nasals; 31) anterior ramus of parasphe-
noid narrow, not reaching palatines, not
keeled medially; 32) parasphenoid alae
long, oriented at right angles to anterior
ramus of parasphenoid, narrowly sepa-
rated from median rami of pterygoids;
33) pterygoids small, median and pos-
terior rami minute, anterior rami long,
reaching to palatines; 34) occipital
condyles small, stalked, widely sep-
arated medially; 36) terminal phalanges
knobbed; 37) alary processes of hyoid
plate wing-like; 39) m. petwhyoideus

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

anterior and m. sternohyoideus insert on
hyoid plate near midline; 41 ) pupil hori-
zontal; 42) males with bilobed subgular
vocal sac, nuptial asperities lacking; 43)
body lacking glands; 44) tongue large,
oval; 45) toes lacking webbing and lat-
eral fringes, metatarsal tubercles not en-
larged, digital tips narrow; 46) larvae
with dextral vent, 2/3 tooth rows, labial
papillae broadly interrupted anteriorly;
48) eggs small, numerous, laid singly or
in small clumps attached to submerged
vegetation; 49) males reach 16 mm., fe-
males reach 19 mm. SVL; 50) tympanum
concealed; 51 ) the antebrachial tubercles
are generically unique.

Composition. — Parker (1927) recog-
nized five species of the genus (ameg-
hini, boliviana, falcipes, pusilh, and sal-
tica). Bokermann ( 1966 ) recognized
five species in the coastal lowlands of
eastern and southern Brasil (ameghini,
falcipes, mystacalis, saltica, and ternetzi),
which Milstead (1963) had pronounced
identical. I consider boliviana and pusil-
la to be conspecific ( see "Remarks" ) and
recognize six species (ameghini, falcipes,
mystacalis, pusilh, saltica, and ternetzi).

Distribution. — The coastal lowlands
of Brasil from Bahia to northeastern
Argentina; Amazonian Bolivia, Paraguay,
and Venezuela, and in the coastal ranges
of Venezuela and the Santa Marta moun-
tains of Colombia.

Remarks. — The genus Pseudopaludi-
cola gained wide acceptance through
the work of Parker (1927) and Barrio
(1954). Parker included five species in
the genus and considered two of these
(boliviana and pusilla) to have T-shaped
terminal phalanges. All species of the
genus have elongate, knobbed terminal
phalanges and agree in all details of
skull ossification. Parker incorrectly
characterized the genus as having a
cartilaginous sternum (see account of
pectoral girdles, pp. 58-60) and has
been followed by all subsequent authors.
I include the genus in the Leptodacty-
linae and consider it closely related to
the paludicoline genera in spite of the

ethological differences pointed out by
Barrio ( 1954 ) . The structure of the
hyolarynx of Pseudopahidicola is not
greatly different from that of Physalae-
mus but very different from that of all
other Neotropical leptodactylids except
Ilydrolaetare and the Leptodactylus mar-
moratus group.

The species of this genus are readily
distinguished from all other small lepto-
dactylid frogs by their slender habitus,
lack of digital webbing, unexpanded
digital tips, concealed tympanum, and
the presence of an antebrachial tubercle.
The skeletons of the five nominal spe-
cies available to me are indistinguish-
able.

Parker separated boliviana and pu-
silla on slight differences in leg length
and coloration. I tentatively consider the
two nominal species identical because I
am unable to separate paratypes and
topotypic material of each from one
another. Rivero ( 1961 ) reported "pusil-
la" from Amazonian Venezuela and
Parker (1935) reported "Z?o/iu/ana" from
British Guiana and Paraguay. In view
of the ethological differences recently
discovered between the Brasilian species
(W. C. A. Bokermann, pers. comm.), it
might prove premature to combine the
cis-Andean populations of Pseudopaludi-
cola as a single species. The specimens
examined by me represent a single mor-
phological species. The Brasilian species
were pronounced conspecific by Mil-
stead ( 1963 ) . Bokermann has since
examined most of the types of this genus
and is familiar with all of the Brasilian
species in the field; he informs me that
most of the previously named kinds rep-
resent valid species, and there are yet
undescribed species living in the coastal
lowlands of southeastern Brasil. All of
the Brasilian species can be separated on
the basis of call, leg length, and color
pattern.

Leptodactylus Fitzinger, 1826

(Figs. 122-24)

Leptodactylus Fitzinger, 1826, Neue Class.
Rept., p. 38 [Type-species by subsequent

LYNCH: LEPTODACTYLOID FROGS

187

designation, (Fitzinger, 1843), Leptodac-
tijlus typhonia ( ^Rana typhonia Daudin,
1803, non Rana typhonia Linno, 1758).
Rami typhonia Daudin is R. sihilatrix Wied,
1824, which Heyer (1968) considered iden-
tical witli Rana fusca Schneider, 1799, for
which lie designated a neotype. However,
at least some of tlie syntypes of Rana fusca
are extant (W. C. A. Bokerniann, pers.
conim.), and study of these must be made
before Heyer's action can be accepted].

Cystignathiis Wagler, 1830, Syst. Amph., p. 202
[Type-species by present designation, Rana
mystacea Spix, 1824].

Gnathophysa Fitzinger, 1843, Syst. Rept., p. 31
[Type-species by original designation, Rana
hihyrinthica Spix, 1824].

Sihilatiix Fitzinger, 1843, Ibid., p. 31 [Type-
species by original designation, Cystigna-
thus gracilis Dumeril and Bibron, 1841].

Plectromantis W. Peters, 1862, Mtber. k. Preuss.
Akad. Wiss. Berlin, 1862:232 [Type-species
by monotypy, Plectromantis wagneri W.
Peters, 1862].

Adenomera Steindachner, 1867, Reise Novara,
Zool. Amph., p. 37 [Type-species by mono-
typy, Adenomera marmorata Steindachner,
1867].

Entomoglossus W. Peters, 1870, Mtber. k.
Preuss. Akad. Wiss. Berlin, 1870:647 [Type-

species by monotypy, Entomoglossus pustii-
latus W. Peters, 1870].

Pachypiis A. Lutz, 1930, Mem. Inst. Oswaldo
Cruz, 23:22 [Proposed as a subgeneric
name of Leptodactylus; no type-species was
designated. Preoccupied by Pachypus Bill-
berg, 1820 (Insecta: Coleoptera), Pachy-
pus d' Alton, 1840 (Mammalia), and Pachy-
pus Cambridge, 1873 ( Arachnida)].

Cavicola A. Lutz, 19.30, Ibid., 23:22 [Proposed
as a subgeneric name of Leptodactylus; no
type-species was designated. Preoccupied
by Cavicola Ancey, 1887 (Mollusca)].

Parvulus A. Lutz, 1930, Ibid., 23:22 [Proposed
as a subgeneric name in Leptodactylus;
type-species by subsequent designation
(Parker, 1932:342), Leptodactylus nanus
L. Muller, 1922].

Diagnostic definition. — 1) sternum
bearing an elongate osseous style; 3)
transverse processes of posterior presa-
cral vertebrae not shortened; 4) cervical
cotylar arrangement type I; 7) omo-
sternum large, elongate, cartilaginous,
manubrium large; 8) sacral diapophyses
rounded; 9) maxillary arch toothed,
teeth frequently pointed; 10) alary proc-

FiGURE 122. Lateral, dorsal, and ventral \iews of skull of Leptodactijlus quadriiittatus (KU 41030, X

4), a member of the fuscus group.

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

esses of premaxillac directed dorsally or
posterodorsally. broad at base; 11) pala-
tal shelf of premaxilla moderately broad;
12) facial lobe of maxilla relatively shal-
low, entire maxilla same depth anterior
to end of tooth row; 13) palatal shelf of
maxilla relatively narrow, no pterygoid
process; 14) maxillary arch complete;
15) nasals large, narrowly separated
medially; 16) nasals usually not in con-
tact with maxillae, never in contact with
pterygoids, nasals have elongate maxil-
lary processes in most species; 17) na-
sals not in contact with frontoparietals;

18) frontoparietal fontanelle lacking;

19) frontoparietals bearing some orna-
mentation posteriorly in old adults of
the larger species; 22) epiotic eminences
well defined posteriorly; 23) cristae
paroticae moderately long, somewhat
stocky; 24) zygomatic ramus of squa-
mosal somewhat expanded, relatively
short; 25) otic ramus of squamosal slight-
ly longer than zygomatic ramus, ex-
panded into narrow otic plate which
usually rests tenuously on crista paro-
tica; 26) squamosal-maxillary angle less
than 45°; 27) columella present; 28)
prevomers large, entire, toothed, nar-

FiGURE 123. Dorsal views of skull of (top)

Leptodactyltis pentadactyliis ( KU 68159, X D

and L. wagncri (KU 104389, X 1).

rowly separated medially; 29) palatines
broad, narrowly separated medially,
sometimes bearing odontoid ridge; 30)
sphenethmoid extending anteriorly to
middle of nasals in marmoraius, mela-
nonotus, oceUatiis, and pentadactyliis
groups, extending anteriorly to a point
anterior to nasals and usually anterior to
premaxillac in fuscus group; 31) anterior
ramus of parasphenoid narrow, not
keeled medially, reaching to palatines;
32) parasphenoid alae deflected pos-
teriorly, narrowly overlapped laterally
by median rami of pterygoids in fuscus,
melanonotus, oceUatus, and pentadac-
tyliis groups, separated in the marmora-
tiis group; 33) pterygoids slender, an-
terior rami reaching to middle of orbit;
34) occipital condyles moderate-sized,
not stalked, moderate to wide median
separation; 36) terminal phalanges
knobbed; 37) alary processes of hyoid
on narrow stalks in fuscus, melanonotus,
oceUatus, and pentadactyliis groups,
wing-like in mannoratiis group; 39) m.
petrohyoideus anterior and m. sterno-
hyoideus insert on lateral edges of hyoid
plate in fuscus, melanonotus, oceUatus,
and pentadactyliis groups, insert on
hyoid plate near midline in marmoratus
group; 41) pupil horizontal; 42) males
with median subgular or paired lateral
vocal sacs or none, males of fuscus and
marmoratus groups lack nuptial asperi-
ties, males of 7nelanonotus, oceUatus, and
pentadactyliis groups have spines on the
thumb, males of the pentadactyliis group
have spines on the chest and thumb;
43) body with diffuse venti'al and flank
glands or not, many species have glan-
dular folds on the dorsum, species of the
pentadactyliis group have inguinal
glands; 44) tongue large, with two long
posterior horns; 45) toes not webbed,
species of the melanonotus and oceUatus
groups have lateral fringes on the toes,
metatarsal tubercles not enlarged, digital
tips narrow, first finger slightly longer
than second in marmoratus group, much
longer than second in other species
groups except for a few species in the

LYNCH: LEPTODACTYLOID FROCS

189

melanonotus group; 46) frogs of mar-
moratus group do not have tadpoles, in
other groups larvae with median vent,
2/3 tooth rows, labial papillae broadly
interrupted anteriorly; 48) eggs laid in
foam nest floating on water in mela-
nonotus, oceUatus, and pentadactijJtis
groups, in these groups the eggs are
small and numerous; in the fiisciis group
eggs are deposited in a foam nest in an
underground burrow and hatch when
the nest is inundated; in the marmoratus
group the eggs are large, few in number,
and are deposited in an underground
nest in foam; development is direct in
the marmoratus group but the other
groups have aquatic larvae; 49) adults
range in size from 20-200 mm. SVL;
50) tympanum visible externally or
concealed.

Composition. — Gorham (1966) listed
60 species in the genus. Of these, three
belong to other genera (L. gaigeae^=
Paratelmatobius gaigeae; L. pulcherz=
Barycholos pulcher; L. tubercluosus:=
Ischnocnema quixensis). Heyer (1969a)

recognized only 32 species. His system
is admittedly conservative, but is a con-
siderable improvement over that pre-
sented by Gorham.

Distribution. — Middle American low-
lands from Sonora, Mexico, and southern
Texas to the Argentine Chaco and Gua-
yas region of Ecuador in South America,
and on the Lesser Antilles and Hispani-
ola. All localities for the species of the
genus are lowland (usually below 1200
meters ) .

Remarks. — No new generic synonyms
are added here. The generic synonymy
of Leptoclactylus has been stable for
many decades, because the genus is
rather morphologically uniform in ex-
ternal characteristics. Heyer ( 1969a )
solved many systematic problems of the
genus, among which was the discovery
of the identity of Plectromantis wagneri.
This species is a widespread species of
the Amazon Basin and has accumulated
nearly a dozen synonyms.

I studied the skeletons of 18 species
of the genus in formulating my concepts

Figure 124. Lateral, dorsal, and ventral views of skull of Leptodactylus hijlaedactyhis (KU 119387,

X5).

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

of the genus. This study illustrated the
remarkable osteological homogeneity of
the frogs of this genus. Accordingly, I
do not advocate the use of subgenera
such as those proposed by A. Lutz
( 1930 ) . Lutz placed the species of the
genus in six subgenera; of these, two are
generic homonyms, one is a valid generic
name of Asiatic ranids (Phityniantis),
and one (Plectromantis) was based on
erroneous data.

A major advancement in the system-
atics of this genus was provided by
Heyer, who divided the genus into five
species groups based on external mor-
phology, thigh musculature, jaw muscu-
lature, developmental patterns and tad-
pole morphology, some osteological
characters, and vocalization. Heyer util-
ized secondary sex characters in his
classification; his classification appears to
be a realistic one. I contend that his
meJononotus group is a composite with
part of the species (those with arched
prevomerine dentigerous processes of
the prevomerine bones) being members
of the oceUatus group.

Heyer advocated placing the mar-
moratus group in a separate subgenus,
Adenomera; this action is one of prefer-
ence, but in keeping with the criteria
followed throughout this review of the
family, I choose not to recognize the
subgenus. The frogs of the subgenus
Adenomera differ from those of the sub-
genus Leptodactijhis in that the former
exhibit direct development whereas the
latter have aquatic tadpoles. Heyer
cited additional characteristics to distin-
guish the marmoratus group from other
Leptodactyhis but these additional char-
acters are shared by some other species
groups. The musculature of the hyo-
larynx of the frogs of the marmoratus
group is like that seen in FhysaJaemus
and Pseudopahidicola (the criniine pat-
tern). The frogs of the marmoratus
group resemble Barycliolos pidclier (see
following account) in some but not all
morphological characters. When the
breeding biology of all of the genera of

the Leptodactylinae becomes known, the
generic status of Adenomera and Bary-
cholos should become more apparent.

Barycholos Heyer, 1969

(Fig. 125)

Banjcholos Heyer, 1969, Cont. Sci. Los Angeles
Co. Mus., 155:6 [Type-species by original
designation, Leptoclactijhis ptilcher Boulen-
ger, 1898].

Diagnostic definition. — 1) sternum
containing a calcified style, bifurcate
posteriorly; .3) transverse processes of
posterior presacral vertebrae slightly
shortened; 4) cervical cotylar arrange-
ment type I; 7) omosternum moderate-
sized, manubrium elongate, partly calci-
fied; 8) sacral diapophyses slightly di-
lated; 9) maxillary arch toothed; 10)
alary processes of premaxillae directed
dorsally, broad at base; 11) palatal shelf
of premaxilla relatively deep, deeply in-
cised; 12) facial lobe of maxillae rela-
tively shallow; 13) palatal shelf of maxil-
la broad anteriorly, narrowing posterior-
ly, no pterygoid process; 14) maxillary
arch complete; 15) nasals large, in broad
median contact; 16) nasals narrowly sep-
arated from maxillae, widely separated
from pterygoids; 17) nasals in tenuous
contact with frontoparietals; 18) fronto-
parietal fontanelle lacking; 19) fronto-
parietals not ornamented; 22) epiotic
eminences moderately well defined; 23)
cristae paroticae relatively short, stocky;
24) zygomatic ramus of squamosal rela-
tively short, broadly separated from
maxilla; 25) otic ramus of squamosal
relatively long, expanded medially to
form small otic plate; 26) squamosal-
maxillary angle 55°; 27) columella pres-
ent; 28) prevomers small, irregular in
outline, widely separated medially, bear-
ing large, arched, transverse, toothed,
dentigerous processes; 29) palatines
slender, lacking odontoid ridge; 30)
sphenethmoid entire, extending anterior-
ly beneath nasals; 31) anterior ramus of
parasphenoid narrow, not keeled, reach-
ing prevomerine dentigerous processes;
32) parasphenoid alae oriented at right

LYNCH: LEPTODACTYLOID FROGS

191

angles to anterior ramus, not overlapped
laterally by median rami of pterygoids;
33) pterygoids small, all rami slender,
anterior rami not reaching middle of
orbit; ;34) occipital condyles small, not
stalked, widely separated medially; 36)
terminal phalanges T-shaped; 37) alary
processes of hyoid plate small, on nar-
row stalks; 38-39); 41) pupil horizontal;
42) males lacking nuptial asperities,
vocal sac large, external, median, sub-
gular; 43) body lacking glands; 44)
tongue round, posterior edge free; 45)
toes lacking webs and lateral fringes,
outer metatarsal tubercle present, inner
metatarsal tubercle not enlarged, digital
tips dilated, no circumferential groove
on discs, first finger much longer than
second; 46-48); 49) adults small, about
25 mm. SVL; 50) tympanum visible
externally.

Composition. — Monotypic, B. ptil-
cher.

Distribution. — Pacific lowlands of
Ecuador (Heyer, 1969b).

Remarks. — In external appearance,
Barijcholos pulcher is simply a small
Leptodactijlus of the marmoratus group.
Heyer ( 1969b ) concluded that Barij-
cholos is not closely allied to Leptodac-
tylus or Lithodijtes, but is most closely
related to Eletitlierodactyhis. He did not

consider the relationship between Barij-
cholos and Eleutlierodactijlus to be close.
Barijcholos pulcher exhibits the fol-
lowing characteristics which are more
Eleutherodactijlus-\ike than Leptodacty-
lus-Mke: 11)," 15), 22), 32), and 36).
Barycholos more closely resembles Lep-
fodactylus in the following characteris-
tics: 1), 13), and 45). Heyer (1969b)
suggested that the life history of Bara-
chijlos is more like that of Eleutherodac-
tijlus than Leptodactyliis. Heyer exam-
ined a single adult female of Barycholos
pulcher which contained 43 ova about
2.8 mm. in diameter and concluded that
the species probably exhibits direct de-
velopment. I agree with Heyer on this
point, but do not agree that this charac-
ter, even in coincidence with T-shaped
terminal phalanges, indicates a closer
relationship to Eleutlierodactijlus than
to Leptodactijlus. Heyer ( 1969a ) char-
acterized the marmoratus group of Lep-
todactijlus in having "4-25 eggs per nest;
egg diameter 2.1-3.0 mm." In preserved
L. hylaedactijlus, females usually con-
tain about 20 eggs. The species of the
marmoratus group were assigned to the
subgenus Adenomera by Heyer ( 1969a,
1969b); the subgenus was in large part
defined on the basis of direct develop-
ment. As pointed out by Cochran ( 1955:
309), species of the marmoratus group

Figure 125. Dorsal and ventral views of skuU of Barycholos pulcher (UMMZ S-2881, X 5.7).

192

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

have enlarged digital pads and some in-
dication of T-shaped terminal phalanges.
The phalanges are intermediate between
the knobbed phalanges of the subgenus
Leptodactyhis and the distinctly T-
shaped phalanges of Eleiitlwrodactylus,
Heleopliryne, Litliodytes, SmintliiUus,
Syrrliopluis, Taudactyhis, and Tomodac-
tyltis. BarycJiolos more closely resem-
bles the subgenus Adenomera than Eleu-
therodactyJus. Direct development has
appeared several times in the course of
leptodactylid evolution (for example,
Crinia, the Eleutherodactylini, the sub-
genus Adenomera of Leptodactyhis, and
the cycloranine genera Kyarraniis and
Pluloria). The leptodactyline genera
LitJiodyfes and Paratelmatohius prob-
ably exhibit direct development.

The sternal style of Barycholos re-
sembles those of the leptodactyline gen-
era (except Paratelmatohius) and is very
different from the sternal apparatus seen
in non-leptodactyline leptodactylid gen-
era. Heyer ( 1969b ) described the style
as calcified in contrast to the osseous
style seen in Leptodactylus. Histolog-

ically, the calcified element is a precur-
sor to an osseous one, and the distinction
between a calcified and an osseous ele-
ment cannot be considered of primary
importance. The character of the sternal
apparatus which can be considered of
primary importance is its shape (style-
like or plate-like) because this character
is not age or size dependent. Because
the sternal style of Barycholos pulclier
is style-shaped, I consider Barycholos to
be a genus of the Leptodactylinae and
to not be closely related to Eleuthero-
dactylus.

The relationships of Barycholos with-
in the Leptodactylinae are not entirely
apparent, but I consider the genus most
closely related to the subgenus Adeno-
mera (Leptodactylus) and Lithodytes.
Osteologically, Barycholos differs from
the former in having larger nasals which
are in broad median contact and in hav-
ing smaller prevomers. Litljodytes has
large nasals which look like those of
Barycholos but the prevomers of Litho-
dytes are like those of the Leptodactylus
(Adenomera) marmoratus group.

PHYLOGENY AND RELATIONSHIPS

Ideally, a discussion of the phylogeny
of a group of organisms should be based
principally on their fossil record. In the
absence of such a chronicle of the evolu-
tion of the group, systematists turn to
study of the living representatives and
infer the phylogeny on the basis of
primitive and advanced characters.
Many fossils from Cretaceous through
Pleistocene horizons have been assigned
to the Leptodactylidae. Each of these is
discussed below; I consider several of
the fossils to represent frogs of families
other than the Leptodactylidae. The
leptodactylid fossils are too recent to be
of any significance in determining phy-
logeny within the family.

My interpretation of the phylogeny
of the Leptodactylidae is principally
based on comparisons of the character-
istics of pelobatid and leptodactylid
frogs. The phylogeny is in large measure

deduced from a study of evolutionary
trends in many characters. Most of these
evolutionary trends were noted by earlier
authors; the principle difficulty was de-
termination of the directions of the
trends.

THE FOSSIL RECORD

Pleistocene: — Giinther (1859b) re-
ported fossils of Ceratophrys aurita (as
C. cormita), Leptodactylus ocellatus,
L. pentadactylus, and Leptodactylus sp.
(as Cystignathus) from Lagoa Santa,
Minas Gerais, Brasil. Rusconi (1932)
named Ceratophrys ensenadensis from a
Pleistocene (Ensenadan) locality near
Buenos Aires, Argentina. Mecham
(1959) reported late Pleistocene cave
deposits in central Texas containing Hy-
lactophryne augusti (as Eleutherodacty-
lus latrans). Tihen (1960b) and Lynch

LYNCH: LEPTODACTYLOID FROGS

193

(1964) reported middle Pleistocene rec-
ords for Syrrhophus marnockii from
northern Texas. Auffenberg (1958)
named some specimens from late Pleis-
tocene cave deposits of Barbuda, British
Leeward Islands, as Hyla barbiidemis,
which Lynch (1966) placed in the genus
EleutJierodactylus.

Pliocene: — Ameghino (1899) named
Ceratophrys prisca from the Upper Plio-
cene of Monte Hermosa, Argentina.

Miocene: — C a s a m i q u e 1 a ( 1963 )
named Wawelia ger]}oldi from the Up-
per Miocene of Rio Negro, Argentina,
and described additional skeletal re-
mains of Caudiverbera catidiverbera (as
Gigantobatrachus parodii) from the
same locality and horizon. The type-
specimens of Gigantobatrachus were col-
lected from Miocene deposits in Santa
Cruz, Argentina ( Casamiquela, 1959).
Holman ( 1965 ) named Leptodactyhis
abavus and reported (1967) Eleuthero-
dactylus sp. from the Arikareean, Lower
Miocene, of northern Florida.

Oligocene: — Ameghino (1901) listed
Teracophrys (a nomen nudum) from the
Upper Oligocene of Patagonia. The
specimens are now apparently lost
(Schaeffer, 1949). Schaeffer (1949) re-
corded Caudiverbera caudiverbera (as a
new species, Calyptocephallela canque-
li), Eupsophus sp., and Neoprocoela
edentata from Lower Oligocene deposits
in Chubut, Argentina.

Eocene: — Schaeffer (1949) named
Caudiverbera casamayorervsis (as a new
genus, Eophractus) from the Lower Eo-
cene of Chubut, Argentina. Hecht
( 1960 ) named Eorubeta nevadensis
from the Lower Eocene of Nevada.
Noble (1930) concluded that Rana pu-
silla Owen, 1847, from the Eocene (In-
tertrappean) of peninsular India was a
myobatrachine leptodactylid and named
a new genus for it (Indobatrachus).

Cretaceous: — Estes (1964) reported
several bones from the Lance Formation
( Upper Cretaceous ) of Wyoming as pos-
sibly representing leptodactylid frogs;
he named none of these. Hecht (1960)

suggested that some of the frog fossils
from the Trinity Sands (Cretaceous) of
Texas were primitive leptodactyloids.

The fossil record is summarized in
Table 3. Some of these records require
special comment, because they are not
leptodactylid frogs. The following is a
systematic summary (by subfamily) of
the fossils I accept as members of the
family Leptodactylidae.

Ceratophryinae: — Upper Miocene to
Recent of South America. Wawelia ger-
holdi is not distinguishable from either
Ceratophrys or Lepidobatrachus but is
clearly a member of the subfamily (see
generic account, p. 112. Three spe-
cies of Ceratophrys are known as fos-
sils; two of these are extinct (C. ensena-
densis and C. prisca), the other species
still lives in the same area from which
the Pleistocene fossils were recovered.
Teracophrys may be a ceratophryine,
but its taxonomic status must await re-
discovery of Ameghino's specimens.

Cycloraninae: No fossil record.

Elosiinae: No fossil record.

Heleophryninae: No fossil record.

Leptodactylinae : Pleistocene to Re-
cent of South America. The three spe-
cies of Leptodactylus {oceUatus, penta-
dactylus and sp.) reported by Giinther
( 1859b ) are the only fossils known for
the subfamily.

Myobatrachinae: Eocene of penin-
sular India. Indobatrachus pusillus is
very similar to Crinia, and is probably a
leptodactylid. However, before the sys-
tematic position of Indobatraclms can be
fully evaluated, an osteological study
must be made of the Arthroleptinae
(Ranidae). See the generic account of
Imlobatrachus for further details (p.
103).

Telmatobiinae : Fossils of three tribes
are known.

Telmatobiini: Lower Eocene to Up-
per Miocene of Patagonia. Caudiverbera
is represented in Lower Eocene, Lower
Oligocene, and Upper Miocene deposits
of south-central Argentina. The Miocene
and Oligocene fossils are regarded as

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

TABLE 3. Geographic and temporal distribution of the fossil frogs currently considered

members of the Leptodactylidae.

NORTH AMERICA

SOUTH AMERICA

OLD WORLD

PLEISTOCENE

Eleutherodactyliis

harhudensis"
Hijlactophryne augiisti

Syrrhophus marnockii

Ceratophrys aurita

C. ensenadensis

Leptodactylus sp.
Leptodactyhis ocellatus
L. pentadactyhis

PLIOCENE

Ceratophrys prisca

MIOCENE

Leptodactylus ahavus
Eleutherodactyliis sp.

Wawelia gerholdi

Caudiverhera caudiverbera

OLIGOCENE

C caudiverbera
Eupsophus sp.
Neoprocoela edentata
? Teracophrys

EOCENE

Eorubeta
nevadensis

Caudiverbera
casamayorensis

Indobatrachus
pusillus

PALEOCENE

CRETACEOUS

[Lance Formation]
[Trinity Sands]

■■' West Indian: Barbuda, Leeward Islands.

identical with the Recent species ( Hecht,
1963), but the Lower Eocene fossils are
here recognized as specifically distinct.
Neoprocoela edentata, an ancestral stock
for Batrachophrynus, is known from the
Lower Oligocene of Patagonia. Tihen
( 1962b ) placed it in Bufo, but his action
is rejected here (see pp. 120-122).

Alsodini: Lower Oligocene of Pata-
gonia. Schaelfer's (1949) Eupsophus s]p.
is the only fossil record for this tribe.
Many specimens of the fossil are avail-
able but all are incomplete. Further
study of these fossils should be made to
assess their specific status. The fossil
species differs from all Recent species of
the genus in apparently having the nasal
bones in median contact.

Eleutherodactylini: Pleistocene of
Texas and Barbuda Island, Leeward
Islands. Three species of this tribe are

known as fossils. The West Indian Eleu-
therodactyliis barbudetisis may be extinct
or may be identical with E. martinicen-
sis (Schwartz, 1967). The Pleistocene
records of Syrrhophus marnockii occur
200 miles north of the present northern
limit of the range of the species ( Lynch,
1970a). The fossil of Hylactophryne
augusti is from a Late Pleistocene cave
deposit within the present geographic
range of the species.

I do not consider the other species
listed in Table 3 to be members of the
Leptodactylidae. The fossils here re-
moved from the family have been re-
ported from deposits in North America
(Lower Miocene to Upper Cretaceous).

Eleutherodactylus sp. : H o 1 m a n
( 1967 ) referred one right ilium from the
Thomas Farm Miocene beds of Gilchrest
County, Florida, to the genus Eleuthero-

LYNCH: LEPTODACTYLOID FROGS

195

dactijhis but because of limited compara-
tive material, did not make a specific
assignment. Many, if not most, mainland
Central and South American Eleuthero-
dactylus have distinct ilial crests in con-
trast to most West Indian species which
have the crest replaced by an ilial ridge.
Holman's fossil is similar to the ilia of
several West Indian Eleutherodactijlus
but is equally similar to some small
specimens of Acris which I have exam-
ined. Referral of the fossil to the Lepto-
dactylidae cannot be justified; it is equal-
ly likely that the fossil represents a
young individual of Acris, especially in
light of Holman's fossil species, A. bar-
houri, from the same locality.

Leptodactylus abavus Holman, 1965:
Leptodactylus abavus Holman (Lower
Miocene of Florida) is a species of Rana
and may not be separable from Rana
miocenica Holman of the same horizon
and locality. The reasons for placing the
species in Rana are presented in the ac-
count on pelvic girdles ( p. 63 ) .

Eorubeta nevadensis Hecht, 1960.
The fossils of this frog are preserved as
organic imprints in an oil well core. In
this condition, the fossils are badly
crushed and must be studied under
ultraviolet light. Hecht (1960) consid-
ered the presence of maxillary teeth and
long transverse processes of the posterior
presacral vertebrae as characters ade-
quate to associate the fossil with the
advanced frogs (Hylidae, Leptodactyli-
dae, and Ranidae ) . The Buf onidae were
not considered because the fossil has a
toothed maxilla. The sacral diapophyses
of the fossil are dilated and oriented at
right angles to the sagittal line; there-
fore, Hecht reasoned that the fossil did
not belong to the Ranidae.

Hecht then looked for hylid and lep-
todactylid frogs with long transverse
processes of the posterior presacral ver-
tebrae and dilated sacral diapophyses.
He characterized hylids as having either
somewhat shortened or very thin, needle-
like transverse processes of the posterior
presacral vertebrae. By process of elim-

ination, Hecht assigned the fossil to the
Leptodactylidae. Among the leptodac-
tylids available to him, he concluded
that the fossil was most similar to Mixo-
plujes and Lechriodus, because both
genera have dilated sacral diapophyses
and long transverse processes of the pos-
terior presacral vertebrae. Hecht stated
that Eorubeta differed from both of
these genera in having seven instead of
eight presacral vertebrae; Eorubeta was
further distinguished from Lechriodus,
because the latter has transverse proc-
esses on the atlas. The "atlas" (in the
sense of Hecht) of Lechriodus is the
fused cervical and second vertebrae.

Hecht rightfully complained of a
lack of skeletons of representative Re-
cent genera in museums and the lack of
comparative osteological studies of Re-
cent frog families. With these restric-
tions, Hecht's action in assigning the
fossil to the Australian section of the
Leptodactylidae seems capricious. My
study of Eorubeta was limited to the
description, remarks, and illustrations in
Hecht's (1960) paper.

Hecht's description is reasonably ac-
curate. However, I consider the fossil to
have eight presacral vertebrae (Hecht
recorded seven). Seven presacral verte-
brae are clearly evident and all of these
bear long transverse processes which are
as long as or only slightly shorter than
the sacral diapophyses. In all Recent
frog genera I have examined, the trans-
verse processes of the second vertebra
(first post-cervical) are invariably short-
er than those of the third vertebra. The
transverse processes of the third vertebra
are as long as, or longer than, those of
any other presacral vertebra, and are
usually somewhat curved. The trans-
verse processes of the right side of the
anterior presacral vertebrae of Eorubeta
are concealed beneath matrix except for
the leading edge of the vertebra I con-
sider to be the third (Hecht considered
this vertebra to be the second). The
remains of the skull overlie parts of the
vertebral column in this region. It is

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

clearly e\ident from an examination of
Hecht's figures, that the transverse proc-
esses of vertebra 2 are very long (as
long as the sacral diapophyses) and
slightly curved. If this vertebra is the
second as Hecht contends, then Eorubeta
has a vertebral column like no other in
the Anura. It is more reasonable to sug-
gest that this vertebra is the third. Just
anterior to the left transverse process of
this vertebra and the scapula, is a small
area of bone which Hecht tentatively
suggested is the coracoid. This structure
might be the left transverse process of
the second vertebra. Another interpre-
tation is possible for the bones Hecht
called the squamosals, occipital con-
dyles, and foramen magnum. If the "oc-
cipital condyles and foramen magnum"
are the centrum and neural arch of the
second vertebra, then the "squamosals"
are of the proper shape and length to
be considered the transverse processes of
the second vertebra. If this interpreta-
tion is correct, then the cervical is not
distinguishable from the bones of the
posterior part of the skull. The structure
designated "atlas" by Hecht is much too
small to be this element and is probably
not a complete bone. Pending the re-
covery of additional specimens, I will
not offer further interpretation of the
osteology of Eorubeta nevadensis.

I concur with Hecht that the fossil
does not represent the Ascaphidae, Dis-
coglossidae, Pipidae, or Rhinophrynidae,
for his reason (the transverse processes
of the posterior presacral vertebrae in
the Recent genera of these families are
very short and often knob-like). The
fossil has dilated sacral diapophyses
which are unlike those of any ranid
known to me. The Dendrobatidae and
leptodactylids of the subfamily Elosiinae
have non-dilated sacral diapophyses and
short transverse processes on all presa-
cral vertebrae. The pelobatids of the
subfamilies Pelobatinae and Pelodytinae
have very short transverse processes on
the posterior presacral vertebrae, as do
some Australo-Papuan leptodactylids,

and thus could not be closely related to
the fossil frog. The other frog families
have dilated sacral diapophyses in some
or all of the included genera ( Bufonidae,
Centrolenidae, Hylidae, Leptodactylidae,
Megophryinae, Microhylidae and Pseu-
didae). The only skeletal elements of
Eorubeta nevadensis which are useful in
comparisons are those of the vertebral
column, ilium, and maxilla. Eorubeta
does not closely resemble any genus in
the seven families listed above. Eoru-
beta has maxillary teeth and therefore is
probably not a bufonid, although this
character need not completely eliminate
the Bufonidae from consideration. The
ilium of Eorubeta has little indication of
a dorsal protuberance, prominence, or
ilial crest; this condition is seen in some
leptodactylids, megophryine pelobatids,
and microhylids. The ilium is not ex-
posed in lateral aspect and therefore
centrolenids and hylids cannot be elim-
inated from consideration. EoruI)eta is
not a bufonid, ceratophryine, telmato-
biine, leptodactyline, or pseudid because
the ilia of Eorubeta lack large ilial prom-
inences and/or dorsal crests.

The vertebral skeletons of the re-
maining groups ( Centrolenidae, Hylidae,
Cycloraninae, Heleophryninae, Myoba-
trachinae, Microhylidae, and Megophryi-
nae) are difficult to separate as units
when the details of the cervical vertebra
are unknown. The cervical cotyles are
widely separated medially in the Centi'o-
lenidae, Hylidae, Microhylidae, and
Myobatrachinae, whereas the cotyles are
narrowly separated in the Cycloraninae,
Heleophryninae, and Megophryinae.
The cervical vertebra is not distinguish-
able in Eorubeta. The only genera of
the seven family groups listed above
with the transverse processes of the pos-
terior presacral vertebrae as long as the
sacral diapophyses are Heinipliractus
and some Hyla (Hylidae), Batraclw-
phrijnus, Lechriodus, Limnodynastes,
and Mixophijes (Leptodactylidae, Tel-
matobiinae and Cycloraninae), and
Megophrys ( Pelobatidae, Megopliiyi-

LYNCH: LEPTODACTYLOID FROGS

197

nae ) . The transverse processes of Batra-
cJwphrynus (Fig. 79) are very similar
in bulk to those of Eorubeta, but the
sacral diapophyses of these two genera
are very different from one another.
Eorubeta is unique, insofar as I am
aware, in having wide, dilated sacral
diapophyses. The only comparable sa-
cral diapophyses known to me are those
of Atelopus and Rlunoderma.

In summary, Eorubeta does not
closely resemble the skeleton of any
known modern frog genus and cannot
be assigned to any presently recognized
family on the basis of its known mor-
phology. Hecht's (1960) assignment of
Eorubeta to the Leptodactylidae is not
defensible and probably in error. The
only reasonable systematic assignment
of the fossil is to "Family incertae sedis.
Order Salientia."

Estes ( 1964 ) described and figured
several Upper Cretaceous frog fossils
from the Lance Formation of Wyom-
ing.^'^ Among the fossils which are of
significance to this discussion are the
following: "Family PPelobatidae; Sub-
order Neobatrachia, Family incertae
sedis, near Hylidae?; and Family in-
certae sedis, near Leptodactylidae?"
(Estes, 1964: 57-61, figs. 30-32). The
coccyx described and figured by Estes
is unquestionably that of a megophryine
pelobatid, although a generic assignment
is not possible at present. The ilium de-
scribed and figured by Estes is not
sharply distinguishable from the ilia of
some species of Pelobates and Scaphio-
pus, but is different from the ilia of most
Megophryinae and Pelodytinae. The in-
complete left maxilla described and fig-
ured by Estes resembles those of Pelo-
bates, Scaphiopus, and, insofar as it is
known, that of Eopelobates.

Estes tentatively assigned an incom-
plete right squamosal to the Leptodac-
tylidae. The squamosal does not resem-
ble that of any extant leptodactylid

'■'' Estes (1970) associated a number of these Late
Cretaceous elements with the genus Eopelobates
(Pelobatidae, Megophryinae).

genus but is similar to the squamosals
of Scaphiopus (ScapJiiopus) and some
Pelobates. Estes remarked that the fossil
has an "opisthotic articulation surface
[which] resembles that of leptodactyl-
ids." This cryptic statement implies that
there is (or are) a characteristic opis-
thotic articulation of the squamosal in
leptodactylids; the statement is not in
agreement with my observations. A
complete spectrum of opisthotic articula-
tions can be demonstrated within the
Leptodactylidae, ranging from species
lacking the dorsal portion of the squa-
mosal (Notaden) to those with the squa-
mosal enclosing much of the crista paro-
tica (Caudiverbera, Ceratophrijs). Estes
(1969) subsequently assigned this squa-
mosal as well as numerous other Late
Cretaceous and Paleocene bones to
Scotiophryne pustulosa ( Discoglossi-
dae). Further comment on this action
is deferred to a separate paper. The
two bones (squamosal, figured; and na-
sal) which Estes (1964) suggested
might be from hylids or leptodactylids,
could equally confidently be assigned to
the Pelobatidae.

The pelobatid centrum from the
Middle Eocene of Wyoming figured by
Hecht {in McGrew, 1959) is clearly that
of a megophryine pelobatid, and as
Hecht pointed out, is very similar to
that of Eopelobates grandis from the
Lower Oligocene of South Dakota.
Hecht's fossil is probably generically
identical with Eopelobates grandis and
may not be specifically distinct. Estes'
megophryine coccyx is possibly repre-
sentative of the same complex.

Hecht (1960:13) referred to some
fossil frogs from the Trinity Sands of
Texas (early Cretaceous) as "a primi-
tive leptodactylid or some close relative."
Until he publishes on them, no further
comment will be made, although it
should be born in mind that Hecht con-
sidered Eorubeta a definite leptodac-
tylid. Hecht and Estes (1960) named a
Jurassic frog as Comobatraclius and
placed it in Reig's (1958) Neobati-achia.

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MISCELLANEOUS PUBLICATION MUSEUxM OF NATURAL HISTORY

They further suggested that the fossil
not be assigned to any recognized family
but that "on the basis of probability [no
confidence limits are given] a leptodac-
tyloid affinity appears more likely [than
a microhylid or hyperoliid affinity]."

The fossil record for the Leptodac-
tylidae can be summarized as follows:
fossils of several stocks are known from
the Tertiary of southern South America;
an Eocene frog from peninsular India
seems to be a myobatrachine leptodac-
tylid; and leptodactylids are not known
elsewhere in the world until the Pleisto-
cene of the West Indies and Texas.

With the exception of Neoprocoela
and Indobatrachus, the fossil record of
the Leptodactylidae is of little use in de-
termining the phylogeny and is of limited
value in discussing zoogeography.

PELOBATID-LEPTODACTYLID
RELATIONSHIPS

Some Papuan leptodactylid frogs
have been confused with pelobatid frogs.
Lechriodus was erroneously believed to
be a pelobatid until Noble ( 1924 ) dem-
onstrated that the pectoral and thigh
musculature were bufonoid, not pelo-
batoid. In general, however, the two
families have always been regarded as
being very different from one another.
This distinction hinged largely upon a
distinction between the Pelobatinae
(Pelohates and Scaphioptis) and the Neo-
tropical leptodactylids. In the former,
the coccyx is fused to the sacral verte-
bra, whereas in leptodactylids the two
bones are separate.

The Pelobatidae may be separated
from the Leptodactylidae by the greater
dilation of the sacral diapophyses in the
former, the sacral-coccygeal articulation
(fused or with a single condyle in for-
mer, double condyle in latter), mid-
dorsal cricoid gap in former, single slip
to m. depressor mandibulae (pars scapu-
laris) in former, and m. semitendinosus
and m. sartorius not separate in former.
Two or three subfamilies are recognized
depending on the author: Pelobatinae,

Megophryinae, and sometimes Pelody-
tinae.

The Pelobatidae are unquestionably
the more primitive group (Griffiths,
1963; Inger, 1967; Kluge and Karris,
1969; Noble, 1922, 1931; and Tihen,
1965), but the distinction between the
two families is not so great as has been
previously befieved, principally because
previous authors have tended to ignore
the similarities between the Megophryi-
nae and the Austi-alo-Papuan leptodac-
tylids and to stress those features which
distinguish the two most abundant and
best known groups of the two families
(the Pelobatinae and Neotropical lepto-
dactylids ) .

The amplectic position of the male in
the Pelobatidae, Cycloraninae and Myo-
batrachinae is inguinal in contrast to the
axillary amplexus in almost all advanced
frogs. The Megophryinae, Myobatra-
chinae and Cycloraninae (part) have
free intervertebral discs, a character that
has been regarded as paedomorphic or
specialized by most authors but which is
more likely primitive (Tihen, 1965).
Several other characteristics, some here-
tofore regarded as of little significance,
occur in the primitive families and in
some leptodactylids and sporadically,
though rarely, in advanced frogs [e.g.,
absence of an outer metatarsal tubercle;
vertical pupil; large, juxtaposed occipital
condyles; small transverse processes on
the posterior presacral vertebrae; imbri-
cate neural arches; and tadpoles with
complete row(s) of labial papillae and
high number of anterior tooth rows
(3-6)].

Griffiths ( 1963 ) contended that ecto-
chordal and stegochordal centra were
primitive to holochordal centra; Inger
( 1967 ) argued that holochordy is prob-
ably primitive since it occurs in most
extinct lepospondylous amphibians and
that therefore ectochordy and stego-
chordy are derived. Ectochordy has
been termed paedomorphic; if so, then
any distinction between frog families
based on the nature of the coccygeal-

LYNCH: LEPTODACTYLOID FROGS

199

sacral articulation cannot be seriously
considered in interpreting the macro-
systematic evolution of frogs. A full
range of variation occurs in the Pelo-
batidae — the ectochordal megophryines
exhibit a monocondylar articulation, the
stegochordal pelobatines exhibit a coc-
cygeal-sacral fusion, and the presumably
stegochordal pelodytines exhibit a bi-
condylar articulation. Imbricate neural
arches occur in all pelobatids and are
found in cycloranines (but not myo-
batrachines ) , in heleophrynines, in cera-
tophryines, and in some telmatobiines
( Odontophrynini and Telmatobiini ) .
The degree of dilation of the sacral dia-
pophyses in pelobatids exceeds that seen
in any leptodactylid except Neoprocoela
(Lower Oligocene, Patagonia). Bufo-
nids, which are generally conceded to
be leptodactylid derivatives, have broad-
ly dilated sacral diapophyses as well. If
Neoprocoela is properly assigned famil-
ially, then all primitive leptodactylids
may have had sacral dilations of the
pelobatid degree — those living leptodac-
tylids with sacral dilations do not ap-
proach the condition seen in bufonids or
pelobatids. The sacral vertebra of Pelo-
dtjtes is more like that of pipids than
other pelobatids.

Hecht (1960) mentioned the lack of
moderate to long transverse processes on
the presacral vertebrae in primitive
frogs. My examination of skeletons of
all ascaphid, discoglossid, pipid, and
rhinophrynid genera confirms his obser-
vation. Among the nine living and three
extinct pelobatid genera some variation
occurs. The transverse processes of the
posterior presacral vertebrae are little
more than bosses or are very short in
Macropehhates, Pelobates and Scaphio-
pus. Miopelodijtes and Felodytes have
short processes strongly sloped anteriorly
as in pipids. In the Megophnjs and
Eopelobates (Megophryinae), the trans-
verse processes of the posterior presacral
vertebrae are of moderate length (Zwei-
fel, 1956a; personal observation) but
they are shortened and directed strongly

anteriorly in at least one species of Lep-
tobrachium (Boulenger, 1908), recalling
the condition seen in Felodytes. The an-
terior presacral vertebrae of all pelo-
batids have elongate transverse proc-
esses ( relative to the widths of the sacral
diapophyses) as well as in some primi-
tive leptodactylids ( Ceratophryinae ) .
This same condition appears in some
bufonids and is variable within Bufo.
Several groups of leptodactylids have
relatively short transverse processes on
the posterior presacral vertebrae. This
is most pronounced in Neobatrachus and
Notaden (Fig. 30) but is also found in
several Neotropical groups (e.g., Telma-
tobiini, Odontophrynini, and Grypiscini)
as well as in Heleophryne and most
Myobatrachinae and Cycloraninae.

The presence of postzygapophyses on
the sacrum and prezygapophyses on the
coccyx must be considered primitive.
The zygapophyses are not present in
Felodytes, and their presence is obliter-
ated by sacral-coccygeal fusion in the
Pelobatinae; they are found in the Mego-
phryinae but not consistently in any
leptodactylid. McDowell (in litt.) ob-
served sacral postzygapophyses and coc-
cygeal prezygapophyses in Metacrinia
and in the enigmatic Sooglossus, but I
have seen them only in Batrachophryntis
and Megophnjs. Zweifel (1956a) de-
scribed coccygeal zygapophyses in Eo-
pelobates.

The tadpoles of pelobatids and lepto-
dactylids, as well as of all advanced frogs
(except the microhylids whose relation-
ships are obscure), are Type IV of Orton
(1953). Admittedly, use of gross tad-
pole morphology as a basis of macro-
systematic ( interfamilial ) classification
is hazardous (e.g., Inger, 1967), but an
examination of intrafamilial variation can
be useful in determining intrafamilial
evolutionary trends. Inger (1967) char-
acterized the Pelobatidae and Leptodac-
tylidae as having tadpoles with median
vents. Orton (1952) characterized the
pelobatid tadpoles as follows: vent me-
dian, beak present, tooth rows usually

200

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

divided with one complete short row
anteriorly and two complete rows pos-
teriorly, and labial papillae complete ex-
cept for a narrow median interruption
anteriorly. The data for pelobatids, in-
sofar as is known, are summarized in
Table 4.

Only half of the pelobatid genera
(and species) have median vents; the
temperate Himalayan genera have dex-
tral vents as does the subtropical and
tropical Leptobrachitim. Inger's (1966:
25) statement that a complete row of
papillae across the upper lip is charac-
teristic of pelobatids is in error. Insofar
as I am aware, it is true for only L.
gracilis and L. pelodyfoides, although
some Oreolalax have large, widely scat-
tered papillae across the upper lip (Liu,
1950 ) . The uppermost tooth row is com-
plete in all pelobatids (if teeth are
present) and is very short in the Pelo-
batinae and Megophryinae. In Pelodytes
this row is almost as wide as the mouth,
recalling the condition seen in most tad-
poles. The Pelobatinae and Pelodytinae
differ from the Megophryinae in having
two complete tooth rows across the fully
papillate lower lip; only a single com-
plete row is found in megophryines.
Most tooth rows are divided medially by
the upper beak in pelobatids. This fea-

ture in combination with the usually
high tooth formula characterizes most
pelobatids. This is not to say that this
condition is not duplicated elsewhere;
for example, Neohatrachtis has a dental
formula of 1:2-2/1-1:11 to 1:3-3/1-1:11,
Mixophijes has a 11:4-4/1-1:11 with three
other very short lateral divided rows pos-
teriorly and Heleophnjne has a IV/1-1:
XII to IV/1-1: XVI formula; most bufo-
noids exhibit a 1:1-1/111 tooth formula.

The tadpoles of pelobatids suggest a
closer relationship between the Euro-
pean and North American genera than
either group has to the Megophryinae.
However, the variation within the Mego-
phryinae suggests that the tadpole can
be a hazardous source for definitive
statements about relationships. Among
bufonoid frogs the high tooth formulae
in cycloranine and heleophrynine lepto-
dactylids is suggestive of the pelobatid
condition in distinction to the relatively
dissimilar mouthparts seen in other bufo-
noid frogs. Tadpole morphology repre-
sents perhaps one of the most useful
and most misused of the available char-
acter complexes to be used in frog clas-
sification. A major problem to be over-
come is the descriptive formulae applied
to the mouth parts. Many authors use
the most simple, least informative system

TABLE 4. Characteristics of Recent pelobatid genera— tadpoles"

Labial

papillae Tooth formula
Vent complete

Genus median anteriorly ( Low - High )

Scaphiopus" X O n:l-l/2-2:II-I:4-4/4-4:n

Pelobates X O 1:3-3/3-3:11

Pelodytes X O 1:3-3/3-3:11

Nesobia ? ? ?

Megophrys X A" 0

Leptobrachium O V 0 -1:7-7/5-5:1

Oreolalax O O 1:4-4/4-4:1-1:7-7/7-7:1

Scutiger O O 1:3-3/4-4:1-1:6-6/6-6:1

Vibrissaphora O O 1:5-5/4-4:1

"Data from Boulenger (1897), Inger (1966), Liu (1950), Orton (1952), Pope ( 1931), and Stebbins

(1951).
'' Including Spea.
'' Absent.

LYNCH: LEPTODACTYLOID FROGS

201

of arable numerals separated by a solidus
(i.e., 2/3) for tadpoles with all of the
rows complete or some of them divided.
Using various systems, the dental for-
mula for Vibrissaphora liui could be
presented as:

(A) 6/5

1

(C) 1:5-5/4-4:1

(D) IC, ID, 4L/1L, 3D, IC

The formula with the greatest informa-
tion content is the last ( D ) , in which
the arable numeral preceding the letters
C, D, and L refers to the number of
complete, divided (but not lateral), and
lateral (and divided) rows of teeth re-
spectively. Lateral rows are separated
by the beak. This system differs from
that of Liu (1950) in distinguishing be-
tween divided and lateral rows and does
not require the use of Roman numerals.
Liu's system is preferable to the 4-
layered formula (B) in requiring less
space on a printed page. The four-
layered formula likewise does not dis-
tinguish divided from strictly lateral
rows although this distinction can be
demonstrated to be two ends of a con-
tinuum.

There can be little argument against
the statement that the Pelobatidae is the
most primitive of the frog families, ex-
cept the archaic Ascaphidae, Discoglos-
sidae, Pipidae, and Rhinophrynidae (Ti-
hen, 1965 ) . An appropriate test of which
of the advanced families is most closely
related to the Pelobatidae would be to
compare them relative to the number of
primitive characters shared. The selec-
tion of characters and determination of
evolutionary direction was made on the
following bases: 1) characters shared
between the Pelobatidae and some or all
of the archaic frog families were re-
garded as unquestionably ancestral; and
2) those characteristics which occur in
these archaic families but also occm* in

some of the bufonoid families were se-
lected as being useful in measuring the
relative primitiveness of each of the bu-
fonoid families and subfamilies (and
tribes of the leptodactylid subfamily
Telmatobiinae ) .

The familv or subfamilv with the
lowest sum of values is judged to be
most primitive, and the higher the sum
of values, the greater is the divergence
of that group from the ancestral stock.
These primitive characters are ( I ) large,
closely approximated occipital condyles
— Types II or III; (II) imbricate neural
arches; (III) anterior presacral vertebral
transverse processes elongate and pos-
terior presacral vertebral transverse proc-
esses shortened; (IV) diapophyses of
sacral vertebrae broadly dilated; (V)
post-zygapophyses on sacral vertebra
and prezygapophyses and /or transverse
processes present on anterior end of
coccyx; (VI) intervertebral discs free;
(VII) ilium lacking dorsal crest, ilial
prominence or protuberance; (VIII) all
skull bones present; maxillaries, premax-
illaries, and prevomers toothed; (IX)
phalangeal formulae 2-2-3-3 and 2-2-3-
4-3; (X) pupil vertical; (XI) outer meta-
tarsal tubercle absent; (XII) amplexus
inguinal; (XIII) eggs small, laid in
water, tadpole free-living; (XIV) tad-
pole vent median; (XV) tadpole dental
formula including at least three or four
rows above and three rows below beak;
and (XVI) pectoral girdle arciferal.
Each of these 16 characters or character
complexes was assigned a value from 0
to 2 for each of the 23 bufonoid and
pelobatid taxa. The character states and
values are summarized in Table 5 and
the scores and sums for 35 family groups
of "non-archaic" frogs in Table 6. A
value of 0 indicates that the primitive
condition is uniform within the group; a
value of 1 indicates the group exhibits
an intermediate condition or is variable
with more than one-half of the included
taxa exhibiting the primitive state; and a
value of 2 indicates that a majority of or
all of the included taxa share the (or a)

202

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

derived state for the character. The sum
of the values for the 16 characters is an
index of how far removed a given taxon
is from the basal stock of the Bufonoidea
— the Pelobatoidea. All three pelobatid
subfamilies exhibit low values — the
Megophryinae with 2, the Pelobatinae
with 3, and the Pelodytinae with 4. The
following leptodactylid groups exhibit
values below 14 — Heleophryninae, Cy-
cloraninae, Ceratophryinae, and Telma-
tobiini ( Telmatobiinae ) . The Heleo-
phryninae are only slightly removed
from the pelobatid zone and are con-
siderably more primitive than any other
group of the "higher frogs." The Cyclo-
raninae are well removed from the pelo-
batid zone but less than one-half as far
removed as are the Myobatrachinae.
The leptodactylids of southern South
America (Ceratophryinae and telmato-
biine and alsodine Telmatobiinae) are
more primitive than any other bufonoid
groups (except the heleophrynine and
cycloranine leptodactylids ) . Of the non-
leptodactylid bufonoid families and sub-
famihes, the Phyllomedusinae are least
unlike the pelobatoid ancestor. The Bu-
fonidae, which are usually regarded as
only slightly advanced over the Lepto-
dactylidae, are the next most primitive
group.

TABLE 5. Sixteen phylogenetically sig-
nificant characters, their character states
and values.

I. Cervical type II =: 0
Cervical type I := 2

II. Neural arches imbricate == 0
Neural arches open = 2

III. Transverse processes of anterior
presacral vertebrae expanded,
those of posterior presacral verte-
brae shortened = 0

All transverse processes as long as
sacral diapophyses or all transverse
processes shorter than sacral di-
apophyses = 2

IV. Sacral diapophyses dilated = 0
Sacral diapophyses rounded = 2

V. Sacral vertebra bearing postzyg-
apopyses and/ or coccyx bearing
prezygapophyses = 0
Sacral vertebra lacking postzyg-
apophyses and coccyx lacking pre-
zygapophyses =: 2

VI. Intervertebral discs free = 0

Intervertebral disc fused to cen-
trum ^ 2

VII. Ilium without dorsal crest or dorsal
protuberance = 0
Ilium with dorsal crest and/ or dor-
sal protuberance ^ 2

VIII. All skull bones present, premaxillae
and maxillae bearing teeth ^= 0

IX. Phalangeal formulae normal = 0
Some skull bones lost and /or pre-
maxillae and maxillae toothless = 2

Intercalary element present or pha-
langes lost =z 2

X. Pupil vertical = 0

Pupil horizontal or round =^ 2

XI. No outer metatarsal tubercle = 0

Outer metatarsal tubercle present
= 2

XII. Amplectic position inquinal = 0
Amplectic position axillary = 2

XIII. Eggs small, usually pigmented, lar-
vae aquatic == 0

Eggs large, development abbrevi-
ated or direct = 2

XIV. Tadpole vent median = 0
Tadpole vent dextral =^ 2

XV. Larval tooth formula at least 3/3 =
0

Larval tooth formula less than

3/3 = 2

XVI. Pectoral girdle arciferal = 0

Pectoral girdle firmisternal = 2

LYNCH: LEPTODACTYLOID FROGS 203

The Pelobatidac are intermediate be- and ranoid) frogs. The leptodactyhds

tween the primitive ( discoglossoid and are the stem bufonoids and are difficult

pipoid) and the advanced (bufonoid to separate from the pelobatids.

TABLE 6. Vakies for each of the sixteen phylogenetically significant characters in

non-archaic frog groups.

Pelobatidae

Megophryinae 0000001000000100 2

Pelobatinae 0000020100000000 3

Pelodytinae 0000220000000000 4

Leptodactylidae

Heleophryninae 00122100000P0000 6

Cycloraninae 0011211001000110 9

Myobatrachinae 2211101102200220 17

Ceratophryinae 0002221001120010 12

Tehiiatobiini 0011221001120020 13

Alsodini 1111221002210020 16

Odontophrynini 1022221002220020 18

LeptodactyHnae 2122222002220120 22

Elosiinae 2222222102220220 25

Grypiscini 2222222002221220 25

Eleutherodactylini . 2222222002222220 26

Bufonidae 0121222202220121 22

Rhinodermatidae 0221222202222022 25

Centrolenidae 2221222122221020 25

Hyhdae

Phyllomedusinae 2020222010221120 19

other HyHdae 2222222112222220 28

Dendrobatidae 2222222202221122 28

Pseudidae 2222222022222220 28

Sooglossidae 222200??020?2022 16+

Ranidae

Raninae 2222222002220202 24

Rhacophorinae 2222222022121202 26

Petropedetinae 2222222002020202 22

Hemisinae ???222?220020202 16+

Hyperohidae

Astylosterninae 2222222021020202 23

Hyperohinae 2222222020020202 22

Arthroleptinae 2222222022222022 28

Microhyhdae

Dyscophinae ?0?0220001020022 11+

Brevicipinae ?0?022?202202022 16+

Asterophryinae 2020220202022022 20

MicrohyHnae 2000221102020022 16

CophyHnae 2220220002022022 20

Phrynomerinae 2220221222020022 23

204

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

INTRAFAMILIAL RELATIONSHIPS
OF THE LEPTODACTYLIDAEi^

Figure 126 reveals that the Lepto-
dactvHdae exhibit the greatest range of
intrafamilial variabiUty or diversity
among the Pelobatoidea, Bufonoidea,
and Ranoidea. In part, this diversity
represents a finer degree of knowledge
about the Leptodaetylidae than about
some other families, but the diversity is
also real. The Leptodaetylidae have been
the convenient "catch-all" for genera of
bufonoid frogs with obscure relation-
ships, and can be defined as "those bufo-
noid frogs that are not members of the
Bufonidae, Centrolenidae, Dendrobati-
dae, Hylidae, Pseudidae, or Rhinoder-
matidae." The intrafamilial relationships
of the Leptodaetylidae are schematically
summarized in Fig. 127.

The Leptodaetylidae can be viewed
as a series of increasingly more special-
ized subfaiuilies and tribes which bridge
the morphological and behavioral gaps
between the Pelobatidae and the smaller
families of the Bufonoidea and the Ra-

" Elsewhere (Lynch, MS) I have proposed unit-
ing the Old World leptodactyHd subfamilies as
the family Myobatrachidae and using Lepto-
dactyhdae for the four Neotropical subfamilies.
As I noted in the paper to be published in
"Evolutionary Biology of the Anura," the Neo-
tropical subfamilies form a well-defined group
which is approached by only one Old World
subfamily, the Cycloraninae. The Myobatrachi-
nae form a rather isolated group from all other
leptodactyloids and could be placed in a family
of their own (the Sooglossidae is probably not
separable from Myobatrachinae ) . The cyclo-
ranine-heleophrynine relationship is somewhat
tenuous and apparently not very close, but these
two groups seem to be somewhat more closely
related to each other than either is to the
myobatrachines or the Neotropical complex.

Using the Leptodaetylidae as a single family
for all seven subfamilies is only a slightly greater
abstraction than to place the Old World sub-
families in one family ( Myobatrachidae ) and
the Neotropical subfamilies in a second (Lepto-
daetylidae). Cladistically more sound would be
to use three familes, one for the Myobatrachinae,
a second for the Cycloraninae and Heleophry-
ninae, and a third for the Neotropical families.
If the latter fragmentation is employed, use of
the term leptodactyloid becomes appropriate.

noidea. The least specialized subfamilies
and tribes of leptodactylids occur in
southern Africa and the Australo-Papuan
region. Other, only slightly more special-
ized, groups occur in temperate South
America, and the very specialized groups
occur in the subtropical and tropical
zones of South America and Middle
America.

The subfamily Heleophryninae con-
tains one genus and is restricted in dis-
tribution to southern Africa. HeJeophryne
is the most primitive leptodactylid genus
in terms of its degree of divergence
from the pelobatids. In some characters,
Heleophnjne resembles some of the gen-
era of Australo-Papuan leptodactylids
(Heleioporus, NeohatracJuis, Notoden,
and Mixophyes) of the subfamily Cyclo-
raninae, but the Australo-Papuan genera
are closely interrelated, and none shows
any evidence of a close relationship with
Heleophnjne. The characteristics shared
by HeleopJiryne, some of the Cyclo-
raninae, the Ceratophryinae, and the
Telmatobiini reflect those of the ancestral
stock(s) of the Leptodaetylidae and may
be used to demonstrate a close relation-
ship among these eleven genera; how-
ever, with the exception of Heleophryne
and the ceratophryines, the other genera
are the primitive members of larger
groups. Heleophryne may represent an
independent line of pelobatid derivatives
which has achieved the leptodactylid or
bufonoid grade.

The Myobatrachinae are a relatively
compact group of Australo-Papuan gen-
era with one Eocene fossil genus from
peninsular India. Although sympatric
with the Cycloraninae, the Myobatra-
chinae are not closely related to the
Cycloraninae. Other than the character-
istics of leptodactylid frogs, the two sub-
families have one unifying character
(inguinal amplexus), but this is shared
with ascaphids, discoglossids, pipids,
rhinophrynids, pelobatids, and Batra-
chyla (Telmatobiinae, Leptodaetylidae).
I consider it likely that two other genera
of leptodactylids will be found to exhibit
inguinal <x\w\Ae\\xs— Heleophryne and

LYNCH: LEPTODACTYLOID FROGS

205

DENDROBATIDAE •

HYPEROL

/

Figure 126. Comparison of the relative primitiveness of 35 groups of non-archaic frogs including the
23 bufonoid and pelobatid subfamilies and tribes. The concentric semicircles are guidelines and are
not intended to have a special significance. Families are enclosed in dashed lines. The abbreviations
used are as follows: A = Alsodini, Ce = Ceratophryinae, Cy = Cycloraninae, Ele = Eleutherodacty-
lini, Elo = Elosiinae, Gr = Grypiscini, He = Heleophryninae, Le = Leptodactylinae, Mego = Mego-
phryinae, My = Myobatrachinae, Pb = Pelobatinae, Pd = Pelodytinae, Phy =: Phylloniedusinae, and
T = Telmatobiini. The separation of taxa l^y degrees of arc is not intended to reflect relationships.

Thoropa. The tadpole mouthparts of the
Myobatrachinae are iinHke those of all
other leptodactylids but are very similar
to those of the bufonids. It must be
stressed that the Myobatrachinae and
Bufonidae share few other characters and
that the similarity in tadpole mouthparts
may well be convergent or parallel. The
species of most myobatrachine genera
are toothless, but the species of some
genera have teeth on the maxillary arch.
The myobatrachines have type I cervical
cotylar arrangement and widely sepa-
rated occipital condyles in contrast to
the type II cervical cotylar arrangement
and narrowly separated occipital con-
dyles of bufonids. Further study must be
made to evaluate the closeness of the
relationship between these two groups,
although among the extant leptodacty-
lids, I consider the mvobatrachines as

least likely ancestors of the nearly cos-
mopolitan Bufonidae.

The Cycloraninae are restricted to the
Australo-Papuan region, as are the Myo-
batrachinae. Parker (1940) suggested
that with further study, the two sub-
families would be shown to be less dis-
tinctive than he had characterized them.
However, the additional osteological and
tadpole characters utilized here have
amplified the distinction between these
two subfamilies. As pointed out above,
the two Australo-Papuan subfamilies
share only one significant character. I
consider the community of ancestry of
these two groups to be very ancient if it
existed at all. The Cycloraninae share
some characters with the Telmatobiini,
Alsodini, and Ceratophryinae, but are not
closely related to any of these groups.
The tribe Cycloranini is less unlike the

206

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

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Figure 127. Dendrogram illustratinG; proposed relationships of the leptodactylid subfamilies and

tribes and derived families. The hatched zone is Cretaceous time during which decreasing equability

occurred. The vertical scale is not intended to indicate the duration or age of any group.

Neotropical subfamilies than is the Lim-
nodynastini which are more specialized
than the Cycloranini.

I consider each of the three subfam-
ilies discussed above to have evolved
independently from a megophryine an-

cestor. One could therefore argue that
the Leptodactylidae are polyphyletic.
The Cycloraninae appear to be ancestral
to the Neotropical leptodactylids, but the
Heleophryninae and Myobatrachinae are
apparently not involved in the phylogeny

LYNCH: LEPTODACTYLOID FROGS

207

of the Neotropical leptodactylids. As
stressed above, the Megophryinae and
Aiistralo-Papuan leptodactyHds are simi-
lar in most character complexes. Taken
in combination, the Cycloraninae, Heleo-
phryninae, and Myobatrachinae fomi an
evolutionaiy grade between the Mego-
phryinae and the Neotropical leptodacty-
lids. However, I am reluctant to accord
both the Heleophryninae and Myobatra-
chinae familial status and consider
equally unrealistic the idea of consider-
ing either or both subfamilies of the
Pelobatidae. The Myobatrachinae could
be more reasonably distinguished famili-
ally from the Pelobatidae than could the
Heleophryninae. From an evolutionary
standpoint, the Heleophryninae are a rel-
ict of the Megophryine stock which gave
rise to the Cycloraninae (see below, pp.
208-9). The Myobatrachinae probably
evolved from a contemporaneous mego-
phryine. Whether this megophryine was
subfamilially distinct from the group
which gave rise to the Cycloraninae can-
not be known in the absence of fossils. I
doubt that these two megophryines were
subfamilially distinct and therefore con-
sider the Leptodactylidae monophyletic
following the reasoning of Simpson
(1961:124).

The subfamilies discussed below seem
to have a common ancestry within the
Leptodactylidae. This common ancestor
was probably a cycloranine which was
not unlike the modern Cycloranini.

The subfamily Ceratophryinae con-
tains only two extant genera and is a
morphologically isolated group. This iso-
lation has been described by several
workers who consider the subfamily to
be a family more closely allied to the
Bufonidae than to the Leptodactylidae
(Cei, Limeses, Reig). In spite of the
morphological isolation of the Cerato-
phryinae there are some striking simi-
larities between the Ceratophryinae and
the Odontophrynini. Boulenger (1882),
Cochran ( 1955), Reig ( 1960b), Reig and
Cei ( 1963), and Reig and Limeses (1963)
suggested that Odontophrynus and Stom-
bus auctorum ( =^Proceratophrys) are very

closely related to the ceratophryines. The
Odontophrynini may have been derived
from a ceratophryine ancestor, but the ex-
tant odontophrynine genera exhibit fewer
primitive characters than do the genera
of the Telmatobiini. The Odontophrynini
represent either the first or second di-
vergence from the original stock of the
Telmatobiinae. The Ceratophryinae seem
to represent the earliest divergence from
the Neotropical leptodactylid stock.

The remainder of the early Neotrop-
ical leptodactylid stock (after the Cerato-
phryinae diverged ) is represented by the
Telmatobiinae and derived subfamilies
(Elosiinae and Leptodactylinae). Some
of the terrestrial genera of the Telmato-
biini resemble the primitive Australo-
Papuan Cycloranini (Cycloraninae) but
more closely resemble the other tribes of
Telmatobiinae.

The Leptodactylinae are derived from
the relatively primitive Alsodini (Eii-
psophiis). The most primitive lepto-
dactyline (Pleurodema) is very similar to
Eupsophus. The two genera differ in the
sternal apparatus, breeding biology, and
loss of the quadratojugal. Pleurodema
has an osseous sternal style (as do all
other leptodactylines) and lays its eggs in
a foam nest (like several other lepto-
dactylines); these two characters clearly
ally Pleurodema with the Leptodactyli-
nae although its close relationship to
Eupsophus is obvious and could be used
to support the argument that the Lepto-
dactylinae are only a tribe of the Tel-
matobiinae.

The Elosiinae are a small, morpho-
logically homogeneous group except for
the cranial adaptations of Megaelosia.
The relationships of the subfamily to
other leptodactylid groups are unclear
and over-shadowed by the relationship
between the Elosiinae and the Dendro-
batidae. The Elosiinae exhibit many
characters in common with the Alsodini,
Grypiscini, and Eleutherodactylini, and
I consider the elosiines to represent an
early division from the alsodine stock
which later gave rise to the Grypiscini
and Eleutherodactylini.

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

I recognize two tribes in the Cy-
cloraiiinae, the Cycloranini and tlie Lim-
nodynastini. Of the two tribes, the
Cycloranini are more primitive. The
Limnodynastini are speciaHzed in their
breeding biology. None of the five lim-
nodynastine genera has vertical pupils
and all have free intervertebral discs, the
cervical fused to the second vertebrae,
and relatively long transverse processes
of the posterior presacral vertebrae.
Adelotus brevis and one species of Lim-
nodynastes have outer metatarsal tuber-
cles, which are otherwise lacking in the
tribe. The foam-nesting habits and the
modifications of the fingers in females of
the Limnodynastini are considered ade-
quate reasons for separating these five
genera from the Cycloranini which do
not lay their eggs in foam nests (except
Heleioporus) and do not have finger
fringes in the females. The foam-nesting
habit of Heleioporus is quite different
from that of the Limnodynastini but sim-
ilar to that exhibited by the frogs of the
Leptodactylus fiisciis group. Heleio-
porus, Neohatrachus, and Notaden are
closely related. Although their skeletons
are similar, the three genera differ in
breeding biology and some external char-
acters. Cyclorana and Mixophyes differ
from Heleioporus, Neohatrachus, and
Notaden in many features of the skull
and vertebral column, but do not closely
resemble each other. They are derived
genera but probably have evolved several
characters in a parallel fashion (ankylosis
of intervertebral discs to centra, separa-
tion of cervical and second vertebrae in
adults, long transverse processes on the
posterior presacral vertebrae, and lack of
a frontoparietal fontanelle). The last
character may be primitive to the pres-
ence of a fontanelle, but in the lepto-
dactylid groups the primitive genera fre-
quently have a frontoparietal fontanelle.

The Ceratophryinae and Heleophry-
ninae do not exhibit a great amount of
intrasu1)familial variability in that they
are small groups. The Myobatrachinae
are morphologically homogeneous in
many characters but are heterogeneous

in many others. With the possible ex-
ception of bufonid and ranoid derivatives
(see below), the myobatrachines do not
figure prominently in bufonoid evolution.
With respect to the Leptodactydae,
the Myobatrachinae are an evolutionary
dead-end.

The Ceratophryinae and Telmato-
biinae apparently evolved from cyclo-
ranine ancestors. The Ceratophryinae
represent an early divergence of the Neo-
tropical stock and are a morphologically
isolated and small group. The Telmato-
biinae are a morphologically diverse and
large group with one fossil and 24 Recent
genera. I divide the Telmatobiinae into
five tribes. The Telmatobiini (5 genera)
and Odontophrynini (2 genera) are the
most primitive tribes and the Alsodini
are but slightly more advanced. The
former two tribes have apparently not
given rise to additional groups and have
the bulk of their species in temperate or
Andean South America. The Odonto-
phrynini share several characteristics
with the Ceratophryinae and may be
early derivatives of that group which
have paralleled the Telmatobiini. At
present, I include the odontophrynines
in the Telmatobiinae because they lack
the distinctive vertebral column and
vertebral shield of the ceratophryines.
Cei ( 1965 ) demonstrated that the skin
proteins of Ceratophrys and Lepidoha-
trachus set these genera off from the
other leptodactylids including Odonto-
phrynus and Proceratophrys. The dis-
tinctive ilia of the ceratophryines are
identical with those of the odontophry-
nines and unlike those of all other lepto-
dactylids. The solution of the problem of
whether the odontophrynines represent
a proto-ceratophryine or a telmatobiine
stock will probably require some fossil
data.

The Alsodini and their derivatives
make up the majority of the Neotropical
Leptodactylidae. The Alsodini occur in
temperate and Andean South America,
with one genus (Thoropa) endemic to
southeastern Brasil. The tribe is some-
what heterogeneous in that two of the

LYNCH: LEPTODACTYLOID FROGS

209

genera (Eupsophus and Hylorina) have
a type II cervical cotylar arrangement
and lay relatively numerous small eggs
in aquatic situations. Both of these gen-
era engage in axillary amplexus and
have outer metatarsal tubercles. One of
them (Hylorimi) has vertical pupils. The
other two alsodine genera (Batrachijla
and Thoropa) have a type I cervical
cotylar arrangement and lay relatively
few large eggs in moist terrestrial situ-
ations. The larvae of both genera be-
come aquatic after the nest is inundated.
Both genera have outer metatarsal tu-
bercles and horizontal pupils. Batrachijla
engages in inguinal amplexus (Barrio,
1967a), but amplectic behavior has not
been observed for Thoropa. As men-
tioned above, the Leptodactylinae seem
to have been evolved from an ancestral
stock with the characteristics of Eu-
psophus. Batrachijla and Thoropa are
probably representative of the alsodine
stock which gave rise to the Eleuthero-
dactylini, Grypiscini, and Elosiinae. I
consider the Elosiinae to represent an
early divergence from this stock because
the Elosiinae have aquatic larvae. Os-
teologically, the Elosiinae are isolated
from the Grypiscini and Eleutherodacty-
lini. I consider the Grypiscini more
primitive than the Eleutherodactylini.
The eggs of grypiscine genera are large
and deposited in moist terrestrial situ-
ations as is the case in the Alsodini, but
in contrast to the Alsodini, the larvae are
not aquatic ( Lutz, 1944 ) . The eleuthero-
dactyline genera exhibit direct develop-
ment. Unlike the grypiscine larvae,
eleutherodactyline larvae never free
themselves from the enclosing egg enve-
lopes. The Grypiscini make up a small
group which is restricted in distribution
to the coastal ranges of southeastern and
southern Brasil. The Eleutherodactylini
range over the entire tropical and sub-
tropical zones of the Americas except in
the arid regions of Central America,
Ecuador, Peru, and Venezuela. The ze-
nith of the Leptodactylidae is Eleuthero-
dactylus which contains probably 400
species and occurs over most of the range

of the Eleutherodactylini. Direct devel-
opment is probably the single adaptive
change made by this tribe which per-
mitted the tribe to evolve into such a
large group. The success of the genus
Eleutherodactijlus is measured by its di-
versity and adaptability. The genus is
rich in species throughout the West In-
dies and occurs in semiarid as well as
veiy moist habitats. Unlike many lepto-
dactylid genera, it is not restricted to
lowland situations where there are ponds
(a requirement for species with aquatic
larvae) but ranges altitudinally to at least
4200 meters in the Andes.

EXTRAFAMILIAL RELATIONSHIPS
OF THE LEPTODACTYLIDAE

I have discussed the relationships be-
tween the Pelobatidae and Leptodacty-
lidae and the relationships within the
Leptodactylidae above. In some cases,
reference was made to the close relation-
ship between a leptodactylid group and
the frogs of other families. I consider the
close relationship between the Pelobat-
idae and Leptodactylidae to be estab-
lished, and also consider the statement
"the Leptodactylidae are the stem bufo-
noid group" to be established. If the
Leptodactylidae are the stem bufonoid
group, then all other bufonoid families
are leptodactylid derivatives or are inde-
pendently derived from a pelobatid
stock. The works of Griffiths (1963),
Inger (1967), Kluge and Farris (1969),
Noble (1922, 1931), and Tihen (1965)
have clearly established that the archaic
frog families (Ascaphidae, Discoglossi-
dae, Pipidae, and Rhinophrynidae ) did
not directly give rise to the "advanced
frogs."

These authors agree that these four
families are clearly primitive and that the
other frog families are advanced. Most
consider the Pelobatidae to bridge the
gap between "primitive" and "advanced"
frogs. There is considerable debate as to
whether the Microhylidae are primitive
or advanced frogs. The proponents of
the primitive position (Hecht, 1963, Inger,

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

1967, Orton, 1957, and Starrett, 1968)
based much of their argument on the
tadpole moiphology, whereas the advo-
cates of the advanced, ranoid position
(Griffiths, 1963, Noble, 1922, 1931, and
Parker, 1934) rely on the pectoral archi-
tecture, thigh musculature, and the vari-
able sacral-presacral central articulation.
Further discussion of the relationships of
the Microhylidae is withheld pending
new data and perhaps another mono-
graph of the family. I have plotted values
for some of the microhylid subfamilies in
the figure illustrating the degree of prim-
itiveness in frog groups ( Fig. 126). The
subfamily Microhylinae appears to be
the most primitive subfamily of this
group on the basis of the characters I
used. Relatively little data of the breed-
ing biologies of microhylids are available,
and I have seen relatively few genera in
computing my values of primitiveness for
the family. In general the values are in
agreement with the idea of the phylogeny
of microhylids advanced by Parker ( 1934)
—that the group represents an early ranid
divergence. The data are not in accord
with the position taken by Hecht (1963)
and Starrett (1968)— that the family rep-
resents one of the archaic families. The
remaining frog families seem to have
some relationship to the varied lepto-
dactylid stocks.

I consider the rhinodermatids to rep-
resent a Neotropical bufonid derivative
and therefore include that genus in a
discussion of the relationships of the
Bufonidae to the Leptodactylidae. As
mentioned above, among the extant lep-
todactylid groups the Myobatrachinae
are most like the Bufonidae and are pre-
sumably the modern representatives of
the proto-bufonid stock. The two groups
agree strikingly in the structure of the
tadpole mouthparts. All bufonids (and
Rhinoderrmi) lack teeth; few leptodactyl-
ids are edentate, but this character-state
is most pronounced in the Myobatra-
chinae. All bufonids and myobatra-
chines have dilated sacral diapophyses.
Myobatrachines have a type I cervical
cotylar arrangement (type II in bufo-

nids), free intervertebral discs (pro-
coelous vertebrae in bufonids), and lack
Bidder's organ (present in bufonids).
Most bufonids lack a prezonal element
in the pectoral girdle ( omosternum pres-
ent in Nectophnjnoides and at least one
Btifo, B. haematiticus), whereas most lep-
todactylids have a large omosternum and
manubrium. The prezonal element in
myobatrachines is small in most genera
and absent in some. Although the Neo-
tropical bufonids have radiated (7 en-
demic genera ) , there is no close relation-
ship between the Neotropical bufonids
and leptodactylids.

The relationships of the Centrolenidae
are not apparent. Before the significance
of the intercalary cartilage was accepted,
they were often placed in the Hylidae or
Leptodactylidae. Centrolenids are arbo-
real, have aquatic tadpoles, intercalary
cartilages, T-shaped tenninal phalanges,
the astragulus and calcaneum fused, and
lack a prezonal element in the pectoral
girdle. Most authors consider them hylid
derivatives (Eaton, 1958). Much of the
argument that centrolenids are hylid de-
rivatives rests on a conviction that all
Neotropical taxa with intercalary carti-
lages are related. The hyolarynx of Cen-
troleneUa is distinctive (figured by Eaton,
1958) and quite different from that of
most bufonoid genera. The variation in
the hyolarynx of hylids has not been
adequately investigated and until it has,
the taxonomic value of the distinctive
hyolaiynx of CentroleneUa remains un-
known.

The Dendrobatidae are an elosiine
leptodactylid derivative and not ranoid
as claimed by Griffiths ( 1959, 1963). This
point was discussed in greater detail in
the section of the Elosiinae (pp. 163-4).
Griffiths ( 1963 ) , in arguing in support of
his contention that the Dendrobatidae
are a subfamily of the Ranidae, cited
the apparent parallelisms in the Petro-
pedetinae ( African ranids ) . My study of
the group is limited to some dissection
and examination of cleared and stained
individuals but results in the conclusion
that the two groups exhibit considerable

LYNCH: LEPTODACTYLOID FROGS

211

similarity in myology and osteology. The
similarities are quite striking and prob-
ably reflect a community of ancestry
rather than parallelism. However, it
should be borne in mind that I have not
studied the other ranid subfamilies and
genera in detail and cannot therefore
convincingly argue that the similarities
are not due to parallelism.

The relationships of the Hylidae are
not apparent, but the family is usually
tacitly considered a leptodactylid de-
rivative. Inger's (1967: Fig. 6) phylogeny
suggests that the Hylidae are the sister
group ( sensii Hennig ) of the Ranidae +
Rhacophoridae. The suggestion is unic(ue
and mentally provocative but needs fur-
ther investigation. Previous authors were
committed to placing the Hylidae and
Ranidae in different suborders because of
convictions that the pectoral architecture
or sacral centrum were characteristic of
basic dichotomies. The same convictions
required the Microhylidae to be closely
related to ranids.

The Pseudidae were considered lepto-
dactylids until Parker (1935) suggested
that they were hylids (because of the
presence of an intercalary phalanx ) . The
intercalary phalanx of pseudids is elon-
gate and osseous instead of short, disc-
like, and cartilaginous as in centrolenids,
hylids, hyperoliids, rhacophorine ranids,
and some microhylids (Phrynomerus).
Savage and Carvalho ( 1953 ) named a
new family for the Pseudidae (Lysapsus
and Pseiidis) because they considered the
accessory phalanx analogous to the in-
tercalated cartilage. As pointed out by
Burger (1954), Savage and Carvalho's
"new family" was authored by Fitzinger
(1S43). Savage and Carvalho (1953)
considered Pseudis more primitive than
Lysapsus and suggested that the family

was derived from the Leptodactylidae.
Burger ( 1954 ) considered Lysapsus more
primitive than Pseudis and suggested
that the group was a hylid subfamily. I
have not studied pseudids in detail, and
much information is lacking for Lysapsus,
but I consider the group more closely
allied to leptodactylids than to hylids.
The pseudids are not closely related to
any leptodactylid group.

The ranoid families ( Hyperoliidae,
Sooglossidae, Ranidae, and Rhacopho-
ridae) are usually considered remote
from leptodactylids. Sooglossus and
'Nesomantis were included in the Sooglos-
sinae, a pelobatid subfamily with ranoid
parallelisms by Noble (1931). The sub-
family is restricted to the Seychelles.
Darlington (1957) doubted on zoogeo-
graphic grounds (with preconceived ac-
ceptance of Matthew's conclusions ) that
the Sooglossinae could be pelobatids,
and Griffiths ( 1960), with similar zoogeo-
graphic biases, demonstrated ranoid af-
finities for the group and elevated the
subfamily to family rank. The Sooglos-
sidae have some pelobatid, myobatra-
chine leptodactylid, and ranoid traits and
are conceivably modern representatives
of a leptodactylid derivative that gave
rise to the Ranidae. Until further study
is made of the numerous hyperoliid,
ranid, and rhacophorid genera, aclditional
comments on the relationships of these
families to leptodactylids are held in
abeyance. Vertical pupils are considered
to be primitive by me and vertical pupils
do not occur in the Ranidae or Sooglos-
sidae. Vertical pupils are common in the
genera of two of the subfamilies of Lau-
rent's (1951) Hyperoliidae ( Astylosteri-
nae and Hyperoliinae). Most of these
genera also lack outer metatarsal tu-
bercles.

ZOOGEOGRAPHY

Almost without exception, previous
zoogeographic studies utilizing or in-
volving anurans have been Matthewian
in analysis and conclusion. Darlington
(1957) voiced numerous suggestions con-

cerning anuran zoogeography but lacked
an understanding of the relationships of
most groups. Noble ( 1924, 1926c, 1930)
attacked many zoogeographic enigmas,
but his approach was strictly Matthew-

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

ian. Many macrosystematic problems of
frogs have been studied and solved in
the past decade, and the major evolu-
tionary patterns of the Anura are only
now beginning to emerge. Many prob-
lems, principally minor ones, remain to
be solved; a major difficulty to be over-
come is the relationships of the families
assigned to the Ranoidea, especially the
Microhylidae. Paleontological work has
continued to force biologists to accept
the antiquity of frogs; the order had
diversified into several families by the
Jurassic (Tihen, 1965). As the known
antiquity of frogs increases, so must the
consideration that continental drift may
have played an important role in estab-
lishing their present distributions. How-
ever, simply because a group is an old
one, continental drift need not be in-
voked to explain distril)ution patterns
(cf. Kluge, 1967).

The present distribution of the Lepto-
dactylidae ( Fig. 1 ) is suggestive of a
southern origin and dispersal. No fossil
evidence is available to establish the
presence of the family in the northern
hemisphere before the late Tertiary ex-
cept for the Lower Eocene Indolxitra-
chus from peninsular India. Noble (1930)
and Darlington ( 1957 ) cited the fossil as
evidence of a northern occurrence of the
otherwise Australo-Papuan Myobatra-
chinae. Both authors assumed that pe-
ninsular India has always been part of
the Asian continent or at least has been
part of it for sufficiently long that, zoo-
geographically, India is part of Asia.
However, contrary evidence is impres-
sive. Paleomagnetic studies place Bom-
bay at 40 "" S in the Jurassic and record a
steady northerly movement of the sub-
continent throughout the later Mesozoic
and Cenozoic. During the Eocene, Bom-
bay was at lO'' S (Takeuchi, Uyeda, and
Kanamori, 1967 ) , or at about the level of
northern Australia, and the present Hi-
malayan region was crossed by the Sea
of Tethys.

The frogs of the family Leptodactyl-
idae presently occur in Australia (and
New Guinea and Tasmania ) , in southern

Africa, and in the Neotropics from south-
ern Chile and Argentina (53-54° S) north
to Arizona, Florida. New Mexico, and
Texas (30-33° N). With the exceptions
of a few leptodactyline and telmatobiine
genera which range northward into Mid-
dle America and the southern United
States, no leptodactylid subfamily occurs
on two continents. The Cycloraninae
and Myobatrachinae occur only in the
Australo-Papuan Region (two genera
reach eastern New Guinea and three
reach Tasmania), the Heleophryninae
(monotypic) occur only in southern Af-
rica, and the Ceratophryinae, Elosiinae,
Leptodactylinae, and Telmatobiinae oc-
cur only in South and Middle America.
With the exception of Indohatnichus, the
fossil record for each subfamily (insofar
as it is known ) is included in the Recent
distribution of the group (on the same
continent). Five genera of the Telma-
tobiinae range outside of South America;
four of these (Hijlacfophryne, Sminthil-
lus, Syrrhophiis, and Tomodactyhis) do
not occur in South America, but the other
(Eleiitherodactijlus) is wide-spread in
tropical South America. Sminfhilhis is a
Cuban endemic, and the other three non-
South American genera are principally
distributed on or around the Mexican
Plateau. Three genera of the Lepto-
dactylinae range outside of South Amer-
ica. Fleurodema occurs in the savannas
of central Panama but has its center of
distribution in temperate South America.
Physakiemus occurs in the Central Amer-
ican lowlands as far northwest as Oaxaca
and Veracruz, Mexico, but has its center
of distribution in subtropical and tropical
Argentina and Brasil. In the case of both
genera, only a single species enters Cen-
tral America. Leptodactylus ranges
northwestward through Central America
to Texas and Sinaloa but all Central
American species are also found in north-
ern South America. Leptodactylus also
occurs on Hispaniola and Puerto Rico
and on several islands in the Lesser An-
tilles, but is not represented by extra-
South American endemics. All of the
Middle American leptodactylines arc

LYNCH: LEPTODACTYLOID FROGS

213

Soiitli American species wliich have
spread northward since the Late Phocene
closmx' of the Panamanian portal (Lloyd,
1963). The Telmatobiine genera all be-
long to the Eleutherodactylini. Three of
the extra-South American genera (Smin-
thillus, Syrrhophus, and Tomodactyhis)
are derivatives of the alpha division of
Eleuthewdactylus. The alpha division of
EJentherodactyhis is centered in the
West Indies ( about 100 species ) but also
occurs in the Andean system in Colom-
bia, Ecuador, Guyana, and Venezuela.
The fourth extra- South American eleu-
theroclactyline (Hylactophryne) is not ob-
viously derived from Eleufherodactylus
and shows a greater affinity with an
Amazonian genus (Ischnocnema). The
paleogeographic maps compiled bv Har-
rington (1962), Jacobs et al (1963), and
especially Lloyd ( 1963 ) strongly suggest
that South America was not connected to
North America during the Mesozoic and
most of the Tertiary. Contact was estab-
lished in the Pliocene (Lloyd, 1963).
Animals could have moved northward
across the volcanic island chain in the
latter half of the Tertiary but the degree
of isolation of animal groups suggests
that the terrestrial South American faima
was isolated in South America until the
Pliocene. No leptodactylid group reached
North America via a Panama-Costa Rica
route until late in the Tertiary. Some
leptodactylids may have reached North
America earlier (perhaps Miocene) by
way of the Antillean arc. Therefore, until
the Middle Tertiary we may ignore
North America insofar as leptodactylids
are concerned. The history of the family
lies in South America and the southern
hemisphere. Vinson and Brineman (1963)
suggested that Middle and South Amer-
ica were connected by a Late Paleocene
land bridge. These authors argued that
this is true because of the lack of Danian-
Paleocenc marine formations in the Isth-
mian region of Panama. This land bridge
was essential to Savage's ( 1966), analysis
of the history of the Middle American
heipetofauna.

The Australo-Papuan and African

leptodactylid groups share a few charac-
ters but each of the three groups has
more characters in common with the
pelobatid subfamily Megophryinae. The
Megophryinae seem to be ancestral to
the Leptodactylidae, and the lack of ob-
vious interrelationships between the
Heleophryninae, Cycloraninae, and Myo-
batrachinae suggests that each subfamily
is an independent derivative of the mego-
phryine stock. The Cycloraninae and
Heleophryninae share one interesting
character ( the fusion of the cervical and
second vertebrae ) which does not appear
elsewhere in the family. The fusion ap-
pears in some African and South Amer-
ican bufonid genera, rhinodermatids,
palaeobatrachids, and pelodytids; the
significance of this character distribution
is not entirely apparent at present.

The Recent distrijjution of the Mego-
phryinae is much smaller than its early
Tertiary distribution. Eopelobates is
known from Europe in the early Miocene
and western North America in the Eo-
cene and Oligocene and is probably rep-
resented in the late Cretaceous deposits
of Wyoming. The Megophryinae are
presently restricted to southeastern Asia
and the Indo-Australian archipelago west
of Wallace's Line but do not occur on
Luzon and Masbate (Philippines).

Both the Cycloraninae (Lechriodus)
and Myobatrachinae (Crinia) occur in
eastern New Guinea, and Lechriodus also
occurs on the Aru Islands. Lechriodus
has three endemic species in New Guinea
and the Aru Islands, but one other spe-
cies also occurs in eastern Australia. The
single Crinia which occurs on New
Guinea is also widespread on the Aus-
tralian mainland. Each subfamily is dis-
tributed over most suitable anuran habi-
tat on the Australian mainland and both
are absent over most of the western
Eyrean desert.

Among living leptodactylids, the Cy-
cloranini are least unlike the primitive
Neotropical leptodactylid genera. The
morphological hiatus between the Neo-
tropical leptodactylids and the several
pelobatid stocks requires an intermediate

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MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

stage whose characteristics are Hke those
exhibited by the Cycloranini. Darhngton
(1957, 1965) considered the temperate
South American frog fauna depauperate
and thus of Httle zoogeographic impor-
tance. Cei (1962a) and Vellard (1957)
contended that the fauna is rehct. Vuil-
leumier (1968) analyzed the amphil)ian
fauna of the Nothofagus forests of tem-
perate South America and noted that the
present anuran distributional patterns are
a result of Pleistocene events but ad-
mitted that the high degree of endemism
is a consequence of a long history in the
Patagonian forests. Vuilleumier ( 1968)
considered the frog fauna of the Notho-
fagus forests to consist of four elements:
1) leptodactylid stocks which are au-
tochthonous and have not subsequently
diversified; 2) autochthonous lepto-
dactylid elements which have subse-
quently radiated into northern South
America; 3 ) Nothofagus endemics which
are derived from tropical South Amer-
ican leptodactylids; and 4) bufonid and
leptodactylid stocks which are wide-
spread in South America and have more
species outside of the Nothofagus forests
than in it. He considered the available
data as inadequate to permit determina-
tion of whether these groups are secon-
dary or primary inhabitants of southern
South America. Vuilleumier's analysis
must be rejected because his conclusions
are in part based upon the erroneous
conclusions of other authors. His second
element [2), above] consisted of En-
psophus. As I pointed out ( Lynch, 1971),
Eupsophus (as used by Vuilleumier and
many other authors) is a composite of
seven genera. Eupsophus is restricted to
western Argentina and Chile, except for
two Andean species in southern
and central Peru. Vuilleumier's third
element has been altered taxonomically
by Barrio (1967a) and Lynch (1971).
Batrachyhi is valid and contains three
species (all Nothofagus endemics); the
genus is not an Eleutherodactylus de-
rivative but an offshoot of a Eupsophus-
Htjlorirm stock. The fourth element of
Vuilleumier was misrepresented in part.

Vuilleumier included the genera Bufo
and Fleurodema. The former is wide-
spread and species-rich in tropical and
subtropical South America but Fleuro-
dema is basically a temperate South
American genus ( defined on the basis of
cool or cold winters and cool summers).
The distributions of eight of the ten
species of the genus are contained in
temperate South America; the other two
species range northward into subtropical
and tropical areas in Brasil, Colombia,
Venezuela, and the Guianas (Fig. 128).
Fleurodema is a temperate zone genus
which has invaded the tropical zone,
whereas Bufo is more a tropical zone
genus which enters the temperate zone.

Batrachophrymis, Batrachyla, Caudi-
verhera, Eupsophus, Hylorina, Fleuro-
dema, Telmatohufo, and Telmatohius are
temperate South American leptodactylid
genera either wholly or principally re-
stricted to the temperate zone. This list
of genera includes most of the primitive
Telmatobiinae and the most primitive
living Leptodactylinae. Ceratophrys,
Lepidohatrachus, Limnomedusa, and
Thoropa are also considered primitive,
and with the exception of Ceratophrys
do not range north of the southern sub-
tropical zone.

The primitive Cycloranini morpho-
logically resemble the primitive Neo-
tropical genera, which are the principally
temperate South American genera. These
coincidences require that we regard the
similarities as convergent or that we con-
sider them products of common ancestry.
The latter conclusion further requires
some land connection between Patagonia
and Australia; the most plausible route is
via Antarctica. To conclude that the
route involved the Holarctic Region re-
quires a massive extinction of the stocks
which passed through the Holarctic and
Neotropical regions, and further requires
that these stocks survived only in tem-
perate South America from which they
gave rise to new groups which then in-
vaded the tropical zone. A possible
causative agent for such a mass extinction
would be climatic zonation and decreas-

LYNCH: LEPTODACTYLOID FROGS

215

Figure 128. Distribution of leptodactylids in South America. (A) Black areas encompass the range
of the Alsodini. Thoropa is restricted to southeastern BrasiL The dotted Hne encloses the range of
the genera Tehnatobius and Batrachophrynus. Two other genera of the tribe (Caiidiverbera and Tel-
matobitfo) occur in the black area in Chile. ( B) Ranges of the genera Limnomedusa and Plevrodema,
the most primitive genera of the Leptodactylinae. (C) Range of the Elosiinae. The area in southeast-
em Brasil encompasses the distribution of Crossodactijlus and Megaelosia; the genus HijJodes also
occurs on Cerro Duida, Amazonas, Venezuela. The range of the Elosiinae in southeastern Brasil

approximates the range of the Grypiscini.

ing climatic eqiial)ility of the northern
hemisphere during Cretaceous time.
However, this explanation results in two
principle difficulties: 1) why did not
leptodactylid stocks survive in areas in
the northern hemisphere which retain
high equabilities?, and 2) the mego-
phryine pelobatids are probably equally
sensitive to low equabilities yet they
persisted in North America until the
Oligocene and survive today in southeast
Asia.

Equability is a property of climate
which expresses departures from 14.0° C
and thus responds primarily to tempera-
ture extremes (Axelrod and Bailey, 1968).
High equabilities reflect little variation
in temperatures about an annual mean of
14.0° C. Equability applies equally well
to warm and cold regions. Bailey ( 1960,
1964 ) also noted a second aspect of tem-
perature that affects biotic composition
—effective temperature, which expresses
warmth of the climate in terms of tem-
perature and the duration of the summer
(Axelrod and Bailey, 1968). Effective
temperature is independent of the mean
annual range of temperatures.

Axelrod and Bailey ( 1968) noted that
many areas in the southern hemisphere
harbor relicts of the Cretaceous flora
(cycads, tree ferns, podocarps, arau-
carias). It is in some of these regions
(south Africa, Australia, southern South
America, southeastern Brasil) that prim-
itive leptodactylids ( Cycloraninae, Myo-
batrachinae, Heleophryninae, Telmato-
biini, and Alsodini) occur. These areas
are characterized bv high climatic equa-
bility ( M = 60+). '

Mesozoic climates were characterized
by high equabilities ( Axelrod and Bailey,
1968). Equability is increased if the
locality is associated with large bodies
of water (for example, the equability of
localities along the Peruvian coast is
higher than might be expected, as are
the equabilities for localities in coastal
Uruguay). The broad marine embay-
ments and epeiric seas of Mesozoic land-
masses probably contributed to the
maintenance of high equabilities in con-
tinental situations. Under a temperature
regime of high equabilities, the dispersal
routes of Mesozoic animals would not be
temperature-limited. The southern hemi-

216

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Figure 129. (A) Distribution of the Odontophrynini. ( B) Distribution of the Ceratophryinae. (C)
Distriliution of the LeptodactyHnae except for the primitive genera Limnomedusa and Fleurodema.

sphere leptodactylid stocks could have
invaded the northern hemisphere were
land connections available because the
northern hemisphere was also under a
regime of high equabilities until the
Cretaceous. During the Cretaceous many
plant and animal groups became extinct
or began to flourish. At the same time
the earth began to experience a marked
climatic zonation. With the development
of climatic zonation, broad areas of the
world experienced a decrease in equa-
bility. Axelrod and Bailey suggested that
the primitive faunas and floras presently
living in areas of high equability survived
in those regions because these groups
are, and their ancestors were, adapted to
climates of high equability.

The part of South America (Figs. 129-
130) with high equabilities (58+) in-
cludes the ranges of almost all of the
primitive genera of Icptodactylids (Ba-
trachophryrms, Batmchijla, Caudwer-
bera, Etipsophus, Hylorimi, Lepidohatra-
chus, Limnomedusa, Fleurodema, Telma-
tobius, and Tehrmtobufo). A few of the
species in these genera occur in areas of
low equability ( for example, Fleurodema

brachyops and P. diplolistris occur in
areas with equabilities of 40-45). The
Leptodactylinae probably evolved in re-
sponse to the decreasing equability of
the Late Cretaceous (see below). Before
returning to the history of the Lepto-
dactylidae in South America, it is con-
venient to review briefly the history of
the African and Australo-Papuan groups
and the Megophryinae and to discuss the
possible role of climatic equability in the
establishment of the Recent distributions
of these groups.

The Megophryinae have survived in
areas with relatively high equabilities.
Even the temperate eastern Himalayas
have equabilities of 60-[-. Regardless of
which stance ( Wegenerian or fixed con-
tinents) one wishes to take, the Mego-
phryinae are probably a group which
evolved in the northern hemisphere. The
Cycloraninae, Heleophryninae, Myoba-
trachinae, and Pelobatinae are the deriv-
atives of the Megophryinae. Two of
these derivatives are Australo-Papuan,
one south African, and one Holarctic.
The proximity of the present distributions
of the Megophryinae, Cycloraninae, and

LYNCH: LEPTODACTYLOID FROGS

217

Myobatrachinae tempts one to assume
that megophryine stocks crossed the
Inclo-AustraHan archipelago into Aiis-
traha in the early Cretaceous with the
Marsupalia and boid and elapid snakes.
However, Inger ( 1954 ) considered the
Megophryinae late invaders of the Phil-
ippine Islands. Of further significance in
this regard is the wider distribution of
the Megophiyinae in the Late Cretaceous
and early Tertiary of North America and
Europe. Proponents of continental drift
argue that the proximity of the Sundan

shelf and Sahul shelf is a relatively recent
phenomenon. The Indo-Australian archi-
pelago is usually cited as the route
whereby northern-evolved groups
reached Australia in the Cretaceous
(Clemens, 1968, Darlington, 1957, 1965).
Biologists seeking support for this route
have cited Audley-Charles (1966), who
after studying the geology of Timor con-
cluded that the relationship between
Timor and the Sahul shelf has not
changed since the Middle Triassic and
that the relationship between Timor and

50.0

45.0

-55.0

Figure 130. Map of South America with isoequaphane h'nes (45, 50, 55, 60 and above) super-
imposed. All dots on the map represent stations for which equability \ alues ( after Axelrod and
Bailey, 1968) were computed. The hatched areas have equabilities of 50 or less.

218

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

tlie rest of the archipelago is apparently
as ancient. As Hallani ( 1967 ) remarked,
". . . something more than island chain
links is required to account for the pres-
ence in Australia of the lungfish Neo-
cerotochis and large Jurassic and Cre-
taceous dinosaurs."

A Jurassic dispersal of megophryinc
stocks through eastern Africa onto penin-
sular India, Antarctica, and Australia
(Fig. 131) would explain the similarities
of the Heleophryninae and some prim-
itive Cycloranini, but would require that
these continental masses be in close prox-
imity instead of being widely separated
as they are today. This route also re-
quires that the megophryines not invade
South America which was in close prox-
imity with Africa until the Neocomian
(Lower Cretaceous). The megophryine
stock probably had the same reproduc-
tive biology as do the living pelobatids
and the Australo-Papuan leptodactylids.
During the Jurassic and early Cretaceous,
the Brasilian and Guiana shields must
have been formidable barriers to pond-
breeding frogs. On the slopes of these
uplifted shields, ponds would be rare if
they existed at all. In contrast, ponds
would be available along the western
edge of Africa in the vicinity of the East
African Gulf of Tethys sea (Hallam,
1967). If the interior of Antarctica was
as forbidding to amphibians as Darling-
ton ( 1965 ) reasons it must have been,
then the only southern route would have
been across that part of Antarctica now
called Enderby Land and the American
Highland. Hallam (1967) suggested that
a deep marine sea separated South Amer-
ica and Africa from Antarctica, Australia,
and Peninsular India in the Neocomian.
Harrington ( 1962 ) noted the occurrence
of marine facies over the southernmost
tip of South America in the Neocomian
but the extent of this marine sea was not
as great: as suggested by Hallam. The
leptodactylid stock that did reach south-
ern South America probably entered the
continent in Callovian (Upper Jurassic)
or Neocomian (Lower Cretaceous) times.
Whether dispersal was by a corridor or

filter bridge (Darlington, 1957) route is
non-consequential; dispersal could even
have been via a sweepstakes route. The
significant point to be made here is that
contrary to Darlington's (1965:38) asser-
tion, one group of terrestrial vertebrates
(leptodactylid frogs) does show special
relationships between the southern tem-
perate fomis on different continents.
Darlington stressed the absence of close-
ly related cold-temperate vertebrates in
Tierra del Fuego and Tasmania. How-
ever, this absence is explained by in-
creased cold and lowered equabilities in
Tierra del Fuego. Once the Australo-
Papuan groups reached Peninsular India
and Australia, and the Australo-Papuan
Cycloranini spread across Antarctica to
Patagonia, continental connections were
no longer necessary because each of the
subfamilies occupied the appropriate
continental masses. The probable time
period was [or perhaps somewhat earlier
than] the Neocomian (Lower Creta-
ceous). After this time, Australia and
South America appear to have been com-
pletely isolated from all other land masses
until the middle Tertiary in the case of
Australia (connection via the Indo- Aus-
tralian archipelago) and the late Tertiaiy
in the case of South America ( connection
via the Panamanian Isthmus).

The early Cretaceous continental
separations were followed by the impo-
sition of a severe climatic regime on the
distribution of the early Cretaceous fau-
nas and floras. Insofar as the Mego-
phryinae and their derivatives are con-
cerned, the development of this new
climatic regime had several important
consequences: 1) evolution of a low-
equability adapted group, the Peloba-
tinae, in the northern hemisphere; 2)
gradual extinction and range restriction
of the Holarctic Megophiyinae; 3) isola-
tion of the Heleophryninae into a high
equability area in south Africa and/ or
extinction of Heleophryninae or Mego-
phryinae in east Africa; 4) extinction of
Myobatrachinae in peninsular India; and
5) evolution of low-equability adapted
groups in Australia and South America.

LYNCH: LEPTODACTYLOID FROGS

219

The high-equability adapted groups were
restricted in geographic distribution to
areas of high equabiHty in southern

South America and AustraHa. The de-
velopment of climatic zonation probably
resulted in physiological stresses on the

Figure 131. Paleozoogeographic maps for three stages of the evolution and dispersal of leptodactyhd
frogs and for related groups. (A) Jurassic (Callovian) — southerly dispersal of Megophryinae. ( B)
Late Jurassic or eariiest Cretaceous — Africa is isolated from southern masses; climatic equability is
beginning to influence dispersal routes. Leptodactylid invasion of South America. The dashed line is
the route of free exchange of other vertebrate groups (e.g., characoid fishes and pipid frogs). ( C) Mid-
dle Cretaceous — Africa-South American connection is tenuous, Neotropical leptodactylids are radi-
ating from Patagonia into low-equability zones. Bufonidae spread into Africa.

220

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

surviving fauna and flora; these stresses
forced a concentration of the Cretaceous
genera into the remaining areas of high
equabihty and may well have been the
principle factors causing a radiation of
the Neotropical Icptodactylids. The de-
velopment of foam-nesting habits in the
breeding biologies of the Limnodynastini
may have been stimulated by decreasing
e(|uabilit)'. The foam-nesting habit en-
ables the frogs of this group to survive
in more xcric, and less equable, climates
than are suitable for the Cycloranini and
Myobatrachinae.

The distribution of Icptodactylids in
South America was greatly restricted
during the later Cretaceous, when cli-
matic zonation was developing and equa-
bility was decreasing. The fauna prob-
ably included only a half dozen genera
(Batrachyla, Caud'werhera, Etipsophus,
Lepidohatrachus, Telmatobiifo or Neo-
procoela, and TJwropo). In response to
decreasing equabilities the range of the
Alsodini was contracted and probably
resulted in the isolation of a relict popu-
lation on the Brasilian shield. This popu-
lation probably gave rise to the Elosiinae
and Grypiscini. The decreasing equabil-
ities also resulted in the evolution of a
group of frogs utilizing foam-nests in
their breeding biology. The ancestral
stock of this group is Etipsophus which
gave rise to Pleiirodema and possibly to
Limnomedusa. The presence of a foam-
nest enabled these frogs to breed in more
xeric environments, but presumably the
adults were adapted to high equabilities
and were therefore unable to successfully
invade the tropical zone. New genera
evolved to occupy this region (Lepto-
dactyhis and Physalaemus). Batrachyla
and Thoropa lay their eggs in moist ter-
restrial situations but require water for
their aquatic tadpoles. The Elosiinae
represents a derivative of this group
which became isolated on the Brasilian
and Guiana shields (high equability).
The Eleutherodactylini also evolved from
an ancestral stock not unlike the Alsodini
and Grypiscini. The Grypiscini and
Eleutherodactylini evolved direct devel-

opment, which enabled these groups to
invade regions lacking ponds. The evo-
lution of these groups may have been
prompted by avoidance of competition
in the larval stage. The evolution of
Eleutherodactyhis is probably partly cor-
related with the Andean orogeny in the
Mid-Tertiary. The Ceratophryinae and
Odontophrynini represent groups that
evolved in response to decreasing equa-
bility but were adapted to xeric, non-
forested habitats. The principle low-
equability adaptations of these groups
involve a heavier, drier skin.

The Dendrobatidae are the low-
equability adapted derivatives of the
high-equability adapted Elosiinae. The
similarities between the Dendrobatidae
and Petropedetinae and the distributions
of these two groups are suggestive of
their evolution being synchronous with
the last stage of separation of eastern
Brasil and Africa. The Petropedetinae
are low-equability adapted frogs. The
radiation of the Dendrobatidae probably
occurred during the Andean orogeny; the
dispersal of the group into Central Amer-
ica is a I^ate Tertiary phenomenon (Sav-
age, 1966). The Bufonidae may represent
a paedomorphic offshoot of the Telma-
tobiinae rather than the Myobatrachinae.
The heavy, frequently dermostosed skulls
and thick, dry-adapted skin of bufonids
suggest that this group evolved in re-
sponse to decreasing equability. The
antiquity of the group is not directly
known, but the earliest fossils are Lower
Miocene (North America and Europe).
The most primitive genus is African
(Tihen, 1960a), but the remainder of the
genera are derived from the widespread
Bufo. Among its derivatives are seven
Neotropical, seven African, and five Ma-
laysian and Philippine genera. Bufo
ranges over all continents except Aus-
tralia and Antarctica. The Bufonidae
probably originated in Africa and South
America when the two continents were
connected. The early evolution and dis-
persal of the Bufonidae is considered
syntopic witli tliat of the Pipidae and
characoid fishes. The Gymnophiona ex-

LYNCH: LEPTODACTYLOID FROGS

221

hibit a distribution and radiation pattern
which is similar to tliat of the Bufonidae
except tliat there is no caecihan genus
which ranges over the Holarctic. The
evidence suggesting that the Bufonidae
represent an offshoot of the Myobatra-
chinae is meager. The mouthparts of the
tadpoles of the two groups are similar.
It is possible that the Bufonidae are a
paedomorphic myobatrachine offshoot
that invaded South America synchro-
nously with the Telmatobiinae. If Ant-
arctica was under a temperate regime of
increasingly lower equability while Aus-
tralia was under one of higher equability,
it is conceivable that a low-equability
adapted myobatrachine group evolved
in Antarctica and dispersed into South
America before the South America-Ant-
arctica land bridge was obliterated in
the Cretaceous. One serious objection to

this hypothesis is the absence of any
high-equability adapted bufonid genera
in South America. Rhinoderma is a high-
equability adapted genus and is related to
the Bufonidae. Rhinodermatidae may
represent this high-equability adapted
proto-bufonid. The bufonid genus Mel-
anophnjmsciis lives in areas of relatively
high equability in northern Argentina
and Uruguay but does not range south-
ward into the zone of very high equa-
bility (5S+). A further difficulty with
the hypothesis is the requirement of a
low-equability corridor through the high
equability zone which presumably cov-
ered all of Patagonia during the Late
Cretaceous and subsequent Tertiary. A
more thorough studv of Rhinoderma mav
prove to be the key in ascertaining the
relationships of the Bufonidae to the
Australo-Papuan or southern South
American leptodactylids.

SUMMARY AND CONCLUSIONS

Based on the variation of behavorial
and morphological characters, 57 Recent
and three fossil genera of the Lepto-
dactylidae are recognized. These 60 gen-
era are placed in seven subfamilies, two
of which are further subdivided into
tribes. Two of the subfamilies (Cyclo-
raninae and Myobatrachinae) occur only
in the Australo-Papuan region, one
(Heleophryninae) in southern Africa, and
four ( Ceratophryinae, Elosiinae, Lepto-
dactylinae, and Telmatobiinae) in the
Neotropical realm. ^■'''

The fossil record of leptodactylid
frogs is of little value in deducing macro-
systematic phylogeny. Fossils are repre-

'" The alternatives to a single family arrange-
ment are cited elsewhere (footnote 14, p. 204).
The alternatives are: 1) two families, an Old
World Myobatrachidae and a Neotropical Lep-
todactylidae; 2) two Old World families, an
African-Australian group for the Cycloraninae
and the Heleophryninae, an Australian group for
the Myobatrachinae, and one Neotropical family;
and 3) one Old World family and two Neo-
tropical families, one for the two genera of the
Ceratophryinae and a second for the other three
Neotropical subfamilies. Other alternatives are
simply permutations of the above.

sented in the Pleistocene of North Amer-
ica, the West Indies, and South America.
Fossils of three phyletic lines are pre-
served in Lower Eocene to Pliocene de-
posits of Patagonia. All of the above-
mentioned fossils are members of genera
living in the same regions today or are
closely related, extinct genera. The only
zoogeographically important fossil is the
Lower Eocene Indohatrachus, a myoba-
trachine from peninsular India. The sub-
family Myobatrachinae is presently re-
stricted to the Australo-Papuan region.

In order to determine the evolutionary
direction ( primitive to advanced ) of sev-
eral evolutionary trends, the following
reasoning was used: character states
shared by some or all archaic frog fam-
ilies and the other lissamphibian orders
are primitive in the Anura; character
states shared by most archaic frogs and
most pelobatids are primitive in the
Anura; and character states which are
rare in the archaic families but always
evident in the Pelobatidae are primitive
in the pelobatid-bufonoid superfamily
complex and may be primitive in all

222

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

frogs. This analysis resulted in the con-
clusion that the following character com-
plexes are useful in ascertaining the rela-
tive primitiveness of frog groups: 1) type
of cervical cotylar arrangement and type
of occipital condylar arrangement; 2)
neural arches— imbricate or not; 3) rela-
tive lengths of transverse processes of
presacral vertebrae; 4) extent of dilation
of sacral diapophyses; 5) presence of
zygapophyses involved in sacral-coccyg-
eal articulation; 6) separation of inter-
vertebral discs from the centra; 7) com-
plexity of the ilium; 8) loss of skull bones
and teeth; 9) modifications of phalangeal
formulae; 10) shape of pupil; 11) pres-
ence of outer metatarsal tubercles; 12)
amplectic position; 13) egg size, oviposi-
tion site, larval development; 14 ) tadpole
vent position; 15) tooth row formula of
tadpole; and 16 ) architecture of pectoral
girdle.

The Megophryinae (Pelobatidae)
gave rise to four principle groups: 1) the
Pelobatinae, 2) the Heleophryinae, 3)
the Cycloraninae, and 4) the Myobatra-
chinae. Each of these groups appears to
be an independent derivative. All of the
Neotropical leptodactylids arc probably
descendants of one invasion of a cyclo-
ranine stock into South America. The
Ceratophryinae and Telmatobiini are the
most primitive Neotropical groups. The
major leptodactylid radiation occurred
within South America and radiated out
of southern South America. One primi-
tive tribe of the Telmatobiinae (Alsodini)
gave rise to the advanced tribes of the
subfamily (Grypiscini and Eleuthero-
dactylini) as well as to two additional
subfamilies (Elosiinae and Leptodactyl-
inae). The Dendrobatidae are a deriv-
ative of the Elosiinae. The Telmatobiini
and Odontophrynini are two tribes of the
Telmatobiinae which appear to be the
most primitive.

The evolution of leptodactylids is best
explained on a superstructure involving
continental drift. The following paleo-
zoogeographic sequence is proposed: the
basal stock, the Megophryinae, origi-
nated on northern landmasses from an

ascaphid-discoglossid ancestor and dis-
persed southward in the Middle Meso-
zoic. A single dispersal corridor was
utilized, the area in east Africa to the
west of the East African Gulf of Tethys
Sea. The megophryine stock could not
invade South America because of the
lack of a lowland corridor between North
and South America or between Africa
and the Brasilian shield. The stock
passed through south Africa onto the
southern landmass composed of penin-
sular India, Australia, and part of Ant-
arctica. Progressive southerly extension
of the East African Gulf isolated the
derived group on India, Australia, and
coastal Antarctica. During the Late Ju-
rassic or early Cretaceous, a cycloranine
stock dispersed along the coast of Ant-
arctica into southern South America.
Concurrently, climatic zonation and the
ensuing decrease in climatic equability
were bringing about the extinction of
many groups of plants and animals. In
the northern hemisphere, the Mego-
phryinae gave rise to a low-equability
adapted group, the Pelobatinae, and
were restricted in distribution to a high-
equability zone in southeast Asia. Lower-
ing equability resulted in the isolation of
the African megophryine derivative in
the high-equability refugium in south
Africa. This group is the Heleophryninae.
The effect of decreasing equability in
Australia was minor. The Cycloraninae
evolved a low-equability tolerant group,
the Limnodynastini. Decreasing equabil-
ities with the northward movement of
peninsular India during the Cretaceous
and Tertiary resulted in the extinction of
the myobatrachines living there. In
South America, the more primitive lepto-
dactylid genera survive in the area of
high equability (often termed temperate
South America). Decreasing equability
resulted in the evolution of a low-equa-
bility adapted group, the Leptodactyl-
inae. The Ceratophryinae and Odonto-
phrynini are arid-adapted frogs and were
therefore able to invade the low-equabil-
ity areas which were arid or semi-arid.

LYNCH: LEPTODACTYLOID FROGS

223

Decreasing equabilities probably initially
led to range restrictions which resulted
in the isolation of the Elosiinae and
Grypiscini in southeastern Brasil (a high-
equability zone). The Dendrobatidae are
low-equability adapted frogs which
evolved from the high-equability adapted
Elosiinae. The African ranids of the sub-
family Petropedetinae are usually cited
as ranids with adaptations paralleling the
Neotropical dendrobatids. I have studied
briefly the anatomy of Petropedetes and
find the similarity to dendrobatids strik-
ing. The two family groups may repre-
sent remnants of a group once ranging
across the South American-African isth-
mus. However, before the relationship
can be established, more study of the
ranids needs to be made.

In the course of this study, many
taxonomic changes have been proposed.
These are summarized below.

1. Geobatrachus Ruthven is removed
from the Leptodactylidae and tentatively
assigned to the Microhylidae, pending
completion of a study of the systematic
position of the genus by Dr. Charles F.
Walker.

2. Htjlopsis pJattjcephalus Werner is
a species of the Hylidae and perhaps not
separable from Hyla.

3. Rhinodernia is placed in a mono-
typic family, the Rhinodermatidae.

4. The subfamily Cycloraninae is rec-
ognized for 10 genera. The subfamily is
divided into two tribes: Cycloranini
(Cyclorana, Heleioponis, Neohatrachus,
and Notaden) and Limnodynastini new
tribe (Adelotus, Kyarranus, Lechriodus,
Limnodynastes, and Philoria).

5. The subfamily Myobatrachinae is
recognized for one fossil and seven Re-
cent genera (Crinia, Glatiertia, Indoha-
trachus, Metacrinia, Myobatrachus, Pseu-
dophryne, Taudactylus, and Uperoleia).

6. Glauertia mjoebergi is transferred
to the genus Uperoleia.

7. Taudactylus diurnus is identical to
Crinia acutirostris; the genus Taudactylus
is worthy of recognition but the only

species in the genus must now be called
T. acutirostris.^^

8. The subfamily Heleophryninae is
recognized for the African leptodactylid,
Heleophryne. The opinions of some
earlier authors that Heleophryne is a
ranid or the only genus of a monotypic
ranoid family are rejected. The subfam-
ily is similar to megophryine pelobatids
but has achieved the leptodactylid grade.

9. The subfamily Ceratophryinae is
recognized for two Recent genera (Cerat-
ophrys and Lepidobatrachus). The group
is not sufficiently different to be accorded
family rank and is not more closely re-
lated to the Bufonidae than to other
leptodactylids.

10. Chacophrys is considered a syn-
onym of Ceratophrys.

11. Stombus Gravenhorst, 1825, is a
synonym of Ceratophrys Wied, 1824;
Rana cornuta Linne is designated the
type-species of Stombus.

12. The Miocene fossil Wawelia ger-
holdi is a ceratophryine but is not sepa-
rable from either Ceratophrys or Lepido-
batrachus and is therefore tentatively
recognized.

13. The subfamily Telmatobiinae is
recognized for one fossil and 24 Recent
genera of Neotropical leptodactylids. The
following family group names are con-
sidered synonymous with the Telmato-
biinae: Hylodidae Giinther, 1859, Al-
sodina Mivart, 1869, Cacotina Mivart,
1869, Grypiscina Mivart, 1869, Cyclo-
rhamphinae Lutz, 1954, Eleutherodactyl-
inae Lutz, 1954, Calyptocephalellinae
Reig, 1960, and Batrachylinae Gallardo,
1965. The subfamily is divided into five
tribes: Telmatobiini (Batrachophrynus,
Caudiverbera, Neoprocoela, Tehnatobius,
and Tebnatobufo), Alsodini (Batrachyla,
Eupsophus, Hylorina, and Thoropa),
Odontophrynini new tribe (Odonto-
phrynus and Proceratophrys), Grypiscini
(Crossodactylodes, Cycloramphus, and

'" M. J. Tyler (in litt.) informed me that this
statement is erroneous. The two species of
Taudacttjhis are distinctive species. The mate-
rials used in this work are T. acutirostris
(Andersson).

224

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

JMchaeniis), and Eleiitherodactylini (Am-
bh/phrynus, EleutJwrodactijhis, Eupark-
erella, Holoaden, Hykictophryne, Isch-
nocnema, Niceforonia, SmintluUus, Syr-
rhophus, and Tomodactyhts). A new
genus, ScythropJirys, is named for
Zachaenus sawayae Cochran but cannot
be confidently assigned to a tribe until
osteological data are available.

14. The generic name Caudiverhera
is used in preference to Calyptocepha-
lella. Eophractiis Schaeffer is placed in
the synonymy of Caudiverhera. Giganto-
hatrachus parodii (Miocene of Patagonia)
and CalyptocephaJeUa canqueli (Oligo-
cene of Patagonia ) are placed in the syn-
onymy of Caudiverhera caudiverhera.

15. Neoprocoela is not a synonym of
Btifo and is recognized as a genus of
leptodactylid frogs which is ancestral to
BatracJwphrynus.

16. Tehiuitohufo is not a synonym of
Aruncus; Aruncus is a synonym of Bufo.

17. Macrogenioglottus is placed in
the synonymy of Odontophrynus.

18. Proceratophrys is the valid ge-
neric name for the supraspecific group
frequently called Stomhus.

19. Craspedoglossa is considered a
synonym of Zachaenus.

20. Zachaenus roseus Cope is not a
member of the genus Zachaenus and is
considered a species inquirenda in the
Leptodactylidae, probably in the Telma-
tobiinae.

21. Nohlella is placed in the synonymy
of Eleutherodactyhis.

22. Pseudohyla, previously considered
a hylid genus or a synonym of Hyla, is a
synonym of Eleutherodactyhis.

23. Eupsophus wettsteini is trans-
ferred to the genus Niceforonia.

24. SyrrJiophus lapkicai is probably a
member of the genus Niceforonia.

25. Syrrhophus is not a synonym of
Eleutherodactyhis, but the separation of
Syrrhoplms and Tomodactyhis is consid-
ered tenuous.

26. The subfamily Elosiinae is recog-
nized for three Recent genera (Crosso-
dactylus, Hylodes, and Megaelosia).
Hylodes Fitzinger is used in preference
to EJosia Tschudi.

27. The subfamily Leptodactylinae is
recognized for ten Neotropical genera
(Barycholos, Edalorhina, Hydrolaetare,
Leptodactylus, Lininomedusa, Litho-
dytes, Paratehiuitobius, PJiysalaemus,
Pleurodema, and Pseudopahidicola).

28. Paratehnatohius pictiventris A.
Lutz, in Lutz and Carvalho, is a nomen
nudum and an obli<i:ate synonym of
Paratehnatohius gaigeae (Cochran).

29. Pseudopahidicola hohviana Park-
er is considered a synonym of P. pusilla
(Ruthven).

30. Leptodactyhis ahavus Holman
(Lower Miocene of Florida) is a Rana
and questionably separable from Rana
miocenica Holman of the same horizon
and locality.

31. Eoruheta tievadensis Hecht (Low-
er Eocene of Nevada) is not a lepto-
dactylid and is considered "Family in-
certae sedis. Order Salientia."

32. The following new taxa are pro-
posed: Limnodynastini new tribe, Odon-
tophrynini new tribe, and Scythrophrys
new genus (type-species Zachaenus
sawayae Cochran, 1953).

LYNCH: LEPTODACTYLOID FROGS

225

APPENDIX

The following is an alphabetical list
of the species and specimens used in
formulating the generic accounts for the
Leptodactylidae. In addition to the mu-
seum abbreviations listed on page 17,
the following abbreviations (and the
name of the institution) are used below:

CAS California Academy of

Science
LHUBA Laboratorio Herpctologica

Universidad Buenos Aires
NMW Naturhistorisches Museum

Wien
WAM Western Australia Museum

Unless otherwise noted, the specimens
listed below are dry skeletons. Speci-
mens which were cleared and stained
are indicated by "CS"; those which were
studied with stereoradiographs are indi-
cated by "SRC."

Adelotus hrevis.—AMNH 59489-90, KU
56242.

Amhltfphryniis ingeri. — FMNH 54591
(SRG).

Barycholos piiI die r.—UMMZ S-2881 (CS).

Bati(iclio])hn/mis maciostomus. — KU 98127-
28.

Batmchyla leptopus.—HMMZ S-2246 (CS).

Batiachyla t aeniat a. —UMMZ S-2247 (CS).

Caudiverbera catidiverhera. — AMNH 23622,
23958, 24016, 51510, FMNH 9703.

Ceratophrys atiritd.—EHT 1415, FMNH
51703-04, KU 98129.

Cciatophrys calcarata. —KU 69844, JDL S-
76, 237-40, 1189.

Ceratophrys omata.—UMMZ 98834 (tad-
pole, CS), 109753 (juvenile, CS).

Crinia simifera.— AMNH 40291, KU 56243-
49 (CS), 56364 (CS).

Crossodactylodes pintoi. — USNM 102611
(CS).

Cwssodactyhis dispar. — KU 92753 ( CS ) .

Crossodactylus gaudichattdi. — KU 92759
(CS), JDL S-265, 283-84, 417-19.

Crossodactylus grandis.—KU 92765 (CS).

Cycloramphus asper. — KU 92773.

Cydommphus duhitis. — KU 92780.

Cycloramphus eleutherodactylus. — KU
92785.

Cycloramphus fulginosus.—KU 92790-91.
Cycloramphus granulosus. — KU 92795 (CS).
Cycloramphus ohausi. — KU 92801 (CS).
Cycloramphus pinderi. — KU 92807.
Cydorana australis. — AMNH 62228, KU

93549 (CS), 93550.

Cydorami cult ripes.— AMNH 67232 (CS),
KU 110.324.

Cydorana dahli.—UMMZ 65250.

Cydorana platycephalus. — KU 110329.

Edalorhina perezi — AMNH 52847, KU
124225 (CS), 124226.

Eleutherodactylus abhotti.— AS X1795 (CS),
X2006 (CS).

Eleutherodactylus achatmus. — KU 119472
(CS).

Eleutherodactylus antillensis. — AS 12083
(CS), 12104 (CS).

Eleutherodactylus arcolatus. — KU 118129
(CS), 119501 (CS).

Eleutherodactylus armstrongi. — KU 110347
(CS), JDL S-236.

Eleutherodactylus atkinsi.— AS \'6263 (CS ) .

Eleutherodactylus audanti. — JDL S-235.

Eleutherodactylus auriculatoides. — AS X-
9200 (CS).

Eleutherodactylus barlagnei. — MCZ 35331.

Eletitherodactylus bhwtatus. — KU 92813
(CS).

Eleutherodactylus hiporcatus. — KU 41047,
105903.

Eleutherodactylus bogotensis. — KU 1 10408-
09 (CS).

Eleutherodactylus bransfordi. — KU 25217
41032, 103036-41 (CS), JDL S-243, 401.

Eleutherodactylus brederi. — KU 77670 (CS).

Eleutherodactylus brevicrus. — KU 108982
(CS).

Eleutherodactylus bufoniformis. — KU 80621.

Eleutherodactylus caryophyllaceus. — KU

68261 (CS).

Eleutherodactylus cerasmus. — KU 103026-27
(CS).

Eletitherodactylus chloronotus. — KU 118130-
33 (CS), JDL S-247, 260, 335.

Eleutherodactylus cochranae. — AS V8014
(CS).

Eleutherodactylus conspicillatus. — KU
108988 (CS).

Eleutherodactylus coqui.—KU 79924 (CS),
79947-50 (CS).

Eleutherodactylus crassidigitatus. — KU

68262 (CS).

Eleutherodactylus croceoinguinis. — KU
109086 (CS).

Eleutherodactylus cruentus. — KU 102998
(CS), 107941-42 (CS), 117353-54.

Eleutherodactylus cundallL— AS 15128 (CS),
15711 (CS).

Eleutherodactylus curtipes. — KU 109052-58
(CS), JDL S-252-54, 261, 310, 337-38, 342-43,
351, 402, 430-31.

Eleutherodactylus devillei.—UIMNH 55833
(CS).

Eleutherodactylus diastema. — KU 41033-35,

68263 (CS), 80636 (CS), JDL S-244, 441.

226

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Eleutherodacttjhis eneidae. — AS 12637 (CS),
12758 (CS).

Eleittherodactyhis engytympanum. — KU
102999-3007 (CS), 117355-57.

Eleutheiodactyhis fitzingeri. — KU 77658,
117358-62, JDL S-407-08.

Eleutheiodactyliis fteischmanni. — KU 65790
(CS), 68157-58, 68264-65 (CS).

Eletitherodactylus flondentiis.—KU 102242-
46 (CS).

Eletitherodactylus f rater. —KV 80637 (CS).

Eletitherodactylus furcyensis. — AS X2018
(CS).

Eletitherodactylus gaigeae. — KU 106297-98
(CS).

Eletitherodactylus gaklL—USNM GOV 8944
(CS).

Eleutherodacttjhis gollmeri. — KU 41036-37.

Eletitherodactylus gossei. — AS 13795 (CS),
14464 (CS), 15677 (CS).

Eletitherodactylus guentheri. — KU 92819
(CS).

Eletitherodactylus haitianus. — AS X8296
(CS).

Eleutherodacttjhis inoptatus. — AS X2356.

Eletitherodactylus jugans.—MCZ 19856 (2)
(CS).

Eleutherodacttjhis karlschmidti. — AS 12760
(CS).

Eletitherodactylus lenttis. — AS V7395 (CS).

Eletitherodactylus locustus. — AS 11867 (CS),
11881 (CS).

Eleutherodacttjlus longirostris. — KU 77671
(CS).

Eletitherodactylus macdotigalli. — UIMNH
40941 (CS).

Eletitherodactylus martinicensis. — MCZ
35321.

Eletitherodactylus melanostictus. — KU
107943 (CS).

Eleutherodacttjlus mexicanus. — KU 55592-
95 (CS), 55596-98, 55599-600 (CS), 55622,
103008-15 (CS), JDL S-1261-62.

Eleutherodacttjlus mmutus. — AS X8939 (CS).
Eleutherodacttjlus molinoi. — KU 80635 (CS).
Eletitherodactylus montanus. — AS X8313
(CS), X8479 (CS).

Eleutherodacttjlus nasutus. — KU 92822 (CS).

Eletitherodactylus nigriventris. — KU 92739
(CS).

Eletitherodactylus nigrovittatus. — USNM
GOV 8108 (CS).

Eleutherodacttjlus ntihicola. — AS 12891 (CS),
12898 (CS).

Eleutherodacttjlus occidentalis. — KU 102598
(CS), 104101 (CS).

Eleutherodacttjlus octavioi. — KU 92828 (CS).

Eleutherodacttjlus orcutti. — AS 13373 (CS).
Eleutherodacttjlus ornatissimus. — KU 119749
(CS).

Eleutherodacttjlus palmatus. — KU 41038.

Eletitherodactylus pahneri. — KU 110913
(CS), 110923 (CS), JDL S-242, 397.

Eleutherodacti/lus pant oni.— AS 13494 (CS),
13747 (CS).

Eleutherodacttjlus parvus.— KV 92834 (CS).

Eletitherodactt/lus patriciae. — KU 79770-71
(CS), 79794 (CS).

Eleutherodacttjlus pertivianus. — ( paratype of
Smmthilltis pertivianus) AMNH not catalogued
(CS).

Eletitherodactylus pictissimus. — AS X2750
(CS),X2813 (CS).

Eleutherodacttjlus planirostris. — KU 92656
(CS), JDL S-230, 245, 1068.

Eleutherodacttjlus podociferus. — KU 41049,
68266 (CS), 80644 (CS), 103017-24 (CS).

Eleutherodacttjlus portoricensis. — AS 11710
(CS).

Eleutherodactylus punctariolus. — KU
117363.

Eleutherodacttjlus pygmaetis. — KU 103025
(CS), JDLS-15, 25, 1311, UIMNH 15141 (CS),
16132 (CS), 40335 (CS), 49268 (CS), 49275
(CS).

Eleutherodacttjlus rhodopis. — UIMNH 14729
(CS), 47996 (CS), 49192 (CS), 49194 (CS),
49211 (CS).

Eleutherodactylus richmondi. — AS 12623
(CS).

Eleutherodacttjlus ricordi.— AMNH 63439
(CS ), 63450 (CS).

Eletitherodactifltis ridens.—KU 102996-97
(CS).

Eleutherodacttjlus rugulosus. — KU 59877-79,
84891, JDL S-17, 67, 1006, 1238-51, UIMNH
14756 (CS).

Eleutherodacttjlus rtithae. — AS V4237 ( CS ) .

Eleutherodactylus spattilatus. — KU 87781
(CS).

Eleutherodactylus stdcatus. — KU 100355,
124227 (CS).

Eleutherodactylus surdtis. — JDL S-268, 357-
58, 360.

Eleutherodacttjlus talamancae. — KU 68267-
68 (CS), 117364.

Eleutherodacttjlus trepidotus. — KU 118134-
35 (CS), UIMNH 55874 (CS).

Eleutherodactylus unistrigatus. — KU 111136-
37 (CS), JDL S-263, 269, 332, 340, 392-95,
530-31.

Eleutherodacttjlus variabilis. — KU 109094
(CS).

Eleutherodacttjlus venancioi. — KU 92839
(CS).

Eleutherodactylus tveinlandi. — AS V1718
(CS), V2466 (CS).

Eleutherodactylus lohtjmperi. — JDL S-248-
51, 270, 446.

Eleutherodacttjlus ivighmanae. — AS 12704
(CS), 12739 (CS).

Eleutherodacttjlus tv-nigrum. — KU 119857
(CS).

LYNCH: LEPTODACTYLOID FROGS

227

Eleutherodactylus ;:(/^/.— AMNH 63269-70

(CS).

Euparkerella brasiliensis. — KU 93192 (CS).

Exipsophiis juninensis. — MCZ 24360 (CS).

Eupsophtis wseiis.—AMNH 22104, KU
84731 (CS).

Ghucrtia russcUi.—\W AM R22920 (CS).

Heleioponis albopunctatus. — UMMZ not cat-
alogued.

Heleioponis australiacus. — AMNH 59491.

Heleioponis eijeri. — UMMZ 124504.

Heleophryne natalensis. — KU 105925 (CS).

Holoaden bmdei.—KU 92868 (CS), 107087-
88 (CS).

Hijdwlaetare schmidti. — KU 110613.

Hiilactophnjnc aupusti.—KU 56187, 56192
(CS),UMMZS-2695.

Htjlodes aspera.—KV 92870 (CS), 92875.

Hijlodes latemtnf>ata.—KV 92878 (CS).

Hijlodes magalhaesi. —KU 92887 ( CS ) .

Hijlodes nasus.—KU 74209 (CS), 92893-94,
JDL S-255-56, 258, 282.

Htjlodes ptdcher. —KU 92899 (CS), 92900.

Hijlorina .sy/uafica.— BMNH 91.29.17,
FMNH7102 (SRC), 7107 (SRC).

Ischnocnema quixensis. — KU 104388,
UIMNH 59643 (CS).

Kyarramis sphagnicola. — AMNH 64294, KU
1103.31 (CS).

Lechriodiis fletcheri.— AMNH 59488, CAS
82221 (CS).

Lepidohatrachus asper. — KU 80783.

Leptodactylus albilabris. — UMMZ S-166.

LeptodacttjJus bolivianus. — KU 41026.

Leptodactylus bufonhis. — KU 92905.

Leptodactylus chaquensis. — KU 80795.

Leptodactylus gracilis. — KU 92913-14.

Leptodactylus hylaedactylus. — KU 119387-
88 (CS).

Leptodactylus labial is. — KU 41027, 68273-
74 (CS).

Leptodactylus macrosternus. — KU 92919-20.

Leptodactylus melanonotus. — KU 68275-76
(CS), JDL S- 1252-58, UMMZ S-858, 1045.

Leptodacttjlus vnjstaceus. — KU 92932.

Leptodactylus mystacinus. — KU 92925-26.

Leptodactylus pentadactylus. — KU 41028-
29, 68159, 84981-82, 117366-68.

Leptodactylus podicipinus. — KU 92938.

Leptodacttjlus prognathus. — KU 80824 (CS),
92944.

Leptodacttjlus piistidostis. — KU 92947.

Leptodacttjlus quadrivittatus. — KU 41030.

Leptodactylus syphax. — KU 92951.

Leptodactyltis wagneri. — KU 104389-90.

Limnodtjnastes dorsalis. — KU 93553, UMMZ
S-165.

Limnodtjnastes fletcheri. — KU 93559 ( CS ) .

Limnodtjnastes peronii. — KU 93566 ( CS ) .

Limnodynastes tasmanensis. — AMNH 60589,
KU 93573-74 (CS).

Limnomedusa macroglossa. — KU 92960 (CS),
92961.

Lithodtjtes lineattis.—KU 104340 (CS).

Megaelosia goeldi.—KU 92965-66, 106271.

Metacrinia nichollsi.—KU 110332 (CS).

Mixophtjes fasciolatus. — KU 56627.

Mtjobatrachus gotddii.—KU 110333 (CS).

Neobatrachus centralis. — KU 93578.

Neobatrachus /;/cf(/.s.— FMNH 97281, KU
69278 (CS).

Niceforonia festae.—KU 118137 (CS),
USNM 160944 (CS), 160950 (CS).

Niceforonia flacomaculata. — KU 119743
(CS).

Niceforottia montia.— MCZ 24352 (CS).

Niceforonia tvettsteini (paratypes of Etipso-
plitts icettsteini).—NMW 15846:1-2 (SRC).

Notaden bennetti.—FMNH 97658.

Notaden nichollsi.—KU 93580 (CS), 93582
(CS).

Odontophrijnus americanus. — KU 92968,
100437.

Odontophrijnus carvalhoi. — KU 100441-42.

Odontophrijnus cidtripes. — KU 92975.

Odontophrijnus occidentalis. — LHUBA 1200,
1218.

Paratelmatobius lutzi.—KU 92981 (CS),
107089 (CS).

Philoria frosti.—KU 50699 (CS).

Phijsalaemtis albonotatus. — KU 92987 (CS).

Phtjsalaemus biligonigerus. — KU 84768-76.

Phijsalaemus cetUralis.—KU 92993 (CS).

Phtjsalaemus cuvieri. — KU 92999 (CS).

Phtjsalaemus ephippifer.—KU 93005 (CS).

Phtjsalaemus fiiscomaculatus. — KU 80811
(CS), 93010 (CS), UMMZ S-2357 (CS).

Phtjsalaemus gracilis.— KU 93016 (CS).

Physalaemus maculiventris. — KU 93022 (CS).

Phtjsalaemus nanus.— KU 93025 (CS).

Phtjsalaemus nattereri.—KU 92844 (CS),
92845.

Phtjsalaemus petersi. — KU 120290 (CS).

Phtjsalaemus pti.stulatus.—KU 118136 (CS).

Phtjsalaemus pttsiulosus.—KU 41031, 68269-
72 (CS), JDL S-1204.

Physalaemus signif ems. —KU 93033 ( CS ) .

Pleurodema bihroni.-YMNYl 3746-47, 3758.
Pleurodema brachtjops.— AMNH 69754, KU
96159, 104318 (CS), UIMNH WLB-725 (CS).
Pleurodema cinerea.—KU 80836, 93038.

Pleurodema diplolistris.—KU 93044 (CS),
UMMZ 108521 (CS), 108895 (6) (CS).

Proceratophrys appendiculata. — KU 93070.
Proceratophrys boiei. — KU 93076.

Proceratophrys cristiceps. — KU 106273,
UMMZ 115658.

Pseudopaludicola ameghini. — KU 93050
(CS).

Pseudopaludicola falcipes.—KU 93056 (CS).

Pseudopaludicola pusilla. — UMMZ 54589(2)
(CS); boliviana topotypes, UMMZ (2), not cata-
logued (CS).

P.seudopaludicola saltica.—KU 93068 (CS).

Fsetidophryne bibroni.—KU 93588 (CS).

228

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

PseiidopJinjne corrohoree. — AMNH 64510-
12.

Scythmphrys- sawayae.—VISNM 125530 (ho-
lotype), not a skeletal preparation; some skele-
tal features were observed through slits in the
skin.

SDiintliillus limhatus.—KV 68684 (CS).

Synhnphtis ^uttilatus. — JDL S-1215.

Synhophus Icpnis— JDL S-992, UIMNH
27130 (CS).

SynJwpJius marnockii. — JDL S-214.

Synhophus poUidus.—KV 80320 (CS).

SyirJwpJitts pipilans. — KU 59950 (CS).

S y rrJiopJius nt J) r imacu hi t ii s. — U I M N H
55313-16 (CS).

Taudactyhis aciitirostris.—K\] 124233 (CS).

rdmatohms hauthaIi.—K\J 72879 (CS),
UMMZ S-164.

Tehnatohhia marmomtus. — UMMZ 68179
(2).

Telmatobius puta^onicus.—K\J 80781 (CS).

Telmatohufo huUocki.—FMNH 23842
(SRC).

Thowpa hdzi.—KU 92850 (CS), 92908
(CS).

Thowpa iuili(iris.—Kn 92855 (CS), 92856.

Thowpa pet wpolitana. —K\J 92862 (CS).

Tomodactyhis alholabris. —K\J 87780 (CS).

Tomodactyhis ^landis. — UMMZ S-963.

Tomodactyhis nitidtis.—Kn 102649 (CS),
JDL S-1308, UIMNII 7830 (CS), 78.32-34
(CS), UMMZ S-2225.

Upewleia riifiosa. — AMNH

109861 (CS).

ZacJiaenus i)aiviihis. — KU

107090 (CS), 107091.

Zacliacnus wseus. — USNM
type).

Zachaenus stejnc^rri. — KU
92747.

13336, KU
9,3082 (CS),
15126 (holo-
92742 (CS),

LYNCH: LEPTODACTYLOID FROGS

229

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1960. A new frog from an Eocene oil-well
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Hecht, M. K. and Estes, R.

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Hoffman, A. C.

1930. Description of the external characters
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1965. Early Miocene anurans from Florida.
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1967. Additional Miocene anurans from
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HOSMER, W.

1962. A new leptodactylid frog of the genus
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Huxley, J. S.

1958. Evolutionary processes and taxonomy

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1958. Comments on the definition of genera,
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1966. The systematics and zoogeography of
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1967. The development of a phylogeny of
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1963. Backbone of Colombia. Mem. Amer.
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1955. Evolutionary trends in the courtship
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Kluge, A. G.

1967. Higher taxonomic categories of gek-

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1969. Quantitati\e phyletics and the evolu-
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KUHN, O.

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This manuscript was submitted as a disserta-
tion in partial fulfillment for the requirements
for the Ph.D. degree in the Department of
Zoology, The University of Kansas, in March,
1969.

■Si^

3 2044 093 361 616

Date Due

BOUND

1972