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Title: Subspeciation in the Meadow Mouse, Microtus pennsylvanicus, in Wyoming, Colorado, and Adjacent Areas

Author: Sydney Anderson

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University of Kansas Publications
Museum of Natural History

Volume 9, No. 4, pp. 85-104, 2 figs. in text
May 10, 1956

Subspeciation in the Meadow Mouse,
Microtus pennsylvanicus, in Wyoming,
Colorado, and Adjacent Areas

BY

SYDNEY ANDERSON

University of Kansas
Lawrence
1956

University of Kansas Publications, Museum of Natural History

Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
Robert W. Wilson Theodore H. Eaton, Jr.

Volume 9, No. 4, pp. 85-104, 2 figures in text

Published May 10, 1956

University of Kansas
Lawrence, Kansas

PRINTED BY
FRED VOILAND, JR., STATE PRINTER
TOPEKA, KANSAS
1956

Subspeciation in the Meadow Mouse, Microtus pennsylvanicus, in Wyoming, Colorado, and Adjacent Areas

BY

SYDNEY ANDERSON

INTRODUCTION

In the region including Wyoming and Colorado, Microtus pennsylvanicus has been divided into two subspecies: the pale M. p. insperatus (J. A. Allen) inhabits the Black Hills of the northeasternmost part of Wyoming; the dark M. p. modestus (Baird) inhabits extensive areas in both Wyoming and Colorado. Initial examination of Microtus pennsylvanicus revealed that specimens from the Big Horn Mountains of north-central Wyoming (within the range of modestus as mapped by Hall and Cockrum 1952:407), in color at least, resemble the subspecies insperatus more than they do modestus, and that specimens from southwestern Wyoming are notably dark. Durrant (1952:363) noted that specimens from Utah are dark, and Davis (1939:315) did the same for specimens from near Pocatello, Idaho. It seemed, therefore, that dark color might characterize populations of a wide geographic region and distinguish them from modestus named from southern Colorado. Also, there seemed to be a hiatus of at least 180 miles between the ranges of modestus in northern Colorado and modestus in eastern Wyoming, and an even greater distance separating populations of modestus in northern Colorado from those in western Wyoming. Microtus pennsylvanicus has not been taken in central or southeastern Wyoming despite extensive collecting there, which yielded numerous records of other kinds of Microtus (M. longicaudus, M. montanus, and M. ochrogaster). Subsequent study revealed a pattern of geographic variation within the range now ascribed to modestus which, in my opinion, can be described best by the recognition of three new subspecies.

MATERIALS, METHODS, AND ACKNOWLEDGMENTS

To study geographic variation in color a method was devised as follows: A single skin (KU 42407, from 1½ miles east of Buckhorn in Weston County, Wyoming) was selected as a representative of the paler mice and arbitrarily given the number 2. A single skin (KU 17491, from 3 miles east of Moran in Teton County, Wyoming) was selected as a representative of the darker mice from the western part of Wyoming and arbitrarily given the number 4. These mice were selected so that they were respectively paler and darker than the estimated average of the total variation within the populations to be studied, but the two mice were not at the extremes of paleness and darkness. Comparisons were based on visual inspection of the dorsal pelage as a whole. Skins were compared with these two mice and given whole numbers from one to five. If paler than the standard for 2, the skin was numbered one; if not distinguishably paler or darker, it was given the number two; if intermediate in color to the standards for 2 and 4 and not definitely more nearly referable to one than the other, it was given the number three; if it resembled the standard for 4, it was numbered four; and if darker, it was given the number five. In this manner skins from a given locality could be evaluated one by one and the results plotted, averaged, and treated statistically. On Figure 1 the average values for color of 32 series are mapped to show the geographic variation of color. The following series of adults are the basis for Figure 1 (abbreviations for collections other than at the University of Kansas are included in parentheses): Each locality is followed by the month (or months) of capture, the number of specimens, and the average value for color.

Figure 1. Geographic variation in color in Microtus pennsylvanicus in the Rocky Mountains. Paler colors are represented by smaller numbers. Numbers are derived from the series of specimens listed in the text by the method described there. The subspecies that occur in the region studied are as follows:

In order to study certain variations, the following "method of pairs" was used. Skulls of two series to be compared were matched in pairs so that they corresponded in size and ontogenetic stage of development. Then the two skulls of each pair were examined for differences in each of the following features: size of circle inscribed by the upper incisor teeth, width of nasal bones relative to their length, curvature of the zygomatic arch, elongation of the braincase relative to its width when viewed from the dorsal aspect, degree of indentation in the anterior edge of the zygomatic arch near the rostrum, degree of depression of the nasal bones when viewed from the side, width in the vertical plane of the zygomatic arch at the suture between the maxillary and jugal bones, length relative to width of the prominent fenestra in the posterodorsal part of the squamosal bone, size of the meatus of the auditory canal, distance between the internal margin of an occipital condyle at its posteriormost point and the tip of the paraoccipital process of the same side of the skull, size of the foramen magnum, vertical height of the supraoccipital bone from the dorsalmost point on the margin of the foramen magnum to the midpoint of the lambdoidal crest, constriction posteriorly or narrowness of the incisive foramen relative to its length, distance across the premaxillary bone from the anteriormost point of the incisive foramen to the posteriormost point of the margin of the alveolus of the upper incisor, area of the maxillary septum (Howell 1926:112, or "zygomatic plate" of Ellerman 1941:1), acuminateness of the anterior border of the palatine opening (internal nares), size of auditory bullae, size of foramen ovale, acuteness of the angle between the basioccipital and basisphenoidal bones at the suture between them (degree to which the area of the suture is raised between the bullae when viewed from the ventral aspect), width of first upper molar tooth, least distance between alveoli of first upper molars. Any differential feature present in more than 75 per cent of the pairs of animals is reported in the discussion of the subspecies concerned. The significance of each difference reported was calculated by the Chi-square test and the confidence limit is given in each case. The probability used in the Chi-square formula is one-half of the percentage of all pairs compared in which the skulls were different in regard to the character being considered. For example, in 68 per cent of the total number of pairs of skulls compared in this study a difference in the size of the auditory bullae was noted. Therefore the probability that a specified skull of a pair will have larger bullae than the other skull was taken as 34 per cent. A different probability for each feature compared was derived in like manner.

Figure 2. Distribution of the subspecies of Microtus pennsylvanicus in Wyoming and Colorado. Solid dots represent localities from which specimens have been examined, and triangles represent localities reported in the literature from which I have not examined specimens. The question mark in southern Colorado denotes a questionable record discussed in the text.

ACCOUNTS OF SUBSPECIES

Microtus pennsylvanicus modestus (Baird)

Remarks.—The dividing line between M. p. modestus and the subspecies to the north on Figure 2 is drawn somewhat arbitrarily because few specimens are available from this area. Actual intergradation, in the form of a geographically intermediate population also morphologically intermediate between these two subspecies, is lacking. However, in most populations of both subspecies some individuals are intermediate between the two subspecies or even more like the other subspecies than the one to which they are referred. Warren (1942:226) states that modestus has been recorded from Lake County, although no reference to a specimen is given. Bailey (1900:21) cites Twin Lakes, in Lake County. That county is near the dividing line as I have drawn it, and therefore specimens from Lake County would be of special interest. An isolated colony of modestus occurs at San Rafael in Valencia County, New Mexico (Bailey, 1932:201). A hiatus of approximately 150 miles separates that colony from the southernmost locality shown in Figure 2. A single specimen, the skin of an immature Microtus without skull, from Trinchera, Colorado, taken by L. R. Hersey in 1912, is in the Colorado Museum of Natural History. No species of Microtine has been recorded from within 50 miles of this locality. The specimen is seemingly more like M. pennsylvanicus than any other species of Microtus. This locality is represented in Figure 2 by a question mark. Ecologically M. montanus fusus Hall and M. longicaudus mordax (Merriam) in the same region occupied by modestus seem to be more montane than modestus. It favors lush grass on the wet floors of alluvial valleys, and also irrigated areas such as that at Manassa.

Microtus pennsylvanicus uligocola new subspecies

Remarks.—M. p. uligocola is more closely restricted to wet situations than M. ochrogaster haydeni (Baird) whose general range lies to the eastward. The numerous lakes, the continuous supply of water from the mountains, and the irrigation systems at lower altitudes along the eastern base of the mountains provide the conditions to which uligocola is suited. It is named for its predilection for water.

The variability in color is relatively greater in topotypes of both uligocola and of modestus than in many of the other series studied. Specimens from Denver and Colorado Springs taken in late autumn and winter (October to February) are paler, more reddish and less blackish, than specimens taken in June and July at Loveland. This reddishness results from longer, and more intensely reddish tips of the hair. The entire hairs also are longer. The average weight of 16 adults (12 males and 4 non-pregnant females) from near Loveland is 49.7 gms. The average length of the ear is 13.4.

Microtus pennsylvanicus finitus new subspecies

Remarks.—The species, Microtus pennsylvanicus, in Pleistocene time ranged onto the plains of Kansas as far southward as Meade County, Kansas (Hibbard, 1940:421). This occurrence indicates a cooler more humid climate then than now in southwestern Kansas. M. p. finitus is more closely associated with water than Microtus ochrogaster, the only other species of Microtus now occupying the same region, although both species have been captured at certain places in the same runways. In Nebraska, a marginal part of the range of the species, M. pennsylvanicus has been taken at scattered localities. This scattered and localized distribution of suitable habitat undoubtedly limits gene-flow between these relict populations. Presumably as a result of this isolation finitus has accumulated and maintained its distinctive characteristics. The subspecies is so named because of its limited range. The average weight of eight specimens (4 males and 4 non-pregnant females) from Dundy County is 57.2 grams.

Microtus pennsylvanicus pullatus new subspecies

Remarks.—In this subspecies there is a cline in color from dark in extreme southwestern Wyoming to pale in north-central Wyoming and Montana as the range of M. p. insperatus is approached. There is thus a broad zone of intergradation in color and the line separating the subspecies must be drawn somewhat arbitrarily. In Wyoming the most distinct break in this cline is in the Big Horn Basin and if a detailed study of the species were made in Montana the break would probably be found where the mountains meet the plains, roughly as shown in Figure 1. There is a similar cline in western Montana in color. The mice are paler farther north as one approaches the Canadian border although they do not become so pale as insperatus. Darkness is a characteristic of several non-adjacent subspecies of Microtus pennsylvanicus, for example M. p. kincaidi Dalquest in central Washington (Dalquest, 1948:347), M. p. finitus, and M. p. nigrans Rhoads in eastern Virginia, but these subspecies presumably differ in other characters.

Some morphological features of the same kind and degree that differentiate subspecies in one place may not vary geographically in another place. Furthermore the geographic variation in one feature may be only partly correlated with the variation in another feature. The variation in M. p. pullatus is an example: Specimens from near Pocatello, Idaho, are darker than topotypes of modestus but specimens from Fremont County, Idaho, are indistinguishable from topotypes of modestus (Davis, 1939:315). I have examined a number of mice from the Bitterroot Valley in western Montana and the color value for 12 adults is 2.7. They are slightly but not significantly paler than topotypes of modestus. This is a result of the cline mentioned above and does not indicate relationship with modestus. Some average measurements of 10 skulls from this series are as follows: condylobasilar length, 24.9; zygomatic breadth, 14.4; interorbital breadth, 3.4; lambdoidal breadth, 11.5; prelambdoidal breadth, 9.1; alveolar length of upper molar teeth, 6.5; and depth of braincase, 7.6. Average external measurements of 9 specimens are as follows: total length, 157; length of tail, 36; length of hind feet, 19.4. Mice from the Bitterroot Valley were compared with topotypes of modestus by the "method of pairs," and modestus had a larger foramen magnum in 6 of 6 pairs (Confidence Limit .97) and larger first upper molar teeth in 5 of 6 pairs (C. L. .75).

A comparison of topotypes of pullatus with modestus shows a similar difference in the teeth, modestus being larger, but in the size of the foramen magnum there is no difference. A comparison of the measurements of pullatus (Bitterroot Valley), pullatus (near topotypes), and modestus (topotypes) shows that the two series of topotypes differ significantly in condylobasilar length (of borderline significance), zygomatic breadth and lambdoidal breadth (both of which vary greatly with age), and length of molar series; the specimens from the Bitterroot Valley agree with pullatus rather than modestus in all of these characters. The specimens from the Bitterroot Valley are smaller than pullatus (topotypes) in total length; they more closely resemble modestus in length of tail, and the hind foot is shorter than in either modestus or pullatus (which do not differ significantly). Specimens from western Montana resemble modestus in certain respects but in most respects resemble topotypes of pullatus and are referred to pullatus. Specimens from Coeur d'Alene, Idaho, and from Blackfoot, Montana (marginal records of modestus, Hall and Cockrum, 1953:410) may be referred to drummondi. Marginal records of pullatus in Montana to my knowledge are: Florence, Ravalli Co.; Highwood Mtns., Chouteau Co.; 7 mi. NE Hilger, Fergus Co.; 10 mi. NW Park City, Stillwater Co. (all represented by specimens in the USBS). The line separating pullatus and drummondi is tentatively drawn as shown in Figure 1.

Microtus pennsylvanicus insperatus (Allen)

Table 1. Average Measurements (in Millimeters) of Adult
Microtus pennsylvanicus

¹ 29 specimens were used in taking measurements of the skull.

¹ 29 specimens were used in taking measurements of the skull.

Remarks.—Bailey (1900:20) had only 7 specimens from northeastern Wyoming and western South Dakota, of M. pennsylvanicus and thought that Arvicola insperatus Allen (1894:347) was not subspecifically distinct from modestus. Subsequently Bailey (1920:72) had adequate numbers of specimens and described Microtus pennsylvanicus wahema from eastern Montana and western North Dakota. Anderson (1943:92) concluded that wahema was not distinct from insperatus and therefore the name M. p. insperatus (Allen) is applicable to this subspecies. On the basis of specimens that I have examined from Montana, South Dakota, and Wyoming I concur with Anderson. Bailey's characterization of wahema is applicable to insperatus as I interpret it. In the Black Hills, Microtus longicaudus longicaudus (Merriam) occurs together with insperatus.

GENERAL REMARKS

The region considered in this paper differs in several regards from the state of Pennsylvania, where variation in the skulls of this species has been studied in detail by Snyder (1954) who referred all populations there to a single subspecies. In some characteristics of the skulls, populations within Pennsylvania differed as much or more than the subspecies from Wyoming and Colorado. In other characteristics of the skulls and of the skins differences are greater between populations in Wyoming and Colorado. The region discussed here is approximately five times as large as the state of Pennsylvania. Populations of M. pennsylvanicus are less continuously distributed than in Pennsylvania owing to major physiographic and climatic barriers and also owing to competition with one or more of the five other species of Microtus occurring in this region. The distribution of three of these species has been discussed by Findley (1945:419). Large areas of relatively greater aridity, such as the region occupied by the subspecies insperatus, occur in Wyoming and Colorado. I have pointed out that the populations which I have designated as subspecies are not absolutely uniform. Also the different subspecies are not of exactly equal degrees of difference. However, there is considerable uniformity of populations occupying conveniently mapped geographic areas. In my opinion, the use of subspecific nomenclature is justified in this case, although not completely unambiguous.

LITERATURE CITED

Allen, J. A.

Anderson, R. M.

Anderson, S.

Bailey, V.

Baird, S. F.

Dalquest, W. W.

Davis, W. B.

Durrant, S. D.

Ellerman, J. R.

Findley, J. S.

Hall, E. R., and E. L. Cockrum

Hibbard, C. W.

Howell, A. B.

Snyder, D. P.

Warren, E. R.

Transmitted June 30, 1955.

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