preceding section, and suppose that it lived in the mesic tropical
environment of the eastern part of the Central American Peninsula (in
what is now Costa Rica and western Panama). Two stocks differentiated,
probably in middle Miocene times; one of these, the ancestral stock of
the _baudini_ group, was widespread on the Caribbean lowlands from the
Nicaraguan Depression to the Bolivar Trough, and the other, the
ancestral stock of the _sordida_ group, was restricted to the Pacific
lowlands of the same region. In late Miocene time the ancestral stock of
the _baudini_ group dispersed northwestward around the deep embayment in
the Nicaraguan depression into upper Central America (in what is now
Honduras and Guatemala) and thence into southern Mexico. Apparently
differentiation took place on each side of the Nicaraguan Depression;
the frogs to the south of the depression evolved into _S. phaeota_,
whereas those to the north of the depression represented the stock from
which _S. baudini_ and _cyanosticta_ arose. Prior to the uplift of the
mountains in the late Miocene and the Pliocene the _baudini-cyanosticta_
stock probably was widespread in northwestern Central America. The
elevation of the mountains resulted in notable climatic changes,
principally the development of sub-humid environments on the Pacific
lowlands. The frogs living on the Pacific lowlands became adapted to
sub-humid conditions and developed into _S. baudini_. The stock on the
Caribbean lowlands remained in mesic environments and evolved into _S.
cyanosticta_.
Possibly in the middle Miocene before the Talamanca Range in Costa Rica
and western Panama was greatly uplifted, the ancestral stock of the
_sordida_ group invaded the Caribbean lowlands of what is now Costa
Rica. The subsequent elevation of the Talamanca Range in the Pliocene
effectively isolated the ancestral stock of _S. sila_ on the Pacific
lowlands from the _puma-sordida_ stock on the Caribbean lowlands. The
former was subjected to the sub-humid conditions which developed on the
Pacific lowlands when the Talamanca Range was uplifted. It adapted to
the sub-humid environment by living along streams and evolving
stream-adapted tadpoles. On the Caribbean side of the Talamanca Range
the _puma-sordida_ stock inhabited mesic environments. The stock that
evolved into _S. puma_ remained in the lowlands as a pond-breeding frog,
whereas those frogs living on the slopes of the newly elevated mountains
became adapted for their montane existence by developing stream-adapted
tadpoles and thus differentiated into _S. sordida_.
Probably the six species of _Smilisca_ were in existence by the end of
the Pliocene; at that time a continuous land connection existed from
Central America to South America. The climatic fluctuations in the
Pleistocene, and the post-Wisconsin development of present climatic and
vegetational patterns in Middle America, brought about the present
patterns of distribution of the species. From its place of origin on the
Caribbean lowlands of lower Central America, _S. phaeota_ dispersed
northward into Nicaragua and southward along the Pacific slopes of
northwestern South America. Perhaps in the late Pleistocene or in
post-Wisconsin time when mesic conditions were more widespread than now,
_S. phaeota_ moved onto the Pacific lowlands of Costa Rica. Its route
could have been through the Arenal Depression. Subsequent aridity
restricted its range on the Pacific lowlands to the Golfo Dulce region.
Climatic fluctuation in northern Central America restricted the
distribution of _S. cyanosticta_ to mesic habitats on the slopes of the
Mexican and Guatemalan highlands and to certain humid areas on the
lowlands. _Smilisca baudini_ was well adapted to sub-humid conditions,
and the species dispersed northward to the Rio Grande Embayment and to
the edge of the Sonoran Desert and southward into Costa Rica. In
southern Mexico and Central America the species invaded mesic habitats.
Consequently, in some areas it is sympatric with _S. cyanosticta_ and
_phaeota_.
_Smilisca puma_ dispersed northward onto the Caribbean lowlands of
southern Nicaragua. Its southward movements probably were limited by the
ridges of the Talamanca Range that extend to the Caribbean coast in the
area of Punta Cahuita in Costa Rica. _Smilisca sila_ dispersed along the
Pacific lowlands and slopes of the mountains from eastern Costa Rica and
western Panama through eastern Panama to northern Colombia. Climatic
fluctuation in the Pleistocene evidently provided sufficient altitudinal
shifts in environments in the Talamanca Range to permit _S. sordida_ to
move onto the Pacific slopes. From its upland distribution the species
followed streams down to both the Caribbean and Pacific lowlands, where
it is sympatric with _S. puma_ on the Caribbean lowlands and _S. sila_
on the Pacific lowlands.
The evolution of the species-groups of _Smilisca_ was effected through
isolation by physical barriers in the Cenozoic; the differentiation of
the species was initiated by further isolation of populations by changes
in physiography and climate. Present patterns of distribution resulted
from Pleistocene and post-Wisconsin climatic changes. Today, sympatric
species have different breeding habits and breeding calls which
reinforce the differences in morphology.
SUMMARY AND CONCLUSIONS
The genus _Smilisca_ is composed of six species of tree frogs; each
species is defined on the basis of adult morphology, larval characters,
and breeding behavior. Keys are provided to aid in the identification of
adults and of tadpoles.
Analysis of the characters and examination of type specimens indicates
that several currently-recognized taxa are synonymous, as follows:
1. _Hyla beltrani_ Taylor, 1942 = _Smilisca baudini_.
2. _Hyla gabbi_ Cope, 1876 = _Smilisca sordida_.
3. _Hyla manisorum_ Taylor, 1954 = _Smilisca baudini_.
4. _Hyla nigripes_ Cope, 1876 = _Smilisca sordida_.
5. _Hyla wellmanorum_ Taylor, 1952 = _Smilisca puma_.
_Smilisca phaeota cyanosticta_ Smith, 1953 is elevated to specific rank,
and one new species, _Smilisca sila_, is named and described.
The skeletal system of developmental stages and the adult of _Smilisca
baudini_ is described, and the skull is compared with that of other
members of the genus.
The tadpoles are described, compared, and illustrated; the larval
development of _Smilisca phaeota_ is described.
Breeding behavior and breeding calls are described and compared. Some
species of _Smilisca_ have breeding choruses. Two species, _S. sila_ and
_sordida_, breed in streams, whereas the others breed in ponds.
The genus is considered to be part of the Middle American Faunal
Element; the species are thought to have differentiated in response to
ecological diversity and historical opportunities provided by Cenozoic
changes in physiography and climate.
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_Transmitted March 14, 1966._
[]
31-3430
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Pp. 121-124. March 7, 1962.
9. Taxonomic status of the free-tailed bat, Tadarida
yucatanica Miller. By J. Knox Jones, Jr., and Ticul
Alvarez. Pp. 125-133, 1 figure in text. March 7, 1962.