University of Kansas Publications
Museum of Natural History

Volume 14, No. 2, pp. 9-27, 1 fig. in text
July 24, 1961

Geographic Variation in the Harvest Mouse,
Reithrodontomys megalotis,
On the Central Great Plains
And in Adjacent Regions

By

J. KNOX JONES, JR. AND B. MURSALOĞLU

University of Kansas
Lawrence
1961

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Robert W. Wilson

Volume 14, No. 2, pp. 9-27, 1 fig. in text
Published July 24, 1961

University of Kansas
Lawrence, Kansas

PRINTED IN
THE STATE PRINTING PLANT
TOPEKA, KANSAS
1961

Geographic Variation in the Harvest Mouse,
Reithrodontomys megalotis,
On the Central Great Plains
And in Adjacent Regions

By

J. KNOX JONES, JR. AND B. MURSALOĞLU

The western harvest mouse, Reithrodontomys megalotis , inhabits most parts of the central Great Plains and adjacent regions of tall grass prairie to the eastward, shows a marked predilection for grassy habitats, is common in many areas, and is notably less variable geographically than most other cricetids found in the same region. R. megalotis occurs (see Hall and Kelson, 1959:586, map 342) from Minnesota, southwestern Wisconsin, northwestern Illinois, Iowa and Missouri westward to, but apparently not across, the Rocky Mountains from southeastern Alberta to Colorado; it is known in Oklahoma only from the Panhandle, thence southward through the Panhandle and Trans-Pecos areas of Texas to southern México, westward across the mountains in New Mexico to the Pacific Coast, and northward to the west of the Rockies to southern British Columbia.

Hoffmeister and Warnock (1955) studied western harvest mice from Illinois, Iowa, northeastern Kansas, Minnesota and Wisconsin, concluded that one subspecific name ( Reithrodontomys megalotis dychei J. A. Allen, 1895, with type locality at Lawrence, Douglas Co., Kansas) applied to all, and relegated Reithrodontomys megalotis pectoralis Hanson, 1944 (type locality at Westpoint, Columbia Co., Wisconsin) to synonymy under dychei . Our study, based upon an examination of 1350 specimens, concerns the area west of the Missouri River from Kansas and Nebraska westward to Montana, Wyoming, Colorado and northern New Mexico. Our objectives were to study variation in R. megalotis in the region indicated and to decide what subspecific names properly apply to populations of the species that occur there.

Aside from the name R. m. dychei , currently applied to western harvest mice from a large part of the region here under study, three other subspecific names need consideration:

" Reithrodontomys aztecus " J. A. Allen, 1893 (type locality, La Plata, San Juan Co., New Mexico), currently applied to specimens from northern New Mexico and southern Colorado (and adjacent parts of Arizona and Utah) east to southwestern Kansas and the Oklahoma Panhandle;

" Reithrodontomys megalotis caryi " A. H. Howell, 1935 (type locality, Medano Ranch, 15 mi. NE Mosca, Alamosa Co., Colorado), proposed for, and currently applied to, harvest mice from the San Luis Valley, Colorado, but possibly a synonym of aztecus according to Hooper (1952:218); and

" Reithrodontomys dychei nebrascensis " J. A. Allen, 1895 (type locality, Kennedy, Cherry Co., Nebraska), proposed for harvest mice from western Nebraska and adjacent areas, but regarded as a synonym of dychei by A. H. Howell (1914:30-31).

Our comments concerning the taxonomic status of these several names appear beyond.

We are grateful to Dr. W. Frank Blair, University of Texas, for the loan of a specimen from the Texas Panhandle (TU), and to Dr. Richard H. Manville, U.S. Fish and Wildlife Service, for the loan of specimens of R. m. caryi from the Biological Surveys Collection (USNM). We are grateful also to persons in charge of the following collections for allowing one of us (Jones) to examine Nebraskan specimens of R. megalotis in their care: University of Michigan Museum of Zoology (UMMZ); University of Nebraska State Museum (NSM); and U.S. National Museum (USNM). A research grant from the Society of the Sigma Xi facilitated travel to the institutions mentioned. Specimens not identified as to collection are in the Museum of Natural History of The University of Kansas. All measurements are in millimeters, and are of adults (as defined by Hooper, 1952:12) unless otherwise noted.

Secondary Sexual Variation

Hooper (1952) did not accord separate treatment to males and females in taxonomic accounts of Latin American harvest mice because (p. 11): "In no species ... does sexual dimorphism in the measurements, if present at all, appear to be sufficient to warrant separating the sexes in the analysis." Hooper did not statistically test the validity of treating the sexes together in R. megalotis . He did test a series of R. sumichrasti from El Salvador, in which he found no basis for separate treatment of males and females.

Some authors (Verts, 1960:6, for instance) have recorded females of R. megalotis as larger than males in external measurements, whereas others (Dalquest, 1948:325, for instance) have recorded males as the larger. In order to learn something of secondary sexual variation, and to decide whether or not to separate the sexes in our study, we compared adult males and females from the southern part of the Panhandle of Nebraska (Cheyenne, Keith, Kimball, Morrill and Scotts Bluff counties) in four external and twelve cranial measurements (see Table 1). The external measurements are those customarily taken by collectors and were read from the labels of the specimens; cranial measurements were taken to the nearest tenth of a millimeter by means of dial calipers, and are those described by Hooper (1952:9-11). Females from our sample averaged larger than males in all external and several cranial measurements, but individual variation greatly exceeded secondary sexual variation in each of these measurements and in no case was the greater size of females statistically significant. Therefore, and because we found no qualitative external or cranial differences between the sexes, males and females have been considered together in each population studied.

Table 1. Analysis of Secondary Sexual Variation in Adult Reithrodontomys megalotis From the Southern Part of the Nebraska Panhandle. For Each Measurement, the Number of Specimens Used, the Average, the Extremes, and One Standard Deviation Are Given.

Pelage and Molt

Western harvest mice that attain adulthood acquire at least three distinct types of pelage in sequence in the course of their development. The first of these, the juvenal pelage, is short, relatively sparse, and characteristically grayish brown. The molt (post-juvenal molt) from juvenal pelage to subadult pelage seemingly occurs at an early age, perhaps frequently before the young leave the nest, as individuals in juvenal pelage are few among specimens studied by us. Judging from study skins alone, the progress of post-juvenal molt in R. megalotis is similar to that described for R. humulis by Layne (1959:69-71). The subadult pelage is thicker, longer and brighter than juvenal pelage and closely resembles the pelage of adults; it differs from adult pelage dorsally in being somewhat duller and in having less contrast between back and sides.

The pelage of adults varies depending on season. In summer the individual hairs are relatively short (5-6 mm. at the middle of the back) and sparse. The over-all color of the dorsum, sides and flanks is brownish to dark brownish, and the venter is grayish. In winter the pelage is dense, long (8-9 mm. at the middle of the back) and lax. The over-all color dorsally in fresh winter pelage in most specimens is paler (more buffy) than summer pelage, the sides are markedly buffy, and the venter is whitish; even the tail is more pilose and more sharply bicolored than in summer. Adults molt, usually completely but occasionally only partially, at least twice a year—once in spring (in May and June in Nebraskan specimens) from winter to summer pelage, and once in autumn (in October and November in Nebraskan specimens) from summer to winter pelage. Of the two molts, the one in spring is most easily discernible because the contrast in color between worn winter pelage and fresh summer pelage is considerably greater than that between worn summer pelage and fresh winter pelage, and because the progress of spring molt is seemingly more regular than that of autumn molt. In spring, molt proceeds posteriorly in a more or less regular line on both dorsum and venter; in most specimens it is completed first on the venter. In autumn, molt is irregular, or at best is coincident over large parts of the body, and frequently is seen only by searching through the pelage with a fine probe or dissecting needle. In both spring and autumn, molt seemingly is delayed in females that are pregnant or lactating.

In both winter pelage and summer pelage, the upper parts have blackish or grayish guard hairs and shorter, more numerous cover hairs. All the cover hairs are gray basally; some have a buffy band terminally and others have a buffy subterminal band with a terminal black tip. The generally darker over-all color of upper parts in summer pelage results (as seen in Nebraskan specimens) from a narrower band of buff on the cover hairs (only approximately one half the width of the band on hairs in winter pelage), a darker buffy band (ochraceous buff rather than pale ochraceous or straw color), and a relative sparseness of the pelage, which allows the gray basal portion of some hairs to show on the surface. The more grayish venter of summer-taken specimens results from much more of the grayish basal portion of the white-tipped hairs showing through than in the longer, denser pelage of winter.

Wear on the pelage seems in general to produce a paler over-all color of upper parts, evidently due mostly to abrasion of the terminal black tip of the cover hairs, but possibly actual fading of the pelage is involved also. Worn winter pelage is especially notable for its paleness; the buffy tones are accentuated and the upper parts, especially posteriorly, may even appear fulvous. The difference in color of upper parts between specimens in worn winter pelage and fresh summer pelage (or for that matter specimens in fresh versus worn winter pelage) from the same locality is greater in our material than the difference between some specimens in comparable pelages from localities more than 500 miles apart.

We have seen no specimens taken in winter in which we could discern that the autumn molt had been incomplete, but three old adult males in summer pelage indicate that spring molt is not always completed. KU 50154, obtained on August 14, 1952, 5 mi. N and 2 mi. W Parks, Dundy Co., Nebraska, has the entire posterior back and sides still in old winter pelage and does not appear to have been actively molting; the entire venter is in summer pelage. KU 50146, obtained on August 22, 1952, 3 mi. E Chadron, Dawes Co., Nebraska, has small patches or tufts of winter pelage remaining on the rump and likewise does not appear to have been actively molting. KU 72085, obtained on October 13, 1956, 4 mi. E Barada, Richardson Co., Nebraska, is in the process of molting from summer to winter pelage, but has tufts of old winter pelage on the rump.

Geographic Variation

Geographic variation, both in color of pelage and in external and cranial dimensions, is less in R. megalotis in the region studied than in most other cricetine species that occur there. Nevertheless, meaningful variation is present. The assumption that variation in R. megalotis paralleled in degree that of other species, Peromyscus maniculatus for example, led to untenable taxonomic conclusions by some previous workers.

Color of Pelage

Color of pelage is remarkably uniform, considering the geographic extent of the area involved, over most of the northern part of the central grasslands. Perhaps this uniformity results partly from the predilection of the western harvest mouse for grassy habitats, for in most areas on the Great Plains the species is restricted to riparian communities, principally along river systems, where soils, cover, and other conditions approximate those of corresponding habitats farther to the east to a much greater degree than do conditions in upland habitats. Differential selective pressure, therefore, theoretically would be less between eastern and western populations of R. megalotis than in an upland-inhabiting species. In any event, specimens from western Nebraska, Wyoming, northern Colorado, and adjacent areas average only slightly paler dorsally than specimens in corresponding pelages from the eastern parts of Nebraska and Kansas, and many individuals from the two areas can be matched almost exactly.

To the southwest, on the other hand, a trend toward paler (pale brownish, less blackish) upper parts is apparent. Specimens from southwestern Kansas and adjacent parts of Colorado and Oklahoma average slightly paler in comparable pelages than specimens from northeastern Kansas and eastern Nebraska, but most specimens from farther southwest, in northern New Mexico and southwestern Colorado, are discernibly, although not markedly, paler than mice from northern and eastern populations.

A "pectoral spot," fairly common in some populations of R. megalotis east of the Missouri River (see Hoffmeister and Warnock, 1955:162-163), is present in only a small percentage of the specimens we have studied, and when present is usually only faintly developed.

External and Cranial Size

Fig. 1. Geographic variation in five measurements of Reithrodontomys megalotis on the central Great Plains. The size of each sample is given, along with total length, length of tail expressed as a percentage of the head and body, length of ear, greatest length of skull, and length of rostrum. The approximate distribution of the species in the region shown and the approximate boundary between the subspecies R. m. aztecus and R. m. dychei also are indicated.

As seen in Figure 1, the tail and especially the ear are longer in mice from New Mexico and adjacent areas than in specimens from northern localities. The ear, only slightly variable in size in the northern part of the region, is markedly longer in the southwest, averaging more than 2 mm. longer in specimens from New Mexico and adjacent southwestern Colorado than in specimens from Nebraska and eastern Kansas; specimens in a zone from central Colorado through southwestern Kansas and adjacent Oklahoma generally have ears of a size between the two extremes. As concerns the tail we note a slight trend toward increasing length (best expressed as percentage of length of body) from north to south throughout the central plains, but in general the trend is more pronounced southwestwardly. Variation in length of tail and length of ear, therefore, appear to be in accord with Allen's Rule. Length of body and length of hind foot seem not to vary significantly in specimens we have studied.

The skulls of specimens examined differed only slightly, except that the rostrum is significantly longer and relatively, if not actually, narrower in specimens from the south and southwest than in mice from the rest of the region under study. The rostrum is longest (average 7.7 mm.) in specimens from the vicinity of the type locality of R. m. aztecus , but is relatively long (7.5-7.6 mm.) in populations from as far north as northeastern Colorado and southwestern Nebraska. An average greater occipitonasal length (greatest length of skull) in specimens from the south and southwest results mostly from the longer rostrum.

Recognition of two subspecies of R. megalotis on the central Great Plains seems justified on the basis of the geographic variation discussed above. One subspecies, for which the name R. m. aztecus is applicable, occurs in the southwest and is characterized by the culmination of trends in the region studied to paler upper parts, longer tail, longer ear, and longer, relatively narrower rostrum—characters that appear at least partly independent of each other as concerns gradation toward the smaller, darker-colored populations to the northward. The latter, while exhibiting some differences in color (slightly paler westwardly) and length of tail (shorter northwardly), stand more or less as a unit in contrast to the mice from the southwest, and represent, in our judgment, a single subspecies, R. m. dychei . The area of intergradation between the two subspecies is relatively broad, considering all the characters mentioned, and assignment of some intergrades is admittedly difficult.

Reithrodontomys megalotis aztecus J. A. Allen

Reithrodontomys aztecus J. A. Allen, Bull. Amer. Mus. Nat. Hist., 5:79, April 28, 1893 (type locality, La Plata, San Juan Co., New Mexico).

Reithrodontomys megalotis aztecus , A. H. Howell, N. Amer. Fauna, 36:30, June 5, 1914.

Reithrodontomys megalotis caryi A. H. Howell, Jour. Mamm., 16:143, May 15, 1935 (type locality, Medano Ranch, 15 mi. NE Mosca, Alamosa Co., Colorado).

Distribution. —Western and southern Colorado, southeastern Utah, northeastern Arizona and northern New Mexico, east to the panhandles of Texas and Oklahoma and to southwestern Kansas.

Remarks. —For comparisons with Reithrodontomys megalotis dychei , geographically adjacent to the northeast, see account of that subspecies.

When Howell (1935:143) named Reithrodontomys megalotis caryi from the San Luis Valley of Colorado he compared it directly only with R. m. megalotis from southern New Mexico and northern Chihuahua. Few adults were available to Howell from the San Luis Valley, accounting for the fact, we think, that the published measurements of caryi average less than those given for R. m. aztecus by Howell ( op. cit. :144) and herein. We have examined 16 of the 23 specimens from Medano Ranch and the single specimen from Del Norte that Howell listed. Unfortunately, none is fully adult. The specimens from Medano Ranch, collected in late October and early November, are mostly in fresh winter pelage or molting from subadult pelage, and closely resemble topotypes of aztecus in comparable pelages. Comparison of skulls of the specimens from Medano Ranch with skulls of topotypes and other individuals of aztecus of approximately equal age indicates that the Coloradan specimens may average slightly smaller and have somewhat shorter rostra. Externally, topotypes of caryi have the relatively long tail of aztecus and approach it in length of ear (measured on dry specimens). To us, they appear to be intergrades between aztecus and dychei , but to bear closer resemblance to the former, and we tentatively regard caryi as a synonym of aztecus . Benson (1935:140) noted that two adult topotypes of caryi were "similar to adult topotypes of aztecus ." Specimens from southern Colorado east of the San Luis Valley, assigned to aztecus , are intergrades between it and dychei , as are two specimens from El Paso County, to the north, which resemble aztecus in color but resemble dychei in other characters and are tentatively assigned to the latter.

Specimens from southwestern Kansas and adjacent Oklahoma, herein referred to aztecus , also are intergrades with dychei . Individuals from Meade County, for example, are intermediate on the average between typical specimens of the two subspecies in color of upper parts (if anything, nearer dychei ), resemble dychei in length of ear, but resemble aztecus in length of tail and rostral proportions (consequently also in length of skull). Although a case could be made for assignment of the specimens from Meade County (and elsewhere in southwestern Kansas) to dychei , they are, everything considered, nearer aztecus , to which subspecies they have been assigned consistently since first reported from the area by Hill and Hibbard (1943:24).

Of two specimens examined from 10 mi. S and 1 mi. W Gruver, Hansford Co., in the Panhandle of Texas, the one adult is clearly assignable to aztecus as is the specimen from 9 mi. E Stinnett, Hutchinson Co., Texas, that was referred to dychei by Blair (1954:249).

Reithrodontomys megalotis aztecus has had a rather unstable taxonomic history. Allen, who originally named the subspecies (1893:79), regarded it two years later (1895:125) as a synonym of R. m. megalotis , the subspecies with geographic range to the south and west of that occupied by aztecus . Howell (1914:30) recognized aztecus as valid, but he, too, questioned its distinctness from megalotis in a later paper (1935:144). Hooper (1952:218), the most recent reviewer, supported the validity of aztecus because specimens available to him averaged "distinctly larger in skull length and size of brain case" than specimens of megalotis . Our comparisons of typical specimens of aztecus with specimens of megalotis from southern New Mexico and southwestern Texas confirm Hooper's observations and indicate also that aztecus has a longer rostrum and slightly longer ear.

Reithrodontomys megalotis dychei J. A. Allen

Reithrodontomys dychei J. A. Allen, Bull. Amer. Mus. Nat. Hist., 7:120, May 21, 1895 (type locality, Lawrence, Douglas Co., Kansas).

Reithrodontomys megalotis dychei , A. H. Howell, N. Amer. Fauna, 36:30, June 5, 1914.

Reithrodontomys dychei nebrascensis J. A. Allen, Bull. Amer. Mus. Nat. Hist., 7:122, May 21, 1895 (type locality, Kennedy, Cherry Co., Nebraska).

Remarks. —From Reithrodontomys megalotis aztecus , geographically adjacent to the southwest, R. m. dychei differs as follows: upper parts averaging darker (especially in summer pelage), owing principally to more suffusion of blackish middorsally; tail slightly shorter; ears markedly shorter, rostrum shorter and relatively broader; occipitonasal length shorter owing to shorter rostrum.

" Reithrodontomys dychei nebrascensis ," named by Allen (1895:122) from Kennedy, Nebraska, was distinguished in the original description from dychei by "slightly larger size, relatively longer ears, and more strongly fulvous coloration." Allen applied the name nebrascensis to harvest mice from Montana south to central Colorado and western Nebraska. Howell (1914:30-31) placed nebrascensis in synonymy under dychei because he found specimens from Kennedy to be "indistinguishable from specimens of typical dychei in comparable pelage." We concur with Howell. Topotypes of nebrascensis that we have examined average only slightly paler than topotypes of dychei in the same pelage (some specimens from each series can be matched almost exactly), and do not differ significantly in any external or cranial measurements. The "fulvous" upper parts of the series from Kennedy (all taken in late April) that was available to Allen resulted from worn winter pelage. We think that Allen was led astray also by his erroneous assumption that geographic variation in color of R. megalotis on the Great Plains paralleled that found in Peromyscus maniculatus . Actually, R. megalotis varies in color much less geographically in the region concerned than does P. maniculatus .

Specimens from the northwestern part of the range of dychei (Wyoming, Montana and western South Dakota), like those from western Nebraska, average slightly paler dorsally than topotypes and other specimens from eastern Kansas and Nebraska (a few approach aztecus in this regard), but do not otherwise differ. Most specimens from northern Colorado, southwestern Nebraska (Hitchcock and Dundy counties) and western Kansas average slightly paler than typical specimens and have longer rostra, approaching aztecus in these particulars, but have the shorter ears and shorter tail of dychei . In general, these intergrades resemble dychei to a greater degree than aztecus and are accordingly assigned to the former. One exception is a series from Muir Springs, 2 mi. N and 2½ mi. W Ft. Morgan, Colorado. Specimens in this series approach typical dychei in color, but resemble aztecus in having long ears and long rostra (average 15.3 and 7.5, respectively, in 13 adults). The specimens from Muir Springs resemble aztecus to a greater degree than dychei , but are assigned to the latter because specimens from farther west and farther south in Colorado are assignable to dychei . Howell (1914:31) earlier noted that specimens from northern and central Colorado were intergrades between the two subspecies.

The geographic range occupied by R. m. dychei (from east of the Mississippi River in Illinois and Wisconsin to the foothills of the Rockies) is large (although not so large as that currently ascribed to R. m. megalotis , which ranges from southern British Columbia to central México). Most other small rodents that occur in the same geographic area occupied by dychei are represented there by at least two subspecies, a dark one in the east and a pale one in the west. Eastern populations of dychei have, it is true, somewhat darker upper parts than mice from western localities, but the differences are slight; also, judging from the literature, the "pectoral spot" is more common in eastern mice.

It should be noted that R. m. dychei probably has extended its range both eastward and westward in the last century as a result of agricultural practices—clearing of land in the east and irrigation in the west.

Table 2. Cranial Measurements of Two Subspecies of Reithrodontomys megalotis.

LITERATURE CITED

Allen, J. A.

Benson, S. B.

Blair, W. F.

Dalquest, W. W.

Hall, E. R., and K. R. Kelson

Hill, J. E., and C. W. Hibbard

Hoffmeister, D. F., and J. E. Warnock

Hooper, E. T.

Howell, A. H.

Layne, J. N.

Verts, B. J.

Transmitted March 30, 1961.