PALEONTOLOGY

The only known, significant, fossil _Proechimys_ comes from deposits in

the limestone caves of Lagoa Santa, Minas Gerais, Brazil. These

deposits, of Late Pleistocene or Recent age, were extensively studied by

P. W. Lund and the results published in a series of French and Danish

papers. F. Ameghino (1934:110) studied another fauna from a deposit of

similar age in the cave of Iporanga, São Paulo, Brazil. _Proechimys_ is

recorded in his account under the inclusive specific name _fuliginosus_.

The molariform teeth of the fossil described by Lund (1841:pl. 21, fig.

14) shows its close relationship to the living form _P. s._ _elegans_

(Lund) which still inhabits the same region. It belongs in the more

specialized subgenus _Trinomys_ which seems to have been derived from

_Proechimys_. _Trinomys_ has the main fold in the molars always greatly

developed and the fold tends to set apart one lamina in the occlusal

surface. The Lagoa Santa fossil, like some specimens of the living

subspecies, has a small main fold in P4. However, the main fold is large

in all upper molars and in the lower molariform teeth which are notably

specialized in the extreme reduction of the number of counterfolds to

only one.

One hypothesis concerning the evolution of the genus is that a more

primitive group of _Proechimys_ lived in all of the Central Plateau of

Brazil in the Pleistocene Time. The climatic conditions at that time

might have been such as to support large forests but, since the

Pleistocene, these climatic conditions may have changed from humid to

the present drier conditions, which support the dominant, savanna,

floral climax. Actually the extinct fauna from the caves includes

animals which have disappeared from the area and now live only in more

humid areas, as for example _Myocastor_, which has shifted to the

lowlands to the west and south.

Possibly climatic changes were responsible for the faunal shift from the

region that is now a plateau in Central Brazil. This climatic change may

have resulted from the gradual uplift of the eastern part of the

continent. This uplift prevents part of the trade winds which come from

the east from carrying the same amount of moisture inland as they did

previously. In fact, the Andean revolution, even if it occurred as late

as Late Tertiary, would have had no perceptible influence on the

amount of water precipitated on the more eastern parts of the continent.

Oliveira and Leonardos (1943:617) point out that after a Cretaceous

submersion of the central part of Brazil, there was a general uplift.

The authors (_op. cit._:689) mention the presence of continental

Cretaceous deposits in the Central Plateau of Brazil, in support of

these changes, and state that "pelo menos em certas zonas do litoral a

elevação do continente prolongou-se até o Pleistoceno."

Berry (1942:373) concluded, among other things, that there was a

southward extension "in South America of equatorial floras in the lower

Miocene," and (_op. cit._:372) that ... "east of the Andean Axis in

the south temperate zone there was a normal mesophytic flora ... instead

... of present day large steppes."

My idea is that a tropical forest still covered the Central Plateau of

Brazil in (early?) Pleistocene times and that populations of

_Proechimys_ of a primitive type, similar to _P. g. steerei_, for

example, lived in that extensive forest-climax. The gradual uplift of

the plateau, however, gradually brought about drier conditions in this

region. As a result a large cliseral change was initiated, which shifted

the forest-climax to the more humid eastern escarpments and lowlands

that were gradually being developed, while the savanna climax was being

established on the plateau. Eventually the effect of the decreasing

moisture was locally accentuated by the erosion of the sandstones

(Oliveira and Leonardos, 1943:690) in northeastern Brazil, thus

depriving it of a natural reservoir of rain water. An arid belt was

developed which now constitutes an efficient geographic barrier to the

distribution of many kinds of animals.

One marginal species may have shifted eastward with the forest-climax

to effect the Recent distribution. The eastern species became completely

isolated from the main group, accumulated mutations, and evolved into

the subgeneric type _Trinomys_. The generic trend that gave rise to

_Trinomys_ probably remained more stable as far as supraspecific changes

are concerned. The lack of barriers in the distributional area of the

original group favored the dispersal and submergence of mutations and,

therefore, there was but little further supraspecific evolution. The

speciation in both subgenera finally resulted from gradual

differentiation of varying populations since they show combinations of

the generic biotypes and possess few truly qualitative characters.

The cliseral changes in the Central Plateau, which developed the dry

belt, a barrier, might explain the evolution of a few more supraspecific

groups of mammals, as indicated by the presence of similar forms in the

Amazonian region and in Southeastern Brazil. Among these _Echimys_ and

_Phyllomys_, in the same family with _Proechimys_, show differences that

are parallel to those observed in _Proechimys_. One of these parallel

changes is the increased lamination of the cheekteeth. Although

_Echimys_, from the Amazonian region, has upper molariform teeth with

the four laminae fused, _Phyllomys_ has the four laminae completely

separated.

None of the genera known from the Upper Oligocene and Miocene of

Argentine deposits seems to be directly ancestral to _Proechimys_.

Speciation in the Brazilian Spiny Rats · 第 18 / 43 题