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Microhabitat Use and Population Decline in Banner-Tailed Kangaroo Rats

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Microhabitat Use and Population Decline in Banner-Tailed Kangaroo Rats Journal of Mammalogy, 84(3):1031±1043, 2003 MICROHABITAT USE AND POPULATION DECLINE IN BANNER-TAILED KANGAROO RATS PETER M. WASER* AND JAMES M. AYERS Department of Biological Sciences, Purdue University, West Lafayette, IN 47907, USA Numbers of banner-tailed kangaroo rats, Dipodomys spectabilis, have declined sharply in Downloaded from https://academic.oup.com/jmammal/article/84/3/1031/903815 by guest on 24 September 2021 some but not all populations monitored in southeastern Arizona over the past 20 years. We describe concurrent changes in vegetation and report the results of microhabitat manipu- lation experiments in which we removed broom snakeweed, Gutierrezia sarothrae, from 1.00-ha (pilot) or 0.56-ha (replicate follow-up) plots. D. spectabilis became extinct on control plots, but populations remained stable on plots where snakeweed was removed. On a larger scale, declines in numbers of kangaroo rats coincided with increases in density of woody plants. The data substantiate the preferences of this species for structurally open microhabitats and document that survival rates are higher in areas that are more open. Large kangaroo rat species like D. spectabilis are often regarded as keystone species, and our results indicate that they are vulnerable to grassland degradation. Key words: desert scrub, Dipodomys spectabilis, grassland, heteromyids, kangaroo rats, keystone species, microhabitat use, population dynamics, snakeweed Kangaroo rats (Dipodomys spp.) exert and the banner-tailed kangaroo rat, D. spec- strong in¯uences on other members of their tabilis. community. As competitors, they exert po- The large size of banner-tailed kangaroo tent negative effects on sympatric grani- rats makes them effective interference com- vores (Bowers 1986; Brown et al. 1986; Er- petitors (Bowers and Brown 1992; Bowers nest and Brown 2001; Harris 1984; Valone et al. 1987; Brown and Munger 1985; Frye and Brown 1996; Wondolleck 1978). Their 1983), but they have positive effects as preferential harvesting of large-seeded an- well: their larder hoards may create oppor- nuals prevents these plants from excluding tunities for smaller species to pilfer seeds other smaller-seeded species (Brown et al. (Brown and Heske 1990; Price et al. 2000), 1986; Guo et al. 1995; Heske et al. 1993; and their extensive burrow systems create Samson et al. 1992) and thus, indirectly, fa- long-lasting impacts on soil characteristics cilitates ant populations (Davidson et al. (and thus plant communities) and shelter, 1984). Soil disturbance caused by their for- nesting, or estivation sites for species rang- aging may in¯uence composition of plant ing from beetles to burrowing owls (Brown communities (Brown and Heske 1990; Cur- et al. 1997; Chew and Whitford 1992; tin et al. 2000; Heske et al. 1993), as does Greene and Reynard 1932; Guo 1996; seed caching and vegetation clipping (Ker- Hawkins and Nicoletto 1992; Moorhead et ley et al. 1997; Schiffman 1994; Valone and al. 1988; Moroka et al. 1982; Mun and Thornhill 2001). These effects are particu- Whitford 1989; Seastedt et al. 1986). larly well documented for larger species, Species of large kangaroo rats have been such as the giant kangaroo rat, Dipodomys subject to marked population declines in ingens, the desert kangaroo rat, D. deserti, many areas (Goldingay et al. 1997; Single * Correspondent: [email protected] et al. 1996; Williams and Germano 1992). 1031 1032 JOURNAL OF MAMMALOGY Vol. 84, No. 3 In some cases, declines are related unam- tabilis spent less time near shrubs than did biguously to anthropogenic modi®cation of their smaller sympatric congeners, D. mer- habitat, but in others, causes are less clear. riami and D. ordii (Schroder 1987). For example, between summer 1983 and Mounds and burrow systems that D. spec- winter 1984 a population of D. spectabilis tabilis occupied more continuously had in the San Simon Valley in southeastern Ar- fewer shrubs within 5 m (Jones 1982). izona declined by 80%, from the 2nd most In this study, we asked whether the con- common rodent on the site to one of the version of grassland to desert scrub caused rarest. Valone et al. (1995) summarized ev- declines in numbers of D. spectabilis.We idence that the decline was precipitated by used 2 sources of data: 1st, censuses of D. Downloaded from https://academic.oup.com/jmammal/article/84/3/1031/903815 by guest on 24 September 2021 an extreme climatic event, a tropical storm spectabilis and of woody plants on 2 long- that dropped half the average year's precip- term study sites, 1 immediately adjacent to itation on the study site during a single the Brown±Valone site and the other in a week. The negative effects of heavy rainfall nearby area in which vegetation remains could have been mediated by disease, by open grassland; and 2nd, experimental ma- premature spoilage or germination of lar- nipulations of woody vegetation, speci®cal- der-hoarded seeds, or by excessive fungal ly removal of broom snakeweed, Gutierre- growth in seed stores, resulting in the pro- zia sarothrae. Preferences for open micro- duction of lethal mycotoxins. Valone et al. habitats among kangaroo rats have been hy- (1995) favored the last of these explana- pothesized to increase survival or fecundity tions. None of these possibilities, however, by enhancing foraging ef®ciency or safety explains why D. spectabilis on the site did from predators. To determine whether mi- not recover, becoming locally extinct in crohabitat preferences have adaptive de- 1995 (Brown et al. 2001; Valone and mographic impacts, we examined the ef- Brown 1996). fects of manipulating woody vegetation on The decline of D. spectabilis at the above immigration, recruitment, and survival of siteÐhereafter the Brown±Valone siteÐco- D. spectabilis and assessed the contribu- incided with a gradual increase in the den- tions of recruitment and survival to changes sity of woody vegetation (Brown et al. in the size of D. spectabilis populations. To 1997; Valone and Kelt 1999). Many species determine whether manipulating woody of kangaroo rat are found preferentially in vegetation in¯uenced food availability as open microhabitats (Bowers et al. 1987; well as microhabitat structure, we also Germano et al. 2001; Kotler 1984; Price monitored responses of herbaceous plants. 1978a, 1978b, 1984; Price and Brown 1983; Reichman and Price 1993; Rosen- MATERIALS AND METHODS zweig and Winakur 1969). Some evidence Long-term population trends.ÐOur popula- suggests that this tendency is more pro- tion data on D. spectabilis come from mark± nounced in larger species, which also tend release±recapture studies begun in 1979. Most to be con®ned on a geographic scale to data we report in this study come from our 64- more open habitats (D. desertiÐBrown and ha ``Portal'' site, the northwest corner of which Harney 1993; Thompson 1982a; D. in- abuts the Brown±Valone site. Historical records gensÐGrinnell 1932; Williams 1987). D. suggest that this area (elevation, 1,350 m) was spectabilis, in particular, has been charac- grassland in l875 (Valone and Kelt 1999). How- ever, deterioration began in the late 1800s, and terized as a species of open grassland in 1944, the area was classi®ed as transitional (Hoffmeister 1986; Vorhies and Taylor between grassland and desert shrub (Darrow 1922). On a local scale, individuals were 1944). In 1979, it was still relatively open, with trapped more often at microsites with lower scattered shrubs (Prosopis glandulosa, Acacia percentage cover (Bowers and Brown 1992; greggi, Flourensia cernua, Ephedra trifurca, Bowers et al. 1987). Radiotracked D. spec- and especially A. constricta) and occasional August 2003 WASER AND AYERSÐPOPULATION DECLINE OF KANGAROO RATS 1033 half-shrubs (especially G. sarothraeÐwe use known alive were as high as or higher than the the term half-shrub to refer to suffrutescent pe- closed-population mark±recapture estimates. rennials, species that are smaller than most We, therefore, report minimum numbers alive. shrubs but woody at the base). The most com- Dipodomys spectabilis breeds during the win- mon grass was Tridens pulchella, but open areas ter, and nearly all individuals breed 1st in the were seasonally dominated by annuals and winter after their birth (Holdenried 1957; Rand- short-lived herbaceous perennials (Jones 1982). all 1991; Vorhies and Taylor 1922; Waser and Our 2nd site, ``Rucker,'' is about 40 km SSW Jones 1991). We report adult survival as the pro- of the Portal site. The Rucker site contains a portion of marked adults in March that survived number of small, relatively discrete populations to the next March. Fecundity was the number of of D. spectabilis; in this study, we discuss data daughters captured in August per adult female Downloaded from https://academic.oup.com/jmammal/article/84/3/1031/903815 by guest on 24 September 2021 from our longest-censused, Rucker 1, popula- in March. Recruitment was the number of new tions, which occupy approximately 24 ha. Com- adult females in March per female in March the pared with Portal, the Rucker site is slightly year before. Nearly 95% of ``new'' adults in higher in elevation (1,600 m), wetter, and on March were animals that were trapped as juve- richer soils. Vegetation is desert grassland, niles the year before; the remainder were adults shrubs are virtually absent, and half-shrubs are (as indicated by descended testes or elongated rare. The commonest grasses are Bouteloua spe- nipples) when initially captured, but given our cies, Hilaria mutica, and Eragrostis lehmanni- high adult capture probabilities, we believe these ana, but areas favored by kangaroo rats are char- were 1-year olds, either immigrants or late-ma- acterized by short grass and forbs interspersed turing juveniles, that we missed in the previous with patches of bare ground. census. Dipodomys spectabilis builds conspicuous Long-term trends in woody vegetation.ÐIn mounds 1±3 m in diameter and up to 0.5 m high, August of 1982, 1988, and 1993, we counted all which makes locating them straightforward shrubs and half-shrubs on 1-m2 quadrats on the (Best 1972; Vorhies and Taylor 1922).
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