[Transcriber's Note: The following error is noted, but left as printed: Page 105, "the must abundant rodent" should be "the most abundant rodent"] UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Volume 14, No. 6, pp. 99-110, 1 fig. December 29, 1961 Natural History of the Brush Mouse (Peromyscus boylii) in Kansas With Description of a New Subspecies BY CHARLES A. LONG UNIVERSITY OF KANSAS LAWRENCE 1961 UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY Editors: E. Raymond Hall, Chairman, Henry S. Fitch, Theodore H. Eaton, Jr. Volume 14, No. 6, pp. 99-110, 1 fig. Published December 29, 1961 UNIVERSITY OF KANSAS Lawrence, Kansas PRINTED BY JEAN M. NEIBARGER, STATE PRINTER TOPEKA, KANSAS 1961 [Illustration] 28-8518 Natural History of the Brush Mouse (Peromyscus boylii) in Kansas With Description of a New Subspecies BY CHARLES A. LONG In order to determine the geographic distribution of the brush mouse in the state, 15 localities, chosen on the basis of suitable habitat, were investigated by means of snap-trapping in the winter and spring of 1959, spring of 1960, and winter and spring of 1961. Variation in specimens obtained by me and in other specimens in the Museum of Natural History, The University of Kansas, was analyzed. Captive mice from Cherokee County, Kansas, were observed almost daily from March 27, 1960, to June 1, 1961. Captive mice from Chautauqua and Cowley counties were studied briefly. Contents of 38 stomachs of brush mice were analyzed, and diet-preferences of the captive mice were studied. Data from live-trapping and from snap-trapping are combined and provide some knowledge of size and fluctuation of populations in the species. Examination of the accumulated specimens and the captive mice reveals the occurrence in southern Kansas of an unnamed subspecies, which may be named and described as follows: #Peromyscus boylii cansensis# new subspecies _Type._--Male, adult, skin and skull; No. 81830, K. U.; from 4 mi. E Sedan, Chautauqua County, Kansas; obtained on December 30, 1959, by C. A. Long, original No. 456. _Range._--Known from 3 mi. W Cedar Vale, _in_ Cowley County, Kansas, and from the type locality. _Diagnosis._--Size medium (see Table 1 beyond); underparts white; upper parts Ochraceous-Tawny laterally, becoming intermixed with black and approaching Mummy Brown dorsally (capitalized color terms after Ridgway, 1912); eye nonprotuberant; tail short but well-haired distally and usually less than half total length; nasals long; cranium large. _Comparisons._--From _P. b. attwateri_, the subspecies geographically nearest _cansensis_, the latter can be easily distinguished by the less protuberant eyes and relatively shorter tail (91 per cent of length of head and body; in topotypes of _P. b. attwateri_ from Kerr County, Texas, 104 per cent; in specimens of _P. b. attwateri_ from Cherokee County, Kansas, 103 per cent). _P. b. cansensis_ is darker than _P. b. attwateri_ and darker than _P. b. rowleyi_, the palest subspecies of brush mouse, which occurs to the westward. The skull and nasals (see Table 1) in adults of _P. b. attwateri_ from Cherokee County average shorter than in _cansensis_. _Specimens examined._--Total, 26. _Cowley Co._: 3 mi. W Cedar Vale, 16. _Chautauqua Co._: type locality, 10. TABLE 1. AVERAGE AND EXTREME MEASUREMENTS OF SPECIMENS OF P. B. CANSENSIS, OF P. B. ATTWATERI FROM CHEROKEE COUNTY, KANSAS, AND OF TOPOTYPES OF P. B. ATTWATERI LISTED BY OSGOOD, 1909. ====================================================================== | _P. b. cansensis_ | _P. b. attwateri_ +-----------+---------+----------+---------+----------- |Three miles| Type | Both |Two miles| Type | west of |locality |localities|south of |locality[A] |Cedar Vale | | | Galena | ---------------+-----------+---------+----------+---------+----------- No. specimens | 11 | 7 | 18 | 20 | 10 | | | | | Total length | 180.5 | 176.7 | 179.1 | 186.2 | 196.0 | 170-199 | 166-188 | ..... | 170-210 | ..... | | | | | Tail-vertebrae | 85.5 | 85.0 | 85.3 | 94.5 | 100.0 | 72-101 | 75-93 | ..... | 83-104 | ..... | | | | | Hind foot | 23.1 | 23.6 | 23.3 | 23.8 | 21.0 | 22-24 | 22-25 | ..... | 22-25 | ..... | | | | | Ear from notch | 18.2 | 19.1 | 18.5 | 18.4 | ..... | 17-19 | 18-21 | ..... | 14-21 | ..... | | | | | Greatest length| 27.9 | 28.3 | 28.1 | 27.8 | ..... of skull | 26.8-29.0 |27.9-28.9| ..... |26.6-29.1| ..... | | | | | Length of | 10.4 | 10.2 | 10.3 | 9.9 | ..... nasals | 9.9-10.8 | 9.5-10.7| ..... | 9.1-10.4| ..... | | | | | Zygomatic | 14.3 | 13.5 | 13.9 | 13.8 | ..... breadth | 13.9-15.0 |13.0-13.9| ..... |13.3-14.4| ..... ---------------+-----------+---------+----------+---------+----------- [Footnote A: From Turtle Creek, Kerr County, Texas, after Osgood (1909:148).] _Distribution of Peromyscus boylii in Kansas_ The subspecies _Peromyscus boylii attwateri_ is known in the state only from Cherokee County, the southeasternmost county in the state. Probably the only locality where the brush mouse occurs in that county is on the systems of cliffs along Shoal Creek, southward from Galena, to the eastward of Baxter Springs. This is the extent of the known range, and in my opinion the probable range, of _P. b. attwateri_ in the state (see Fig. 1). Cockrum (1952:fig. 49) by mistake mapped the species from west of Baxter Springs in Cherokee County. Osgood (1909:149) recorded the subspecies _P. b. attwateri_ from Cedar Vale, Chautauqua County, Kansas, but the specimen from there must now be assigned to _cansensis_ on geographic grounds. Probably the specimen was not obtained from Cedar Vale itself for the habitat is not suitable there. Numerous specimens are known from 3 mi. W Cedar Vale, _in_ Cowley County, Kansas, all of which are assigned to _cansensis_. Osgood's recorded locality is situated between this locality and the type locality of _cansensis_, which is 4 mi. E Sedan, Chautauqua County, Kansas. The distribution of _cansensis_ also is shown in Fig. 1. [Illustration: FIG. 1. Distribution of the brush mouse in Kansas. The southernmost row of counties includes from left to right Cowley, Chautauqua, Montgomery, Labette, and Cherokee. Black dots represent trapping localities from which brush mice were not obtained. Triangles represent localities from which brush mice were obtained. The stippled area contains suitable habitat for the brush mouse, but was not investigated. The easternmost triangle represents a place 2 mi. S Galena, Cherokee Co., Kansas, from which _P. b. attwateri_ is known. The westernmost triangle represents a place 3 mi. W Cedar Vale, _in_ Cowley Co., Kansas, from which _P. b. cansensis_ is known. The triangle of intermediate position represents the type locality of _P. b. cansensis_, a place 4 mi. E Sedan, Chautauqua Co., Kansas. Many of the trapping localities have been investigated more than once.] The probable geographic range of _P. boylii_ is based on trapping data (see Fig. 1). The brush mouse is confined to systems of wooded cliffs in Kansas. The two subspecies seem to be separated by more than 80 miles of grasslands. Blair (1959) has postulated that in the northeastern part of its range _P. b. attwateri_ is represented by disjunct, relict populations formed by diminishing montane or cool, moist environmental conditions. He has implied that the critical climatic change occurred during post-Wisconsin times, and that the isolation of these populations occurred so recently that no morphological differentiation has resulted in them. Inasmuch as the species is widely distributed in México, the southwestern United States, and in California, and has been recorded from the Pleistocene of California (Hay, 1927:323), it is reasonable to suppose that the species immigrated into Kansas from the southwest and that the immigration was in a generally northward or eastward direction. If long tail and large eyes are specializations for a scansorial mode of life (discussed below), then _P. b. cansensis_ must be considered more primitive than _P. b. attwateri_ for the eyes are less protuberant and the tail is shorter in _P. b. cansensis_ than in the latter. I suggest that _P. b. cansensis_ occurred in what is now known as Kansas before _P. b. attwateri_ entered this area by way of the Ozark Mountains. The occurrence of a mouse of "the _truei_ or _boylei_ group" (Hibbard, 1955:213) in southwestern Kansas in the Jinglebob interglacial fauna of the Pleistocene adds little to support the thesis outlined above, but is not inconsistent with the thesis. Incidentally, the geographic distribution of _P. boylii_ may differ somewhat from that shown by Blair (1959:fig. 5); whereas he has mapped the distribution of _P. boylii_ to show disjunctivity in _P. b. attwateri_ and homogeneity in the distribution of other subspecies of the brush mouse to the westward and southward, disjunctivity actually occurs frequently also in the western and southern subspecies. _Ecology_ In Kansas the brush mouse is confined to systems of cliffs, the faces of which range in height to at least 40 feet. The highest cliffs--some approximately 100 feet--on which brush mice are known to occur in Kansas are along Shoal Creek, Cherokee County. The brush mouse is found on low bluffs that are parts of higher systems, but in Cherokee County the mouse was not obtained from low bluffs separated by even a few miles from the cliff-system along Shoal Creek. As implied above the brush mouse is adapted for a scansorial mode of life; but other mice and rats inhabit the rocky crevices of low bluffs. Whereas the brush mouse is well adapted for living on high cliffs it seems that the other rodents are better adapted for life on low cliffs. _Sigmodon hispidus_ was obtained from the low, limestone cliffs mentioned previously. From most low bluffs in southeastern Kansas (and on some high bluffs outside the geographic range of _cansensis_) _Peromyscus leucopus_ was obtained. In Cowley County the brush mouse was abundant when _P. leucopus_ was not and _vice versa_ during this study. _Sigmodon hispidus_ did not associate with the brush mouse in any area, although _S. hispidus_ was often trapped in grassy areas adjacent to cliffs and on the grassy crests of the hills. Except at the locality in Cherokee County, the pack rat, _Neotoma floridana_, was found in association with the brush mouse. _Microtus ochrogaster_ was the must abundant rodent in adjacent southwestern Missouri (Jackson, 1907) before _Sigmodon_ thoroughly infiltrated this area and southeastern Kansas. Activities of other rodents may have confined the brush mouse ecologically to cliffs. Although the grasslands are a barrier to further intrusion by the brush mouse into Kansas, one cannot assume that they alone confined the brush mouse to cliffs. Such an assumption would not explain its absence on systems of cliffs similar to and near other systems of cliffs on which it is found in the non-grassy Ozarkian habitats of Arkansas, as was noticed by Black (1937). Such an assumption would not indicate why the size of the cliff-systems is correlated with the absence or presence of the brush mouse on the northeastern margin of its geographic range. Parasites found on _P. b. attwateri_ include three individuals of the laelapid mite, _Haemolaelaps glasgowi_. Two of these mites were removed from a live mouse. Two larval Ixodid ticks, _Ixodes_ possibly _cookei_, were removed from the pinnae of the ears of a specimen of _cansensis_ from the type locality, 4 mi. E Sedan, Chautauqua County. Four larval Ixodid ticks, _Dermacentor_ possibly _variabilis_, were removed from the pinnae of the ears of a live specimen of _cansensis_ from 3 mi. W Cedar Vale, in Cowley County. TABLE 2. STOMACH CONTENTS OF 38 BRUSH MICE FROM SOUTHEASTERN KANSAS IN WINTER AND SPRING. ====================================================== Localities and | | |Acorn| number of stomachs | Month |Empty| pulp|Seeds -------------------+----------------+-----+-----+----- 2 mi. S Galena | | | | 10 | May, 1959 | 2 | 6 | 2 11 | December, 1959 | 1 | 10 | 0 3 | March, 1960 | 1 | 2 | 0 | | | | 4 mi. E Sedan | | | | 3 | December, 1959 | 3 | 0 | 0 2 | April, 1961 | 1 | 1 | 0 | | | | 3 mi. W Cedar Vale | | | | 6 | December, 1959 | 1 | 3 | 2 3 | December, 1960 | 0 | 3[B]| 0 -------------------+----------------+-----+-----+----- [Footnote B: Judged to be acorn pulp or hickory nut pulp.] Black (1937:195) and Cockrum (1952:180-181) reported stomach contents of _P. b. attwateri_ from Cherokee County containing acorn pulp, seeds, and insects. Analysis of 38 stomachs of the brush mouse (Table 2) show acorns to be the most commonly used food in winter and spring. Seed coats were only rarely found, and insects were absent. Two captive females preferred acorns. Live beetles and grasshoppers of numerous kinds were decapitated and their inner parts eaten. Seeds (wheat, corn, and oats) were also eaten. Inasmuch as acorns appear to be the chief food, it is not surprising that the brush mouse is usually found on cliffs that support stands of blackjack oak (_Quercus marilandica_). Other oaks are present, but I have no evidence that the brush mouse eats their acorns. A. Metcalf told me that he observed in December, 1960, a released brush mouse interrupt its movement toward a hole in a cliff-face along Cedar Creek, Cowley County, in order to pick up an acorn (judged to be from the blackjack oak) in daylight. The mouse carried the acorn into the hole in the cliff. I have observed that captive brush mice eat acorns of the blackjack oak but not some other kinds of acorns. _Behavior_ The chief differences observed between the brush mouse and other species of the genus _Peromyscus_ in Kansas can be summarized as follows: the brush mouse is a superior and more cautious climber; seldom jumps from high places when under stress; is capable of finding its way better in partial darkness; has a stronger preference for acorns; and sometimes buries or hides seeds or acorns. These are all behavioral adaptations that seem in harmony with its mode of life. Buck, Tolman, and Tolman (1925) showed the balancing function of the tail in _Mus musculus_. Climbers (for example, squirrels) often possess long, well-haired tails. It is reasonable to suggest (as did Hall, 1955:134) that the long, tufted tail is an adaptation for a scansorial existence. Little observation is necessary to observe how such a tail is used in balancing. Furthermore, it is used as a prop when the mouse is climbing a vertical surface. Dalquest (1955:144) mentioned tree-climbing in _P. boylii_ from San Luis Potosí, México. It may occur in _P. b. attwateri_ or in _P. b. cansensis_ also, but there is no evidence as yet to prove it. The brush mouse can seldom be induced to jump from heights of two feet or more. Rather it tends to scamper downward or to remain in place. It often swings itself over an edge, holding to it by its hind feet, and sometimes to it lightly with its tail, and reduces a short jump by almost the length of its body. Such caution seems to be an adaptation in a mouse that lives as a climber. Many animals of cavernous habitats have small eyes (see Dobzhansky, 1951:284). Some nocturnal animals (for example, owls) have large eyes. The brush mouse has large, protuberant eyes; it lives in the deep crevices and fissures of the cliffs on which it is found, but it is not strictly a cave-dwelling animal. Perhaps large eyes aid the brush mouse in performing activities in the partial darkness of a deep crevice or hole in a cliff. Brush mice experimentally placed in what appeared to be total darkness fed, built houses of cotton, and ran and climbed in the usual manner. On several occasions the captive brush mice hid surplus seeds and on other occasions hid acorns by burying them and sometimes by placing them in a small jar. The mice never carried the surplus food into their house. Black (1937:195) has claimed that the brush mouse builds a nest similar to that of the nest of the pack rat, _Neotoma floridana_. Hall (1955:134) doubts this to be the case. Dalquest (1953:144) described a nest of _P. boylii_ from San Luis Potosí as seven inches in diameter, made of leaves, and found in a hollow tree. Drake (1958:110) noted that _P. b. rowleyi_ lives in holes and crevices in rocky bluffs in Durango, México. I have found this to be the case for _P. b. attwateri_, as did A. Metcalf (unpublished) for _P. b. cansensis_. Nests of sticks and leaves were taken apart by Metcalf, and all sign indicated only the presence of the pack rat. I have observed that there are no such houses on the cliffs along Shoal Creek, Cherokee County, and that no pack rats have been obtained from there (pack rats have not been reported from Cherokee County). Blair (1938) found two brush mice (_P. b. attwateri_) in the house of a pack rat in Oklahoma. Nests of the brush mice that occur in Kansas have not been found. A lactating, pregnant female (KU 81833) of _P. b. attwateri_, containing three embryos, was obtained on December 24, 1959, and shows that this subspecies breeds in winter. Accumulated records for the subspecies indicates year-round breeding (see Cockrum, 1952:181). Another female obtained on March 27, 1960, was probably lactating. Pregnant females of _P. b. cansensis_ (KU 84892, 84895, and 84890) were obtained from the type locality on April 1-2, 1961, containing 3, 4, and 5 embryos respectively. This indicates, perhaps, increased breeding in spring; five was the highest number of embryos found in brush mice in Kansas. _Population Studies_ In the period of my study the populations of brush mice became smaller, perhaps owing to the severe winter of 1959-1960. In Cowley County, _P. leucopus_ is now abundant and _P. boylii_ rare where in December of 1959, the opposite was true. It is also possible, of course, that trapping has depleted the populations. Conclusions 1. A new subspecies of brush mouse is named and described from southern Kansas. 2. The new subspecies has smaller eyes and a shorter tail and may be more primitive than _P. b. attwateri_. 3. No significant sexual dimorphism was noted in _P. boylii_. 4. In Kansas, _P. b. attwateri_ is known only from a single locality; _P. b. cansensis_ is known from only two localities, both in Kansas. 5. The cliff-dwelling habit of _P. boylii_ probably isolates populations from one another. 6. The grasslands constitute a barrier for the brush mouse. 7. In Kansas, _P. b. cansensis_ probably is an older population than _P. b. attwateri_. 8. In Kansas the brush mouse is confined to systems of cliffs that are wooded and that are at least 40 feet in height. 9. The brush mouse may be confined to cliffs in part by activities of other rodents. 10. The brush mouse commonly associates with the pack rat. 11. Laelapid mites have been found on specimens of _P. b. attwateri_. 12. Larval ixodid ticks were found on specimens of _P. b. cansensis_. 13. Acorns seem to be the chief food of the brush mouse; insects and seeds are also commonly eaten. 14. The brush mouse is adapted for climbing and probably for a partly subterranean life. 15. _P. b. attwateri_ breeds in winter, as well as in other parts of the year. 16. _P. b. cansensis_ is known to breed in early April. 17. The highest number of embryos obtained from a brush mouse in Kansas is five. Acknowledgments I am indebted to Prof. E. Raymond Hall and to Mr. J. Knox Jones, Jr., for suggestions and editorial assistance. Prof. R. H. Camin identified the ticks and mites recorded herein. Mr. A. Metcalf, Mrs. C. F. Long, and Mr. D. L. Long helped with the field studies and in other ways. Literature Cited BLACK, J. D. 1937. Mammals of Kansas. 30th Biennial Report, Kansas State Board of Agri., 35:116-217. BLAIR, W. F. 1938. Ecological relationships of the mammals of the Bird Creek Region, Northeastern Oklahoma. Amer. Midl. Nat., 20:473-526. 1959. Distributional patterns of vertebrates in the southern U. S. in relationship to past and present environment. Zoogeography, pp. 463-464 and Fig. 5, January 16. BUCK, C. W., TOLMAN, N., and TOLMAN, W. 1925. The tail as a balancing organ in mice. J. Mamm., 6:267-271. COCKRUM, E. L. 1952. Mammals of Kansas. Univ. Kansas Publ., Museum of Nat. Hist., 7:6, 180-181. DALQUEST, W. W. 1953. Mammals of the Mexican state of San Luis Potosí. Louisiana State Univ. Studies, Biol. series No. 1, 232 pp. DOBZHANSKY, T. 1951. Genetics and the origin of species, 3d ed. New York, Columbia Univ. Press, x + 364 pp. DRAKE, J. D. 1958. The brush mouse, Peromyscus boylii, in southern Durango. Museum Publ., Michigan State Univ., 1:97-132. HALL, E. R. 1955. Handbook of mammals of Kansas. Univ. Kansas Mus. Nat. Hist. Publ. No. 7, 303 pp. HAY, O. P. 1927. The Pleistocene of the western region of N. America ... Carnegie Inst. Washington, 346 pp., 12 pls. HIBBARD, C. W. 1955. The Jinglebob interglacial (Sangamon?) fauna from Kansas ... Museum of Paleo., Univ. Michigan, pp. 179-228, 2 pls. JACKSON, H. H. T. 1907. Notes on some mammals of southwestern Missouri. Proc. Biol. Soc. Washington, 20:71-74. OSGOOD, W. H. 1909. Revision of the mice of the American genus Peromyscus. N. Amer. Fauna, 28:1-285, April 17. RIDGWAY, R. 1912. Color standards and color nomenclature. Washington, D. C., 43 pp., 53 pls. _Transmitted June 30, 1961._ 28-8518