short-term responses of small mammals to experimental fire in tallgrass prairie

Short-term responses of small mammals to experimental fire in tallgrass prairie

BRYON K. CLARK' AND DONALD W. KAUFMAN Division of Biology, Ackert Hall, Kansas State University, Manhattan, KS 66506, U.S.A.

Received August 1, 1989

CLARK, B. K., and KAUFMAN, D. W. 1990. Short-term responses of small mammals to experimental fire in tallgrass prairie. Can. J. Zool. 68: 2450-2454.

Short-term responses of small mammals to experimental fire were examined in a 13-ha site on the Konza Prairie Research Natural Area, Kansas. A 6.1 -ha portion was burned in spring 1987, whereas the 6.9-ha control site was left unburned in 1987, as was the entire 13 ha in spring 1986. In the absence of fire, < 1 % of all small mammals (n = 130) caught in both March and April 1986 completely shifted from one area to the opposite area. No differences occurred in appearance of new individuals between the two areas in April 1986. In April 1987, all Reithrodontomys megalotis (n = 14) and Microtus ochrogaster (n = 6) caught in the burned area before fire and recaught after fire moved to the unburned area, but no individual from either species moved the opposite way. Most Synaptomys cooperi (75%, n = 4) moved away from the burned area, but no animals were available to test for movements into the burned area. Most new individuals for fire-negative species were captured in the unburned area after fire: 93% of R. megalotis (n = 46), 96% of Blarina hylophaga (n = 27), 100% of M. ochrogaster (n = 4), and 100% of S. cooperi (n = 1). In contrast, Peromyscus maniculatus exhibited a fire-positive response, with one of four moving from unburned to burned and none of five moving the opposite way, and most new individuals recorded after fire were in the burned area (88%, n = 24).

CLARK, B. K., et KAUFMAN, D. W. 1990. Short-term responses of small mammals to experimental fire in tallgrass prairie. Can. J. Zool. 68 : 2450-2454.

Les rCactions a court terme de petits mammiferes a des feux expkrimentaux ont CtC CtudiCes sur une surface de 13 ha dans la rCserve naturelle Konza Prairie Research Natural Area, au Kansas. Une portion de 6,l ha a CtC brQlCe au printemps de 1987, alors que la portion tCmoin de 6,9 ha est restke intacte en 1987, comme I'avait CtC la totalit6 des 13 ha en 1986. Avant le feu, moins de 1% de tous les petits mammiferes (n = 130) capturCes en mars ou en avril 1986 a abandonnC complittement une portion pour migrer dans l'autre. La proportion de nouveaux arrivants Ctait Cquivalente dans les deux portions en avril 1986. En avril 1987, tous les Reithrodontomys megalotis (n = 14) et tous les Microtus ochrogaster (n = 6) capturCs dans la region brQlCe avant le feu et recapturks aprits le feu avaient migrC vers la rCgion intacte, mais aucun individu de I'une ou l'autre espitce n'a fait de migration en sens inverse. La plupart des Synaptomys cooperi (75%, n = 4) ont quittC la zone briiICe, mais il n'y avait pas d'animaux disponibles pour vCrifier les dCplacements vers la zone brOl6e. La plupart es nouveaux arrivants d'espitces craignant le feu ont CtC capturks dans la zone intacte aprits le feu : 93% des R. megalotis (n = 46), 96% des Blarina hylophaga (n = 27), 100% des M. ochrogaster (n = 4) et 100% des M. cooperi (n = 1). En revanche, Peromyscus maniculatus semble prCfCrer les zones briiICes, puisque un individu sur quartre a migrC de la zone non brOlCe a la zone brii1Ce et aucun des cinq autres individus n'a quittC la zone brii1Ce. La plupart des nouveaux arrivants aprits le feu ont CtC capturks dans la zone brulCe (88%, n = 24).

[Traduit par la revue]

Introduction Previous studies have shown that fire typically has a species-

specific impact, either positive or negative, on populations of small mammals in grassland habitats (reviewed by Kaufman et al. 1990). Although these general responses to fire are known, most studies reported differences in abundance between burned and unburned sites several months to years after fire, and little information is available on mechanisms behind changes in population numbers. Two previous studies documented the immediate movements of small mammals after fire and reported that meadow voles (Microtus pennsylvanicus) emigrated from newly burned to unburned prairie (Vacanti and Geluso 1985; Geluso et al. 1986), whereas deer mice (Peromyscus maniculatus) exhibited a net movement from unburned to burned prairie (D. W. Kaufman et al. 1988).

We present data on the postfire movements and numerical responses between burned and unburned prairie for western harvest mice (Reithrodontomys megalotis), Elliot's short-tailed shrews (Blarina hylophaga), prairie voles (Microtus ochrogaster), southern bog lemmings (Synaptomys cooperi), and

' ~ u t h o r to whom correspondence should be sent at the Oklahoma Cooperative Fish and Wildlife Research Unit, Department of Zoology, Oklahoma State University, Stillwater, OK 74078, U.S .A.

deer mice (P. maniculatus). We also present data for net movements and changes in numbers of small mammals for the same area in the absence of fire.

Materials and methods This study was conducted on Konza Prairie Research Natural Area, a

3487-ha ungrazed tallgrass prairie site in northeastern Kansas. Grass- land habitats on Konza Prairie are characterized by tallgrass species, including big bluestem (Andropogon gerardii), Indian grass (Sorghas- trum nutans), switch grass (Panicum virgatum), and little bluestem (A. scoparius; Freeman and Hulbert 1984). Topographic relief con- sists of upland plateaus interspersed with wide valleys. Shallow soils with chert fragments and exposed limestone scarps occur along broad hilltops and hillsides, whereas deep mesic soils occur in lowland areas.

The study site included four watersheds with different prescribed spring fire treatments (Fig. 1A). Watersheds 4A (burned in 1984 and 1980) and 4B (burned in 1987, 1983, and 1979) have been burned at 4-year intervals. Watershed 2A has been burned in odd-numbered years since 1973 and WA has been burned only after wet years (defined as those years with 1.2 times the median expected precipitation from 1 October to 30 September), including 1983, and 1985-1987. The entire area was unburned during our censuses in 1986, although 2A and WA were burned on 1 May 1986. In 1987, watersheds 4B, 2A, and WA, a total of 139 ha, were burned on 8 April. This fire burned the 6.1-ha experimental portion of the study site, leaving the 6.9-ha control site.

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NOTES 245 1

FIG. 1 . Watersheds 4A, 4B, 2A and WA that comprised the 13-ha trapping grid (A), and distance moved by R . megalotis (B, C , and D), M. ochrogaster ( E and F), S . cooperi (G and H ) , and P. maniculatus (I and J) between the last capture in March 1986 or 1987 (a) and first capture in April 1986 or after fire in 1987 (0). The cross-hatched portion denotes the experimental area that was burned in April 1987, and figures without cross-hatching are 1986 data when no fire treatment was applied. Animals recaptured at the same trap location are denoted by half-filled circles. Animals that used both areas, in either March or April, are not included.

The control area was part of 152 ha of contiguous prairie left unburned. on the control area before and after fire (Fig. 1B; Table 3; Small mammals were studied on a 576-station (24 X 24 stations Mann-Whitney U-test; U = 157, n l = 12, n2 = 14, P <

set at 15-m intervals) livetrap grid. One large Sherman livetrap (7.6 X 0.001). In the absence of fire in 1986, distance moved by 8.9 x 22.9 cm) was placed at each station. Animals were trapped on 15- 16 and 29-30 March and 12- 13 and 26-27 April 1986, and on 21-22 and 28-29 March and 12- 13 and 28-29 April 1987. Traps were set on every other row of the grid during each census period and alternate rows were set the following period. Traps were prebaited by placing peanut butter on the open backdoor of traps 2 days prior to each census period. A mixture of rolled oats and peanut butter (wrapped in weighing paper) and a small ball of polyester fiber were placed in each trap when set. Standard capture-mark-recapture techniques were used, and all animals toe-clipped and most rodents ear-tagged as well.

All analyses of possible differences between observed and expected distributions were tested with the log likelihood ratio test (G-test) with Yates correction for continuity, except that Fisher's exact test was used with 2 X 2 contingency tables when sample sizes were small. Expected values were based on the relative sizes of the experimental (6.1 ha) and control (6.9 ha) areas. We also tested for a shift in the proportion of individuals, from before to after fire, with a 2 x 2 contingency table. Proportions of new individuals that first appeared on the experimental and control areas after fire were compared with those expected from the sizes of the two areas and prefire distributions of animals. Similar analyses were used to test for nonrandom distributions of individuals on the two areas, shift in proportion of individuals between March and April, and appearance of new individuals in April 1986 in the absence of fire.

Results Western harvest mice exhibited a fire-negative response

within 3 weeks of the fire (Table I). The proportion of mice that moved from the experimental to the control area after fire was greater than the proportion that moved the opposite way (Table 2; Figs. IB, 1C). This differential movement did not occur in the absence of fire in 1986 as only one of 54 animals moved between the two areas (Table 2; Fig. ID). Minimum linear distance moved by the 14 individuals that shifted from the experimental to the control area in 1987 (Fig. 1C) was greater than the distance moved by the 12 individuals that were caught

animals was not different between the experimental and control areas (Fig. ID; Table 3; P > 0.10). Captures of new animals in April were greater on the control (CO) than experimental (EX) area in 1987, but not in 1986. Recapture success in April of animals that were marked in March was different between years (P < 0.01) but did not suggest fire-induced mortality as the cause (1986: CO = 82%, EX = 74%; 1987: CO = 46%, EX = 52%).

The fire-negative response of short-tailed shrews was due to appearance of new individuals onto the control area following fire as no movements of known shrews occurred between the two areas (Tables 1,2). Neither of the two shrews caught in both March and April 1986 moved between the two areas. Compari- son of recapture success between 1986 and 1987 did not suggest any effect of fire mortality (1986: CO = 14%, EX = 20%; 1987: CO = lo%, EX = 0%).

Prairie voles avoided the burned area as all 12 individuals captured after fire were caught only in the control area (Table I). The proportion of voles moving from the experimental to the control area was greater than the proportion of voles moving in the opposite direction (Table 2; Fig. IE). In the absence of fire, no voles (n = 39) shifted between the two areas (Table 2; Fig. IF). Distance moved by these 39 voles (Fig. IF) was similar to that moved by the two voles caught only on the control area in 1987 (Fig. 1E; Table 3; P > 0.10). Distance moved by all voles remaining on the control was less than that moved by the six voles that shifted from the experimental to the control area after fire (Figs. lE, IF; Table 3; U = 242, nl = 6, n2 = 41, P < 0.001). All new voles captured after fire appeared on the unburned control (Table I). Recapture success was different between the experimental and control areas in 1986 (P < 0.01), but no obvious effects of fire mortality were found (1986: CO = 74%, EX = 35%; 1987: CO = loo%, EX = 75%).

Southern bog lemmings tended to be more numerous on the

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CAN. J . ZOOL. VOL. 68, 1990

TABLE 1. Number of individuals captured on the control (CO) and experimental (EX) areas in March and April 1986 and before (March) and after (April) fire in 1987; new animals were those individuals first captured in April

Marc ha April" New animals

Species Year CO EX pb CO EX pb PC CO EX pb PC

R . megalotis 1986 34 35 1987 26 27

B. hylophaga 1986 7 5 1987 10 10

M. ochrogaster 1986 34 40 1987 2 8 0.07

S. cooperi 1986 7 7 1987 1 8 *

P . maniculatus 1986 12 25 * 1987 8 14

NOTE: Only probability values of P < 0.10 are given. *, P < 0.05; **, P < 0.01. "Does not include those individuals using both areas in March 1986 or March 1987. bExpected values based on the relative sizes of the control (6.9 ha) and experimental (6.1 ha) areas and can be calculated by multiplying the total

number of individuals by 0.53 and 0.47 for the control and experimental areas, respectively. "Expected values based on proportion of individuals captured on the control and experimental areas in March. dFisher's exact test used because of small sample size.

TABLE 2. Capture locations in April 1986 and after fire in 1987 (con- was greater than that expected from sizes of the two areas, but trol = CO; experimental = EX; both areas = BO) for those individuals not from the proportion of individuals caught on the two areas first caught on the control or the experimental area in klarch of each Year (Table 1). None of five deer mice captured on the experimental

area before fire moved from the burned area after fire, whereas First caught First caught one of four mice moved from the control to the experimental on control on experimental area (Table 2; Fig. 1 I). No difference was found in the distance

moved by deer mice caught both before and after fire on the Species Year Co EX Bo Co EX Bo experimental area, control area, and both areas (Fig. 11; Table 3;

R . megalotisb 1986 28 0 0 1 25 0 P > 0. lo), and the same was true in 1986 (Fig. 1J). Distance 1987 12 0 0 10 0 4 **c moved by deer mice was not different between 1986 and 1987

B . hylophaga 1986 1 0 0 0 1 0 (Figs. 1 I, 1 J; Table 3; P > 0.10). Most new deer mice (21124)

1987 1 0 0 0 0 0 captured after fire were on the experimental area, similar to the pattern observed in April 1986 (Tables l ,2) . Recapture success,

M. ochrogaster 1986 25 0 0 0 14 0 although lower in 1987 than 1986 (P < 0.01), did not indicate 1987 2 0 0 6 0 0 ** fire mortality as a major impact (1 986: CO = 92%, EX = 76%;

S . cooperi 1986 3 0 0 0 2 0 1987 0 0 0 3 1 0

P. maniculatusb 1986 10 0 1 0 18 1 1987 3 1 0 0 5 0

NOTE: **, P < 0.01. "Significance levels for differences in proportion of shifts between the two areas. bDoes not include animals using both habitats in March 1986 and March 1987. 'Significance level holds for 10 of lOcomplete shifts of 14 of 14completeplus partial shifts.

control than the experimental area after fire, with a shift in distribution from before to after fire (Tables 1, 2). No shift in distribution from March to April was evident in 1986. Three of four bog lemmings caught on the experimental area before fire and recaptured after fire moved to the control area (Table 2; Fig. 1G). The fourth individual was recaptured on the experimental area once after fire but was then never recaptured. Dis- tance moved by the three individuals that shifted from the experimental to the control area in 1987 (Fig. 1G) was greater than the distance moved by five individuals living on the grid during March and April 1986 (Fig. 1H; Table 3; U = 15, nl = 3, n2 = 5, P = 0.05). The one new bog lemming captured after fire was on the control area (Table 1). Recapture success in 1986 and 1987 did not suggest an obvious fire-mortality effect (1986; CO = 43%, EX = 29%; 1987: CO = 0%, EX = 50%).

After fire, proportion of deer mice on the experimental area

Discussion The short-term numerical responses following fire by

R. megalotis, B. hylophaga, M . ochrogaster, S . cooperi, and P . maniculatus were consistent with the general response to fire reported for these species in a variety of North American grasslands (Kaufman et al . 1990). Although rapidity of the response to fire had been documented for M. pennsylvanicus (Vacanti and Geluso 1985; Geluso et al . 1986) and P . maniculatus (D. W. Kaufman et al. 1988), little was known about the chronology of the fire response for other species. Most studies cited by Kaufman et al. (1990) were point in time samples that occurred several months to years after a fire and could not address the immediate response of small mammals to fire (e.g., Cook 1959; Kaufman et al . 1983). We documented that the fire- negative response exhibited by R . megalotis, M. ochrogaster, B . hylophaga, and S . cooperi occurred within 3 weeks of a fire and although somewhat weaker, the fire-positive response of P . maniculatus occurred during this same period.

Emigration was an important component of the fire-negative response of rodents as nearly all individuals captured in the area to be burned prior to fire moved to unburned prairie after fire (100% of the R. megalotis and M . ochrogaster, 75% of the S . cooperi). No individual for any of these species moved from

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TABLE 3. Median linear distance (m) between trap locations for those individuals captured in both March and April 1986 and before (March) and after (April) the fire in 1987

1986 1987

Control Experimental Shifted Control Experimental Shifted

R . megalotis Distance Range n

M . ochrogaster Distance Range n

S . cooperi Distance Range n

P. maniculatus Distance Range n

NOTE: Does not include individuals that were using both control and experimental areas in March of either year and recaptured in April.

unburned to burned prairie during this time. Emigration also waF important in the fire-negative response of M. pennsylvanicus (Vacanti and Geluso 1985; Geluso et al. 1986). For B. hylophaga, emigration may be a component of the fire-negative response we observed, but our trapping data were insufficient to support or refute this possibility. Emigration was a small component of the fire-positive response documented for P. maniculatus as only 25% of the deer mice captured on the unburned area before fire moved into the burned area after fire (see D. W. Kaufman et al. 1988). In the absence of fire, only 1 of 130 animals of the five common species shifted between the two areas, further suggesting that movements were in response to fire.

Although all individuals on the study site may not have been captured during March of either year, one would expect similar numbers of unmarked individuals to occur on both sides of the study area. However, location of captures for new animals after fire were not random as over 94% of all new individuals of fire-negative species appeared on the unburned area, whereas 88% of the new individuals for fire-positive species occurred on the burned area. In contrast, no differences were noted in the appearance of new animals between the two areas in April 1986 for either fire-positive or fire-negative species. This further supports the notion that the appearance of new animals in April 1987 was influenced by fire.

Recapture success was variable among species, but no obvious fire-induced differences in recapture success were noted for any species. Furthermore, no carcasses of trappable- size individuals were found when the burned prairie was searched. The only fire-induced mortality we documented was two young bog lemmings (<3 g) that were found in an intact spherical nest and probably had died from either heat or asphyxiation. Many fire-negative species use aboveground nests and are probably more vulnerable to fire than fire-positive species. However, adult rodents and shrews, whether fire- positive or fire-negative, are probably infrequently killed by fire except under unusual conditions (Geluso et al. 1986; Erwin and Stasiak 1979; Lawrence 1966).

The only reported exception to the general fire response was that M. ochrogaster were more abundant in burned prairie in Illinois (Schramm and Willcutts 1983). However, M. pennsylvanicus were common in nearby unburned prairie and interac- tions between the two species may have contributed to the fire- positive response of prairie voles. Similar numbers of R . megalotis, B. hylophaga, M. ochrogaster, and S . cooperi were captured on the two areas in spring 1986; however, P. maniculatus were more abundant on the experimental than the control area in March, but not April. This difference was probably due to the 2A and WA portion of the experimental area being burned in 1985.

Avoidance of recently burned prairie by fire-negative species was undoubtedly related to removal of litter as these animals need litter and herbaceous layers for construction of runways and (or) nests (see Birney et al. 1976 for M. ochrogaster; Linzey 1983 for S . cooperi; Webster and Jones 1982 for R. megalotis; Choate and Fleharty 1975 for B. hylophaga). In contrast, P. maniculatus typically nest below ground and prefer areas with little ground cover and vertical structure, conditions created by prairie fires (Clark 1989; Kaufman et al. 1989; G. A. Kaufman et al. 1988). Small mammals in a newly burned area will detect changes in habitat structure within hours of a fire and fire- negative individuals should begin to seek new home sites within hours to days of the fire. For individuals of fire-positive species to move into a burned area, they are not cued by a deterioration in habitat quality in their home range, but by the perception that conditions are better elsewhere. This may result from olfactory cues of a nearby fire (smoke during fire or odors of ash residues left after fire). Differences in ability to detect fire-induced changes in habitat quality may result in dissimilarities in movement patterns, magnitude of numerical changes, and timing of response between fire-negative and fire-positive species.

Acknowledgements This is Contribution No. 89-405-5, Kansas State Agricultural

Experiment Station (KAES), Kansas State University, Manhat- tan, KS. Research was conducted at the Konza Prairie Research

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2454 CAN. J . ZOOL. VOL. 68, 1990

Natural Area, a preserve of The Nature Conservancy, operated by the Division of Biology, Kansas State University. Research was supported by National Science Foundation grants BSR- 830757 1 and BSR-85 14327 and KAES project R-23 1 (D. W.K.) and grants-in-aid from Sigma Xi and American Society of Mammalogists (B . K. C . ) . Konza Prairie Research Natural Area provided B .K.C. with graduate summer fellowships in 1986 and 1987. C . Owensby. T . Seastedt, and C . Smith provided helpful suggestions and criticisms of an earlier version of this manuscript. We thank B. Clark, E. Finck, R . Fridell, and especially S . Bixler for help with the field work. J. Briggs, D. Gibson, and C . Kinderknecht provided computer assistance. Data and documentation forms are stored in the Konza Prairie Research Natural Area Data Bank (data code = ZSM06 and ZSM07).

BIRNEY, E. C., GRANT, W. E., and BA~RD, D.D. 1976. Importance of vegetative cover to cycles of Microtus populations. Ecology, 57: 1043- 105 1.

CHOATE, J. R., and FLEHARTY, E. D. 1975. Synopsis of native Recent mammals of Ellis County, Kansas. Occas. Pap. Mus. Texas Tech. Univ. 37: 1-80.

CLARK, B. K. 1989. Influence of plant litter and habitat structure on small mammal assemblages: experimental manipulations and field observations. Ph.D. dissertation, Kansas State University, Manhattan.

COOK, S. F., JR. 1959. The effects of fire on a population of small rodents. Ecology, 40: 102- 108.

ERWIN, W. J., and STASIAK, R. H. 1979. Vertebrate mortality during the burning of re-established prairie in Nebraska. Am. Midl. Nat. 101: 247-249.

FREEMAN, C. C., and HULBERT, L. C. 1984. An annotated list of the vascular flora of Konza Prairie Research Natural Area, Kansas. Trans. Kans. Acad. Sci. 88: 84- 1 15.

GELUSO, K. N., SCHRODER, G. D., and BRAGG, T. B. 1986. Fire avoidance behaviour of meadow voles (Microtus pennsylvanicus). Am. Midl. Nat. 116: 202-205.

KAUFMAN, D. W., KAUFMAN, G. A., and FINCK, E. J. 1983. Effects of fire on rodents in tallgrass prairie in the Flint Hills region of Kansas. Prairie Nat. 15: 49-56.

KAUFMAN, D. W., GURTZ, S. K., and KAUFMAN, G. A. 1988. Movements of Peromyscus maniculatus in response to prairie fire. Prairie Nat. 20: 225-229.

KAUFMAN, D. W., KAUFMAN, G. A., and FINCK, E. J. 1989. Rodents and shrews in ungrazed tallgrass prairie manipulated by fire. In Prairie pioneers: ecology, history, and culture. Proceedings of the 1 lth North American Prairie Conference, University of Nebraska, Lincoln, 7-1 1 August 1988. Edited by T. B. Bragg and J. Stubbendieck. pp. 173-177.

KAUFMAN, D. W. FINCK, E. J., and KAUFMAN, G. A. 1990. Small mammals and grassland fires. In Fire in North American grasslands. Edited by S. L. Collins and L. L. Wallace. University of Oklahoma Press, Norman. pp. 46-8 1.

KAUFMAN, G. A., KAUFMAN, D. W., and FINCK, E. J. 1988. Influence of fire and topography on habitat selection by Peromyscus maniculatus and Reithrodontomys megalotis in ungrazed tallgrass prairie. J. Mammal. 69: 342-352.

LAWRENCE, G. E. 1966. Ecology of vertebrate animals in relation to chaparral fire in the Sierra Nevada foothills. Ecology, 47: 278-291.

LINZEY, A. V. 1983. Synaptomys cooperi. Mamm. Species, 33: 1-4. SCHRAMM, P., and WILLCUTTS, B. J. 1983. Habitat selection of small

mammals in burned and unburned tallgrass prairie. In Proceedings of the 8th North American Prairie Conference, Western Michigan University, Kalamazoo, 1-4 August 1982. Edited by R. Brewer. pp. 49-55.

VACANTI, P. L., and GELUSO, K. N. 1985. Recolonization of a burned prairie by meadow voles (Microtuspennsylvanicus) . Prairie Nat . 17: 15-22.

WEBSTER, W. D., and JONES, J. K., JR. 1982. Reithrodontomys megalotis. Mamm. Species, 167: 1-5.

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short-term responses of small mammals to experimental fire in tallgrass prairie

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