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SPHENOPHRYNE - American Museum of Natural History

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

Fig. 54. Lower surfaces <strong>of</strong> feet and hands <strong>of</strong> Liophryne. A. L. rubra, UPNG 9290. B. L. allisoni,<br />

AMNH A81221. C. L. similis, AMNH A130577. D. L. schlaginhaufeni, AMNH A77589. E. L. dentata,<br />

AMNH A87205. Scale bars marked in 1-mm intervals.<br />

ing skin. More <strong>of</strong>ten the annulus is indistinct<br />

and the covering skin undifferentiated. In a<br />

semiaquatic, riparian species Austrochaperina<br />

palmipes there is no external sign <strong>of</strong> the<br />

ear. I noted no sexual dimorphism <strong>of</strong> tympanum<br />

size in the species studied for this report,<br />

but Tyler et al. (1991) reported that females<br />

<strong>of</strong> an Australian species, A. adelphe,<br />

have a strikingly larger tympanum than do<br />

the males. This is unusual because where dimorphism<br />

is present in frogs, the male typically<br />

has the larger tympanum.<br />

External appearance and variation in the<br />

relative size <strong>of</strong> the tympanic annulus notwithstanding,<br />

the ear in the 13 species dissected<br />

(including examples <strong>of</strong> all four genera)<br />

is complete, with the tympanum distinct<br />

from the overlying skin (fig. 60). Wever<br />

(1985) examined four species <strong>of</strong> three microhyline<br />

genera: Gastrophryne carolinensis, G.<br />

olivacea, Hypopachus variolosus (‘‘cune-<br />

us’’), and Kaloula pulchra. He found that<br />

‘‘All four are closely similar in general ear<br />

structure, lacking a true tympanic membrane<br />

but with an effective substitute in the form<br />

<strong>of</strong> a cartilaginous plate (the pars externa <strong>of</strong><br />

the columella in a flattened form) located immediately<br />

beneath the skin.’’ Earlier authors<br />

(e.g. Parker, 1934) have generally referred to<br />

these species as having a ‘‘hidden tympanum.’’<br />

Presumably the condition Wever described<br />

is derived.<br />

Thirteen species treated in the present<br />

study that I dissected share the primitive<br />

structure <strong>of</strong> a true, albeit concealed, tympanum,<br />

and probably the rest do, too. But Wever’s<br />

description suggests that, in comparative<br />

studies <strong>of</strong> other genera and subfamilies, examination<br />

<strong>of</strong> the ears should be more than<br />

skin deep. For example, Smirnov (1986, fig.<br />

1) illustrated the middle ear apparatus <strong>of</strong> five<br />

species <strong>of</strong> Microhyla: three with a tympanum

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