Microtus Ochrogaster) from the Southern Plains of Texas and Oklahoma
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OCCASIONAL PAPERS Museum of Texas Tech University Number 235 5 May 2004 HISTORICAL ZOOGEOGRAPHY AND TAXONOMIC STATUS OF THE PRAIRIE VOLE (MICROTUS OCHROGASTER) FROM THE SOUTHERN PLAINS OF TEXAS AND OKLAHOMA FREDERICK B. STANGL, JR., JIM R. GOETZE, AND KIMBERLY D. SPRADLING The prairie vole (Microtus ochrogaster) today is with distinct cranial characters andpale pelage colora considered a species ofthe Great Plains, whose range tion, they described M. o. taylori. extends from the interior grasslands ofsouthern Canada to central Oklahoma and the northernTexas Panhandle, Choate and Williams (1978) perfonned a mor and is generally bordered by the Rocky Mountains to phometric analysis ofthe prairie vole, which included the west and the eastern deciduous woodlands (Bee et material reported onby Hibbard and Rinker (1943) as al. 1981; Hall 1981; Hoffmann andKoeppl 1985; Caire M. o. taylori. Each of their external body measure et al. 1990; Stalling 1990). Duringthe late Pleistocene, ments and four of the seven cranial characters (skull the species occurred throughout much ofTexas, leav length, zygomatic breadth, mastoidal breadth, and least ing fossil evidence as far south as the Edwards Pla interorbital breadth) exhibited significant geographic teau and parts ofthe GulfCoast (Dalquest and Schultz variation. While no geographic patterns were evident 1992; Graham 1987). The species retreated north to suggest subspecific levels of differentiation, popu ward with the advent ofincreasingly arid climate dur lations from the Dakotas to Kansas demonstrated a ing the Holocene. At least one isolated population, M. slight north-to-south elinal increase in size of cranial o. ludovicianus, was stranded during this time, and is characters. Among their conclusions were that the known to have survived in extreme southeasternTexas population referred to as M. o. taylori by Hibbard and and adjacent Louisiana until the earliest 1900s (Bailey Rinker (1943) no longer exists. They hypothesized 1905). that either the type locality and vicinity was colonized by adjacent populations following extirpation of the Hibbard and Rinker (1943) reported a series of original taylori stock by drought, or that genetic M. ochrogaster taken in 1942, which extended the swamping enveloped the few remaining drought sur known range ofthe prairie vole southward into Meade vivors. Consequently, in their view, all M. ochrogaster County of southwestern Kansas. The animals were from the Central Plains (which would include all prai found to be restricted to the few bogs and meadows, rie voles of Nebraska, Kansas, Colorado, and Okla having survived increasingpressures ofagriculture and homa) should be assigned to the subspecies M. o. a recent drought. On the basis of these specimens, haydenii. considered by them to represent a relictual population 2 OCCASIONAL PAPER", MUSEUM OF TEXAS TECH UNIVERSITY Since 1986, populations ofthe prairie vole have ofthe late Pleistocene or early Holocene. He referred been reported from south-central Oklahoma (Choate his specimens to M. o. haydeni ongeographic grounds. 1989; Smith 1992a), and the panhandles ofOklahoma Even at a glance, the pelage coloration of these ani (Dalquest et al. 1990a; Reed and Choate 1988) and mals is dark--comparable to reference series from Texas (Choate and Killebrew 1991; Jones et al. 1988; Kansas, Nebraska, and Missouri--and in striking con Manning and Jones 1988). Each ofthese populations trast to the pale pelage of specimens from southern represents marginal extensions to the south ofthe pre Colorado and the panhandle regions ofOklahoma and viously mapped ranges ofthe species. Reed and Choate Texas. (1988) suggested that their Oklahoma panhandle speci mens represented an invasion byM. o. haydenii stock, This studycharacterizes the morphometric varia but others (Dalquest et al. 1990a; Manning and Jones tion ofsouthern peripheral populations ofprairie voles, 1998) have tentatively referred specimens from the and provides the ftrst critical review ofthe species to Oklahoma andTexas panhandle regions to M o. taylorl. include specimens from Texas and Oklahoma. We examine the historical zoogeography and subspeciftc Presence of the prairie vole in southern Okla status of the species from those states, with special homa first was documented by Choate (1989), who attention to the question ofM. o. taylori as an extant considered the south-central Oklahoma prairie vole taxon and its validity as a subspecies. population from the Wichita Mountains to be a relict MATERIALS AND MEmoDs Specimens of Microtus ochrogaster from the lary toothrow) were taken with digital calipers and re collections ofMidwestern State University, Texas Tech corded to the nearest 0.01 mm. We depart from their University, and Oklahoma State University were sur protocol by excluding standard body measurements veyed from southern localities of the species' range. on the basis ofinescapable variations in methodology Of these, 129 specimens representing adult individu and skills of different preparators. als (two oldest of four age classes, sensu Choate and Williams 1978) taken from three populations at the Sexes were pooled for all subsequent analyses southern periphery ofthe species' range in Oklahoma after two-way (sex, age) multiple analysis ofvariance and Texas (Figure 1), and reference series from Colo (MANOVAs) for eachmeasurement revealed that sexual rado (western peripheral population) and Kansas (in dimorphism and interactions between age and sex were terior sample) were selected for ftnal study. Each of not signiftcant. the three nominal subspecies (sensu Hall 1981; Hoffmann and Koepp11985; Stalling 1990) are repre Age variation between the two adult age classes sented in the analyses: M o. taylorl from the Texas in our study was signiftcant for each character except and Oklahoma panhandles; M o. haydenii from south least interorbital breadth and prelambdoidal breadth ern Colorado and southern Oklahoma; and M o. (one-way MANOVAs; P = 0.2987 and P = 0.1779, ochrogaster from eastern Kansas. However, we note respectively), and only the least interorbital breadth that Choate and Williams (1978) have synonymized (one-way MANOVA, P:s 0.0001) among young adult M o. ochrogaster with M o. haydenii, and we follow specimens exhibited signiftcant geographic variation. their revision. Based on these fmdings, further geographic analyses were restricted to four samples comprised of speci Seven cranial measurements described by Choate mens representing the oldest of the four age catego and Williams (1978; length ofskull, zygomatic breadth, ries. Only for the Oklahomaffexas panhandles sample mastoidal breadth, least interorbital breadth, was it necessary to pool individuals from more than prelambdoidal breadth, length of rostrum, and maxil- one collecting locality. STANGL ET AL-PRAnm: VOLE (MICROTUS OCHROGASTER) 3 --- H ;~ --- I 36 , \ 3t- I H f \ 0 300 : + KM N I I 102 96 Figure 1. Southern distribution and superimposed locality records for Microtus ochrogaster. Holocene records from Dalquest et al. (1990b) are indicated by "H". Site represented by skulls examined from owl pellets is indicated by "X". Nominal subspecific taxa (sensu Hall 1981 ) repre sented are: a) M. o. haydenii; b) M. o. taylori; and c) M. o. ochrogaster. Principal components analysis ofthe covariance (MWSU 14219, 14221, 14222, 14224, 14226, 14431, matrix was usedto assess patterns ofgeographic varia 14463, 14474, 14476, 14569, 14632, 14633, 14741, tion between populations, and discriminant functions 14776, 14777, 14854-14856, 14858-14860, 14862, analyses were applied to assess uniformity ofdistinct 14872, 14873, 15018, 15019, 15068, 15069, 15594, ness ofthose samples. Duncan's Multiple Range tests 15741); Mo. ochrogaster-KANSAS: Franklin Co., were used to examine morphometric associations be 1-5 mi. S Ottawa,19 (MWSU 5367, 5368,7176-7180, tween populations. All computations were accom 7191, 7483-7485, 9186, 9188, 9192-9197, 15960); plished with NCSS, Version 5.3 (Hintze 1990). M o. taylori--OKLAHOMA: Beaver, Co., 3.8 mi. W Bryan's Comer, 1 (MWSU 16508); Texas Co., 11.1 Specimens examined.--Following is the list of mi. E Guymon, 1 (MWSU 15674); 4 mi. NE Tyrone, "old adult" individuals (sensu Choate and Williams, 2 (MWSU 15715, 15735); TEXAS: Hansford Co.: 8 1978), listed by nominal subspecific designation, that mi. S Speannan, 4 (ITU52211,52449,52450,52423). comprise the basis for this study: M. o. haydenii- Specimens from Texas are deposited in The Museum COLORADO: Huerfano Co., 4 mi. W Walsenburg, 6 of Texas Tech University (TTU), and all others are (MWSU 18156-18159, 18161,18162); OKLAHOMA: from the Collection of Recent Mammals at Midwest Comanche Co., Fort Sill, Wichita Mountains, 30 ern State University (MWSU). 4 OCCASIONAL PAPERS, MUSEUM OF TEXAS TECH UNIVERSITY REsULTS Each ofthe cranial characters, except zygomatic tion for five ofthe seven measured characters. These breadth in old adult Microtus ochrogaster, exhibited differences are significant (Duncan's Test at P ~ 0.05 highly significant geographic variation between the level) for length ofskull, rostral length, and length of studiedpopulations (Table 1). The interiorsample from maxillary toothrow. Colorado is represented by an as Kansas averaged smaller than any peripheral popula- semblage ofparticularly robust specimens, averaging Table 1. Comparison ofcranial measurements (in mm) of63 "old adult" specimens ofthe prairie vole (Microtus ochrogaster) from four southern populations. Descriptive