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SPHENOPHRYNE - American Museum of Natural History

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94 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 253<br />

paucity <strong>of</strong> the description and poor condition<br />

<strong>of</strong> the juvenile holotype, Microbatrachus<br />

pusillus cannot be identified with any known<br />

BODY FORM AND PROPORTIONS<br />

SEXUAL DIMORPHISM: Typically the sexes<br />

cannot be distinguished externally. The male<br />

subgular vocal sac is not evident except<br />

when the frog is calling. The only male external<br />

secondary sexual character, found in a<br />

some species <strong>of</strong> Austrochaperina, is a tendency<br />

for an elongation <strong>of</strong> the snout that is<br />

paler than the dorsal surface <strong>of</strong> the head. Females<br />

grow slightly larger than males. In 25<br />

species the largest males average 90% <strong>of</strong> the<br />

SVL <strong>of</strong> the largest females, (range 81–<br />

100%). In only one species with an adequate<br />

sample (i.e. Liophryne allisoni) are the largest<br />

male and female <strong>of</strong> equal size.<br />

HANDS AND FEET: Hands range in size<br />

from about 19% to 27% <strong>of</strong> SVL. Smaller<br />

hands appear to be associated with secretive<br />

or burrowing habits, whereas scansorial species<br />

are at the high end <strong>of</strong> the range.<br />

Relative sizes <strong>of</strong> feet in different species<br />

range more widely than do those <strong>of</strong> hands,<br />

from about 33% to 51% <strong>of</strong> SVL. As would<br />

be expected, the burrowing and litter-dwelling<br />

species fall at the lower end <strong>of</strong> the range,<br />

roughly 40% <strong>of</strong> SVL, whereas species<br />

known to be active on the forest floor have<br />

the relatively longest feet. The lengths, both<br />

relative and absolute, <strong>of</strong> the tibia and foot<br />

(and presumably other leg segments not measured)<br />

are typically similar, but some exceptions<br />

are notable.<br />

The relative finger lengths are the same in<br />

all species: 3 4 2 1. The first finger<br />

is typically long; only in Sphenophryne cornuta<br />

is it conspicuously reduced (fig. 52).<br />

The toe lengths <strong>of</strong> all species show no variation<br />

from the presumably primitive order 4<br />

3 5 2 1, and most are webless.<br />

Only one species, Austrochaperina palmipes,<br />

has moderately webbed toes (fig. 56D), while<br />

one other, A. basipalmata, has basal webbing<br />

(fig. 56C). A slight trace <strong>of</strong> webbing appears<br />

in some A. derongo.<br />

The fingertips and toe tips may be narrow<br />

MORPHOLOGY<br />

genus and species, and that future taxonomic<br />

stability will be enhanced if the name is set<br />

aside as a nomen dubium.<br />

and rounded with no trace <strong>of</strong> expansion or<br />

<strong>of</strong> terminal grooves, or they may be broadened<br />

into adhesive organs up to more than<br />

twice the width <strong>of</strong> the penultimate phalanx<br />

and with a prominent terminal groove, or any<br />

degree in between (figs. 49, 51). Intermediate<br />

conditions include discs present on toes but<br />

not on fingers, but never the reverse, or discs<br />

on toes and longer fingers only. The adhesive<br />

pad cells <strong>of</strong> one species, Austrochaperina derongo<br />

(fig. 50), are perhaps more similar to<br />

the cuboidal cells <strong>of</strong> terrestrial Liophryne<br />

rhododactyla (Green and Simon, 1986: fig.<br />

8) than to those <strong>of</strong> scansorial Cophixalus riparius<br />

(Green and Simon, 1986: fig. 9).<br />

Most species have discs on fingers and<br />

toes, typically with those <strong>of</strong> the toes being<br />

distinctly larger. For comparative purposes I<br />

use the discs on the third finger and fourth<br />

toe, which normally are the largest. Two species,<br />

S. cornuta and A. palmipes, have the<br />

largest finger discs. In S. cornuta the finger<br />

disc is markedly wider than the toe disc by<br />

about 1.2. The finger disc <strong>of</strong> A. palmipes<br />

is virtually the same size (relative to body<br />

size) as that <strong>of</strong> S. cornuta, but in contrast the<br />

toe disc is also large, usually almost equal to<br />

the finger disc.<br />

There is a clear correlation between the<br />

state <strong>of</strong> the digital tips and habits. The species<br />

with the largest discs (and most pronounced<br />

T-shaped terminal phalanges; see<br />

Osteology) are, so far as their habits are<br />

known, scansorial: S. cornuta climbs into<br />

shrubs or low into trees when active at night;<br />

A. basipalmata, A. palmipes, A. rivularis,<br />

and A. macrorhyncha are riparian forms with<br />

need for adaptation to traversing slick, sometimes<br />

steep rock surfaces. Those species with<br />

small to intermediate degrees <strong>of</strong> expansion<br />

are terrestrial, active on and within the leaf<br />

litter and in some cases doing limited climbing.<br />

The narrow, rounded tips seen in Oxydactyla<br />

alpestris and O. stenodactyla are associated<br />

with a burrowing mode <strong>of</strong> life.<br />

Most species have subarticular elevations

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