[66] Carnegie Museum.

[67] Univ. California, Mus. Vert. Zool.

[68] Museum of Natural History, Louisiana State University.

EVOLUTION AND SPECIATION

The history of the genus dates back to the early late Pliocene, but morphological change since then has been slight insofar as can be judged from lower jaws. Baiomys seems to have been relatively conservative also in types of habitat occupied.

According to Wilson (1937:59), the late Pliocene was a time of decided expansion of myomorph rodents, more particularly cricetines. Furthermore, at this time, the climate in the interior basin of southwestern North America presumably was becoming arid, if we can judge from the spread of elements of the Madro-Tertiary flora. Axelrod (1950:266) points out that the drier, continental climate initiated in the early Tertiary probably had its culmination in middle Pliocene time. Some floras of early late Pliocene of the southwestern United States reflect a climate slightly cooler and more moist than the climates of the middle Pliocene. However, late Pliocene times reflect an arid climate. The flora of the southwestern interior basin of North America in early late to late Pliocene was intermediate between the previous grassland floras of the middle Pliocene and the savannah flora of upper Pliocene. Axelrod (loc. cit.) suggests that this intermediate flora of the interior basin of southwestern North America resulted from the folding of the Cascades and uplifting of the Sierra Nevada and Peninsular ranges to the south. The development of these mountains produced greater aridity to the lee of the mountains, thus accounting for the grassland-savannah flora. Pygmy mice probably originated in that time, I judge in México, and moved northward and southward in a grassland-savannah habitat that seemingly existed as far north as what is now Meade County, Kansas (where the Sawrock fauna lived). Further evidence for occupancy of a grassland-savannah habitat by ancestral pygmy mice stems from the distribution of the living species, B. taylori, that at present occupies territory adjacent to parts of the Sonoran and Chihuahuan deserts. B. taylori seems to be morphologically more specialized for life in an arid grassland than was B. sawrockensis.

The geographic range of ancestral pygmy mice possibly extended farther south in late Pliocene time than the range of B. musculus does now. Anyhow, B. sawrockensis of the early late Pliocene dwelt in a more mesic type of habitat than B. musculus does, and such habitat may have existed from the Pacific lowlands of Central America to the Caribbean lowlands of northern South America (see Duellman, 1958:136, and Dunn, 1940:156) during late Pliocene times. An ancestral stock of hesperomine mice, not greatly different from Baiomys, may have emigrated from the North American continent into South America across the continuous land connection, which Simpson (1950:395) suggests was formed in the Chapadmalalan age (= Blancan age of North American terminology). The length of time of interchange of genes between northern and southern populations of mice across the Central American land connection probably was brief. Duellman (op. cit.:129) pointed out that once the Panamanian portal was closed, the warm counter equatorial current, El Niño, combined with the uplifting of the Andes, began to produce heavy rain forests in Central America and northern South America in late Pliocene or early Pleistocene times. These forests presumably isolated the stock in North America from that in South America where the latter probably evolved rapidly into kinds that differed from one another and from Baiomys in shape of body, type of pelage, and shape of skull. Internal structures such as hyoid apparatus, auditory ossicles, and baculum remained almost unchanged, as for example in Calomys now living in South America. The present resemblance in internal morphological features between it and Baiomys, I judge, reflects taxonomic relationships more accurately than do shape and conformation of body and skull that seem to respond more rapidly to external environmental changes. The cranial characters distinguishing Baiomys musculus from Calomys laucha are as follows: posterior lacerate foramina between second, rather than first, upper molars; parapterygoid fossa shallower; mesopterygoid fossa as wide or wider, instead of narrower, than parapterygoid processes; burr for attachment of superficial masseter muscle hypertrophied instead of well-developed. In other cranial characters studied, the two genera closely resemble each other. Such similarities of crania between Calomys and Baiomys may reflect convergence, but the total of internal and external morphological characters shared, I think reflects true relationships.

Peromyscus has a large number of living and extinct species and exhibits a wide range of morphological variation, whereas Baiomys has a small number (7) of species and exhibits a narrow range of morphological variation. The small number of known species of pygmy mice suggests their conservatism in elaboration of morphological characters. Possibly this is because the habitat, or even the ecological niche, occupied in geological time by these mice was restricted, geographically and in kind. If the habitat of the pygmy mice oscillated between savannah and arid grassland, then an hypothesis can be made possibly accounting for the origin of species of these mice. My idea is that the geographical distribution of Baiomys today reflects a predilection on the part of these mice for a relatively uniform warm climate. Therefore, in the past, in times of warmer continental climate, these mice moved toward favorable habitat northward from an area in central and northern México. In cooler periods, the mice moved southward as habitats to the north became unfavorable.

Dr. W. B. Davis (in. litt.) informs me that B. taylori was uncommon in Brazos County, Texas, approximately 15 years ago, and suggests that the abundance there now of this mouse and my taking it in 1958 northward nearly to the southern border of Oklahoma reflects a definite movement northward. Movement in the same direction in late years has been suggested for the nine-banded armadillo and the hispid cotton rat (Hall, 1959:373) that are associated with warm climates to the south. These movements possibly reflect only minor fluctuations of climate, but in a long period of warmth movements northward would be expected to be pronounced and extensive.

Extinct species of Baiomys may have originated as a result of extension northward of the geographical range and subsequent retreat southward of the northern populations, as follows: (1) the range of the genus moved northward in a warm period; (2) in cooler times, most of the mice in the north disappeared and only isolated colonies remained in small patches of remaining habitat still favorable to the mice; (3) the small populations of isolated pygmy mice after a time changed through mutations, recombinations and subsequent selection to a degree that prevented crossbreeding once populations from the south again moved northward and came in contact with previously isolated stocks; (4) then competition caused further divergence in morphological characters. Such an hypothesis would account for the morphological differences between the extinct B. kolbi and B. rexroadi. The extinct B. brachygnathus, presumably a dweller of a xerophytic grassland, may have had its origin from a B. minimus-like stock in the manner outlined.