[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]

The detection of differences of systematic worth between populations of animals, represented by skins and skulls, is a step preliminary to deducing the factors responsible for the differences. Ordinarily the factors which cause heritable differences have to do with geographic isolation and adaptation to ecological conditions. When differences in the structure of the animal are known, a person is led to speculate on the factors which could cause them. For one thing, does the observed degree of difference tend to isolate animals possessing the "new" character from the other animals? It would seem to me that the isolation once started by one of these differences tends to be accentuated with time and the difference itself thus then becomes a factor responsible for further differentiation.

Whether or not transition from one character to another occurs gradually, in its geographic expression, and thus whether or not intergradation occurs between two subspecies, can be ascertained by the analysis of a series of population-samples appropriately distributed geographically. If two characters of systematic worth are known to blend in one part of the geographic range of a subgenus, and if the same two characters are seen in two other populations, far removed geographically from each other and without any samples of annectent populations to provide actual evidence of intergradation, then such intergradation is to be inferred.