identifying general patterns in the respons­ Conservation Issues es of arthropod species and arthropod com­ munities to burning. I used a descriptive model of the burn process and postburn succession to aid in organizing the infor­ Responses of Prairie Insects and Other mation. The model allows identification of the information needed to predict the Arthropods to Prescription Burns responses of arthropod species and com­ munities to burning. The conditions undel' which responses to burning vary among Catherine C. Reed sites and among species can be predicted, and possible trade-offs related to manage­ Entomology Department ment for multiple species or groups can be 219 Hodson Hall identified. University of Minnesota St. Paul, Minnesota 55108 USA [email protected] RESULTS AND DISCUSSION

Phases of a Burn ABSTRACT: The results of many field studies provide ample evidence that burning has Warren et al. (1987) reviewed the respons­ a major influence on the presence and persistence of arthropod species on prairie sites. es of grassland arthropods to burning, from The variation in fire tolerance of species and the changes in the physical environment and a pest management perspective, and divid­ plant communities following burns result in the development of distinctly different ed the burn into phases. Burn phases in­ arthropod communities on frequently burned sites compared to sites protected from clude the preburn phase, the combustion burning. Changing successional stages following burns also support distinctive arthropod phase (the burn itself), the shock phase species and groups. In general, a landscape containing sites at different successional (time until the vegetation starts regrowth), stages, and sites varying in burn frequency, will support the most species; if a single site and the recovery phase (time until the sys­ is burned at intervals, a cycle of arthropod species richness, species composition, and tem returns to the preburn state or reaches numbers of individuals will occur. It is possible to predict the responses of a prairie a new steady state). Fire characteristics, arthropod community or of a single species to a controlled burn or a series of burns if the the arthropod species, the timing of the fire history of the site, and the burn tolerance, colonizing ability, and basic biology of the burn relative to arthropod phenology, host! species present, are known. parasite and predator/prey ratios follow­ ing the fire, postburn weather, and the di­ Index terms: insect conservation, fire management, prairie insects; prairie burning rection and extent of postburn succession Natural Areas lournal17:380-385 can influence a species' response to burn­ ing and must be understood in order to predict this response. The mortality, re­ production, immigration, and emigration INTRODUCTION AND METHODS tributions of species among available sites of arthropod taxa during each phase deter­ and wide fluctuations of popUlations from mine the species composition of any site. Whether, when, and how frequently to burn year to year are common among arthro­ prairie preserves have become increasing­ pods. The small size of many prairie pre­ The Preburn Phase ly difficult decisions for managers. Con­ serves has limited the areas available for trolled burns have been used to maintain well-replicated studies. A few groups, pri­ The insect species composition of prairie prairie vegetation and prevent the invasion marily butterflies, have been relatively well areas is poorly known. Tallgrass prairie of woody plants, but concern has devel­ studied, but many taxa have received little sites contain a mixture of rare and com­ oped that these burns may damage or even or no attention from researchers. mon species, including prairie obligates, endanger arthropod species. Many have grassland species, woodland species, and studied the effects of controlled burns on Despite these limitations, many field stud­ widespread species, as observed for bees plants, but arthropod responses to burning ies of arthropod responses to burning have (Reed 1995b); butterflies (Swengel 1996); have not been well understood. Natural been completed in a variety of prairie hab­ features of the prairie arthropod commu­ itats. (A bibliography of 176 published nity challenge researchers. Populations are and unpublished reports and the literature Papers in the Conservation Issues section difficult to census, especially those of spe­ survey on which this paper is based [Reed are reviewed by members of the Editorial cies with short flight periods such as many 1995a] are available from the author.) I Board but are not refereed as are other butterflies and specialist bees. Patchy dis- reviewed these studies with the goals of articles in the Natural Areas Journal.

380 Natural Areas Journal Volume 17 (4),1997 leafhoppers (Hamilton 1995); grasshop­ Wisconsin prairie burn (Reichert and Reed~ sence of one of their predators, Phymata pers, crickets, and katydids (Ballard and er 1970), and crab spiders in Minnesota sp., following a burn. Greenlee 1994); and other insect groups survived an early spring bum (Dana 1991). (Panzer et al. 1995). Insect ~pecies and The Recovery Phase individuals are patchily distributed within Burn tolerance may depend on burn sea­ and among prairie sites (references above). son: for example, Hesperia dacotae and Relative to unburned areas, forbs and grass­ Herbivores, predators, and parasitoids are H. ottoe, prairie skippers (Lepidoptera: es develop and flower earlier in the season present and range from extreme special­ Hesperiidae), showed low larval mortality during the first growing season following ists to extreme generalists in their choice in a test burn early in the season when the a spring burn, and vegetation productivity of host plants, prey, or hosts. Prairie in­ larvae were in burrows; later fires caused increases (Hulbert 1986). Generalist and sects that are dependent on vertebrates, higher mortality because the larvae had specialist plant-feeding insects are attract­ such as dung-feeding scarabs, tenebrion­ moved to surface shelters (Dana 1991). ed to the new growth to feed on leaves, ids that use animal burrows as shelters, sap, or nectar, or for oviposition on plants and vertebrate ectoparasites, are often ab­ Burns are usually patchy; only part of a that are their larval food (Cancelado and sent from small prairie remnants; their con­ site is burned. Savannas and woodlands, Yonke 1970, Nagel 1973, Halvorsen and servation is of special interest (Opler 1981). and areas of heterogeneous topography, Anderson 1980, Van Amburg et al. 1981, generally have more unburned patches than Hansen 1986). more uniform sites such as prairie restora­ The Combustion Phase tions on former agricultural fields In general, insect biomass, numbers, and Some arthropod species come to burns (Schwarzmeier 1994). Unburned patches relative representation of orders and fam­ because they are attracted to smoke or sometimes allow survival of arthropods ilies on small burned sites adjacent to un­ heat. Buprestids (wood-boring beetles) and such as larvae of Karner blue butterfly burned areas return to levels similar to cerambycids' (long-horn beetles) are most (Lycaeides melissa samuelis) (Swengel those of unburned areas by the middle or often reported (Evans 1971, Hansen 1986), 1995), chinch bugs, cutworm larvae, and end of the summer following a spring burn but some spider species (Reichert and centipedes (Rice 1932). (Tester and Marshall 1961, Cancelado and Reeder 1970), silphids (carrion beetles) Yonke 1970, Halvorsen and Anderson (Evans 1971, Hansen 1986), and tabanids, 1980, Van Amburg et al. 1981, Hansen The Shock Phase empidids, and platypezids (all Diptera or 1986, Dunwiddie 1991). Studies with long­ fly families) (Evans 1971) have also been During the shock phase following a burn, er follow-up and studies of specific taxa reported from fires. soil temperatures are increased, soil mois­ reveal more complex patterns. ture is reduced, and litter is reduced or elim­ Mortality and population reduction rates inated (Glenn-Lewin et al. 1990). Diapaus­ Studies of the effects of a single burn on during the combustion phase vary greatly ing arthropods emerge earlier on burned arthropod groups for a single season some­ among arthropod taxa. Individuals of many areas: these include spiders (Reichert and times differ in their results, but many spi­ taxa die during the combustion phase, in­ Reeder 1970) and grasshoppers (Evans der taxa (Araneae) have been found to cluding the stem-boring larvae of the gall 1984). Adult insects that survive the com­ leave the burned area (Rice 1932, Bulan wasp Anistrophus silphii Gillette (Fay and bustion phase may remain on the area and and Barrett 1971, Nagel 1973, Halvorsen Samenus 1993), spider species that were survive or die, or they may leave the site in and Anderson 1980, Dunwiddie 1991). active on the surface at the time of the burn search offood, prey, or cover. For example, Millipedes (Chilopoda) tend to be more (Reichert and Reeder 1970), ambush bugs rove beetles, flower beetles, and chinch bugs numerous on burned areas compared to (Phymata spp.) (Dana 1991), tenebrion­ left an Illinois site following a burn (Rice unburned controls during the recovery ids, carabids, spider "cocoons," and cut­ 1932). Spiders with high moisture require­ phase (Rice 1932, Nagel 1973, Van Am­ wonn larvae (Rice 1932); adult and nymph­ ments and those needing structural support burg et al. 1981, Seastedt et al. 1986). al grasshoppers are killed, especially for webs sought unburned areas (Reichert Hemiptera and Homoptera are reduced in flightless species (BQck and Bock 1991). and Reeder 1970). Bock and Bock (1991) population numbers during the combus­ Very few insects were collected in malaise found that while most grasshopper species tion and shock phases but tend to invade traps for 2 weeks following a Utah range declined temporarily following a wildfire in recently burned areas where the vegeta­ fire (Hansen 1986). ungrazed Arizona grassland, a few species tion is regrowing (Cancelado and Yonke that prefered bare ground andlor herbaceous 1970, Nagel 1973,Halvorsen and Ander­ Other taxa have been reported to survive foods increased immediately after the fire. son 1980, Van Amburg et al. 1981, Seast­ the combustion phase. For example, ants, edt et al. 1986, Anderson et al. 1989, Dun­ gryllacridids, carabids, and tenebrionids Shock phase population changes influence widdie 1991). Coleoptera, Lepidoptera, survived the combustion phase of a range species interactions such as predator-prey Orthoptera, and Diptera vary among taxa fire in Utah sagebrush (Hansen 1986). relationships; these have not been well in their responses to burning. Ants (Hy­ Spiders in burrows, sacs under rocks, and studied, although Dana (1991) noted that menoptera: Formicidae) tend to survive . clumps of dense vegetation survived a prairie skippers may benefit from the ab- burning well (Warren et al. 1987). Few

Volume 17 (4), 1997 Natural Areas Journal 381 statistically significant observations have 3 years earlier, and Poweshiek skipperling Responses of.Arthropod Communities been made on other insect groups. was most common on sites last burned 2 to Frequent Burning or Fire years earlier. The common wood-nymph Suppression (Cercyonis pegala), cabbage white, and Arthropod Responses to Postburn Frequent burning of wooded sites in prai­ orange sulphur (Colias eurytheme) showed Successional Stages rie areas causes prairie plants to increase little preference for any successional stage. and woody plants to decrease; on prairie Controlled burns initiate successional pro­ sites, grasses, especially C4 grasses, in­ cesses rather than produce stable plant and crease with frequent burning. Aboveground animal popUlations. The arthropod com­ Differences in insect popUlations at differ­ plant biomass appears to reach its maxi· munity of a site at any stage of the post­ ent stages of the postburn succession also mum in about 6 years if no burn occurs, burn succession includes taxa that survived Were observed among grasshoppers on the and fire reduces litter depth for up to 5 the burn on the site and taxa that invaded Konza prairie in Kansas, which has a long years in the northern tall grass prairie the burn site from other areas (some spe­ history of annual burning. Sites that had (Ehrenreich and Aikman 1963). Timing of cies may do both, depending on the timing been protected from fire for 4 years or burns (spring, summer, or fall) strongly and completeness of the burn). As the plant more showed higher grasshopper and forb influences forb species presence and community changes over the growing sea­ species richness than sites burned annual­ blooming (Glenn-Lewin et al. 1990). These sons following a burn (the postburn suc­ ly or biennially. Following a burn, grasses habitat changes, as well as the different cession), the arthropod community chang­ and grass-feeding grasshoppers were most fire tolerances of arthropod species, allow es also. If burning is repeated, a succes­ numerous; later, forbs and forb-feeding different taxa to become established on sional cycle of plant and arthropod species grasshoppers increased, and grass feeders frequently burned sites than on sites pro­ composition and abundance occurs. decreased (Evans 1984, 1988a, 1988b). tected from burning. For example, in Mis­ souri and Illinois "the litter-inhabiting Swengel (1996) observed butterflies on Unpublished data from studies now in cryptic species and twig-, stem-, and acorn­ prairie sites of varying length-of-time­ progress suggest that some prairie special­ nesting species which make up a large elapsed-since-Iast-burn. Overall, sites that ist leafhopper species reach their highest portion ofthe woodland ant fauna are vir· had been burned since the last growing populations the second year after a fire tually lacking in regularly burned prairies, season had 42% of the individual butter­ (R.H. Henderson, terrestrial ecologist, Wis­ while the prairie is much richer than the flies, although only 22% was expected conisn Department of Natural Resources, woodlands in mound-building and subter· based on the observation time spent on Monona, pers. com.), while carabid bee­ ranean root-aphid tending species" (Trag­ these sites. The increase in individuals was tles on prairie sites in Iowa are most abun­ er 1990: 104). Prairie ants, which are all based on large populations of monarchs dant the summer following a burn (Larsen soil-dwelling, are unaffected by burning (Danaus plexippus), which did not repro­ 1996). In both cases, species differed in frequency, as long as the open, native­ duce on the site. Eleven of thirty-three their response to burning. dominated character of the vegetation is species were proportionally more com­ maintained (Trager 1990). mon on the recently burned sites than on Colonization or recolonization of a site by a all sites combined, including red admiral species depends on two factors: the ability The relative abundance of grass-feeding (Vanessa atalanta), cabbage white (Pieris of a species to invade (reach the site), and its grasshoppers as a group increased with rapae), and coral hairstreak (Satyrium ti­ ability to become established once it arrives. increasing fire frequency and consequent tus). In contrast, 22 species were less com­ Insect species are more likely to invade new increase in grass biomass on the Konza mon on recently burned sites than on all sites if they have populations close to the prairie (Evans 1984, 1988a, 1988b). Forb­ sites combined. The Poweshiek skipper­ site to be colonized, if their populations are feeding grasshopper species, as well as the ling (Oarisma poweshiek), Ottoe skipper large, and if their mobility is high; thus the forbs they eat, are more susceptible to fire (Hesperia oUoe), arogos skipper (Atrytone composition of the colonizers for a site will than are grass-feeding grasshoppers. With­ arogos), and silver-bordered fritillary depend on the species present in surround­ in these groups, relative abundance of in­ (Eoloria selene) showed especially low ing areas (Dempster 1991). These principles dividual grasshopper species was related representation on recently burned sites. have clear implications for planning of bum­ to burn frequency. For example, Orphule· When sites were classified by years since ing programs but need validation for prairie lla speciosa was most abundant on sites last burn, monarch, painted lady (Vanessa species. Very little is known about what fac­ burned every year, while Phoetaliotes ne­ cardui), and coral hairstreak were rela­ tors determine the ability of prairie arthro­ brascensis was most abundant on unburned tively most common on sites burned since pod species to colonize or recolonize sites, sites. Both species are grass feeders. the last growing season (compared to sites or even about how far they are able to move. unburned for 1 or more years), arogos As prairie fragmentation continues, popula­ Ballard and Greenlee (1994) studied ortho­ skipper was most highly represented on tions of prairie specialist insects may be too pteran abundance and species diversity on sites unburned for 4 or more years, ottoe far apart to allow recolonization to occur, 30 Missouri sites including prairies and skipper and silver-bordered fritillary were and repeated local extinctions will lead to glades. Microhabitat features such as the relatively most common.on sites last burned species extinctions.

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presence of rock outcrops were required for cies include insects whose larval food again, bum-sensitive arthropod species that certain species, while other species required plants respond positively to fires, such as are dependent on postburn successional tall grasses. Annually burned prairies showed the Karner blue (Andow et al. 1994) and plant species colonize the site if they are consistently higher orthopteran abundance some prairie specialist Lepidoptera (Swen­ present nearby or if they have survived in and diversity than comparable unmanaged gel 1996). Among prairie Orthoptera spe­ unburned patches. This may be the time of sites. The annually burned sites also had cies in Missouri, the advantages of the highest insect species richness. more diverse and healthier grasses and forbs, postburh habitat appear to outweigh the and lacked a heavy thatch layer. Unexpect­ dangers of burning even for species with If no further burn occurs, burn-sensitive edly, species of Orthoptera that were poten­ low mobility and aboveground oviposi­ plants and arthropods colonize the site and tially susceptible to fire damage due to flight­ tion (Ballard and Greenlee 1994). the burn-tolerant species decline. If the lessness or aboveground oviposition, among site remains unburned, and if no drought other. factors, were more diverse in fire­ The effects of a single bum or a series of occurs, over the course of years or de­ managed sites than in unburned sites. The burns on the insect community of a prairie cades, woody plants invade, prairie plant authors concluded that there is no evidence site can be predicted if one knows (1) what species decline, burn-sensitive arthropod that frequent or even'annual burning poses a species are initially present on the site, and and plant species invade, and arthropod threat to orthopteran species in Missouri whether they are reproducing on the site species that are dependent on the postburn habitats. Faunal composition and diversity or colonizing the site from adjacent areas; habitat decline as the prairie becomes were highly divergent among tracts with (2) what insect species will invade and forest. different fire history and site quality, even colonize the site, both immediately after among the same community type. the bum and during various stages of veg­ The highest arthropod species richness will etation regrowth (this depends on what occur in landscapes containing sites at dif­ Johnson (1995) found that annually burned species are present in the surrounding ar­ ferent successional stages and sites vary­ and long-term unburned Spartina pectina­ eas and on their mobility, as well as their ing in burn frequency. On the landscape ta wetlands contained similar spider spe­ attraction to various postburn successional scale, the tall grass prairie may be main­ cies. Annually burned wetlands had great­ stages); (3) the interactions of surviving tained as a patchwork of areas including er spider species richness and higher spider species with invading species; and (4) what parts that are burned every year, parts density than the unburned sites; these ob­ other changes will occur in the habitat, burned every few years, and parts burned servations are consistent with the greater especially in the plant community, with or rarely, so that the whole range of spe­ productivity of the burned areas. without further burning. cies-from the most to least burn toler­ ant-is maintained in the landscape. Burn­ The following is a general description of sensitive species can recolonize burned PREDICTING THE EFFECTS OF bum effects derived from the studies cited areas from their popUlations on other sites. BURNS above. Further research will be needed to To predict the effect of a burn on a single validate this model. Sites and communi­ On isolated prairie sites that are burned at arthropod species one needs to understand ties vary, but in general, following a burn, intervals, insect species richness and in­ that species' basic biology, including (1) populations of bum-sensitive species are sect community composition follow a suc­ its bum tolerance or sensitivity during the reduced or eliminated during the combus­ cessional cycle. Managers attempt to max­ season when the burn is done; (2) its pop­ tion phase. Burn-tolerant species are not imize arthropod species richness by ulation dynamics on the postburn site­ affected immediately. Thus sites with a burning every few years to allow a mix of that is, the potential for the population to high proportion of burn-sensitive species burn-sensitive and burn-tolerant insect increase or decline, and the speed at which decline in arthropod species richness im­ species to persist. Unburned areas are left the change will occur; (3) its colonization mediately following a single burn; these to allow survival of burn-sensitive species potential-What other populations of the are sites where fire has been suppressed on parts of each site, permitting internal species exist, how far away are they, and for at least a few seasons. Sites with a high recolonization of burned areas (Moffat and how rapidly can they spread onto the bum proportion of burn-tolerant species show McPhillips 1993). site?; and (4) its response to all phases of little change in insect species richness or composition following a single burn; this the postburn succession. IMPLICATIONS FOR occurs on sites burned regularly in the MANAGEMENT AND RESEARCH These factors vary in importance depend­ past, as noted above for grasshoppers ing on the site and its surroundings as well (Evans 1984). The studies cited here provide ample evi­ as on the species; interactions among fac­ dence that burning has a major influence tors may occur. Insect species whose pop­ As regrowth of the vegetation begins, burn­ on the presence and persistence of arthro­ ulations are reduced by bums, but that also tolerant species persist and species attract­ pod species on prairie sites. The effects of require postburn successional habitats, may ed by fire or by the immediate postburn burning vary among species and among be especially difficult to conserve, espe­ habitat invade. Widespread, mobile spe­ sites, and differ depending on the season cially if their mobility is low. These spe- cies invade also. If the site is not burned of the burn and its completeness, and with

Volume 17 (4), 1997 Natural Areas Journal 383 the length of the interval between burns. they lost from unburned sites? How can Ballard, H.E. and S. Greenlee. 1994. Monitor­ The delicate balance among insect and burn-sensitive species be maintained on ing responses of orthopteran populations to plant population responses to the burn and sites where external recolonization is un­ fire management in Missouri natural areas: baseline data and initial analyses. Report to postburn phases, based on insect popula­ likely? For single sites, what is the role of The Nature Conservancy, Missouri. 66 p. tion increases and decreases on the site as burn survival compared to postburn recol­ Bock, C.E. and J.H. Bock. 1991. Response of well as on colonization, make each year onization? All factors related to coloniza­ grasshoppers (Orthoptera: Acrididae) to different, especially for insect species that tion and recolonization need more study. wildfire in a southeastern Arizona grass­ are dependent on successional plant spe­ Do generalist invaders influence the sur­ land. American Midland Naturalist 125:162- cies and whose populations are influenced vival and growth of prairie obligate spe­ 167. by the burn. Past management practices cies, and if so can buffer zones be created Bulan, C.A. and G.W. Barrett. 1971. The ef­ may be maintained to allow whatever in­ around prairie sites? Studies of mutual­ fects of two acute stresses on the arthropod sect species are currently present on a site isms, such as those between specialist bees component of an experimental grassland to survive (presumably they are adapted to and their flowers, or among parasitoids ecosystem. Ecology 52:597-605. this managment, although they may be and their hosts, and of the effects of timing Cancelado, R and T.R. Yonke. 1970. Effect of threatened by habitat change). and extent of burns on interactions will prairie burning on insect popUlations. Jour­ .interest ecologists and managers alike. In nal of the Kansas Entomological Society 43(3):274-281. Biological features must be considered in particular, studies of several insect groups connection with the goals and scale of on the same site are required. Monitoring Dana, RP. 1991. Conservation management of the prairie skippers Hesperia dacotae and of insect responses to management is es­ prairie management. The goal of arthro­ Hesperia ottoe: basic biology and threat of pod management on prairie sites may be sential, and study sites must be replicated. mortality during prescribed burning in to protect individual species, to maintain a spring. Minnesota Agricultural Experiment typical prairie insect community, to max­ The successional nature of the prairie hab­ Station Bulletin 594-1991 (AD-SB-5511- imize biodiversity, or some combination itat in the northern tallgrass region, and S), University of Minnesota, St. Paul. of these. No site can be managed for all the dynamic and patchy nature of insect Dempster, J.P. 1991. Fragmentation, isolation species simultaneously. The scale of man­ populations, make prairie insect conserva­ and mobility of insect populations. Pp. 143- agement may be small (a single site) or tion a continuing challenge for managers 154 in N.M. Collins and lA. Thomas, eds., large (a whole landscape). and an exciting area for researchers. The Conservation of Insects and their Hab­ itats. Academic Press, London. Much more research is needed to describe Dunwiddie, P.H. 1991. Comparisons of above­ ACKNOWLEDGMENTS ground arthropods in burned, mowed and prairie insect communities and the factors untreated sites in sandplain grasslands on that influence them, and to devise man­ This project was funded in part by the Nantucket island. American Midland Natu­ agement strategies that can be applied to Partnerships for Wildlife Program, U.S. ralist 125:206-212. the prairie sites that are now being protect­ Fish and Wildlife Service. I thank Rich Ehrenreich, J.H. and J.M. Aikman. 1963. An ed and restored. Many taxa have received Henderson and Dave Andow for their help, ecological study of the effect of certain very little study. It would be especially and thanks also to the researchers who management practices on native prairie in valuable to know whether certain sites shared unpublished data. Iowa. Ecological Monographs 33(2):113- support high species richness of all insect 130. Evans, E.W. 1984. Fire as a natural distur­ groups, or whether sites rich in some Catherine Reed is a Research Entomolo­ groups are poor in others. (For example, bance to grasshopper assemblages of gist at the University of Minnesota. She tallgrass prairie. Oikos 43:9-16. Orthoptera appear to be relatively burn studies the insects that visit prairie flow­ tolerant, and Lepidoptera relatively burn Evans, E.W. 1988a. Grasshopper (Insecta: Or­ ers, and the effects ofmanagement on these thoptem: Acrididae) assemblages of tallgrass sensitive: Can we see high diversity of insects as they interact with the prairie prairie: influences of fire frequency, topog­ both on one site?). This information would ecosystem: raphy and vegetation. Canadian Journal of help identify possible trade-offs in man­ Zoology 66: 1495-1501. agement for different groups, and would Evans, E.W. 1988b. Community dynamics of aid in determining whether any species LITERATURE CITED prairie grasshoppers subjected to periodic exist that can be used as indicators of the Anderson, R.C., T. Leahy, and S.S. Dhillion. fire: predictable trajectories or random walks health and diversity of the prairie arthro­ 1989. Numbers and biomass of selected in time? Oikos 52:283-292. pod community. insect groups on burned and unburned sand Evans, W.G. 1971. The attraction of insects to prairie. American Midland Naturalist forest fires. Proceedings of the Tall Tim­ Other important questions include the fol­ 122:151-162. bers Conference 3:115-127. lowing: At what time during the postburn Andow, D.A., R.I. Baker, and C.I. Lane (eds.) Fay, P.A. and Rl Samenus, Jr. 1993. Gall succession is species richness highest? How 1994. Kamer blue butterfly: a symbol of a wasp (Hymenoptera: Cynipidae) mortality long does the whole successional process vanishing landscape. Miscellaneous publi­ in a spring tallgrass prairie fire. Environ­ cation 84-1994, Minnesota Agricultural mental Entomology 22:1333-1337. take? Which arthropod species require or Experiment Station, University of Minne­ benefit from burns, and how rapidly are sota. st. Paul. 222 p.

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Glenn-Lewin, D.C., L.A. Johnson, T.W. Jurik, Moffat, M. and N. McPhillips. 1993. Manage­ Seastedt, T.R, D.C. Hayes, and N.J. Peterson. A. Akey, M. Loesch)ce, and T. Rosburg. ment for butterflies in the Northern Great 1986. Effects of vegetation, burning and 1990. Fire in central North American grass­ Plains: a literature review and guide for mowing on soil macroarthropods of tallgrass lands: vegetative reproduction, seed germi­ managers. U.S. Fish and Wildlife Service, prairie. pp. 99-102 in G.K Clambey and nation and seedling establishment. Pp. 28- South Dakota State Office, Pierre. RH. Pemble, eds., The Prairie: Past, Present 45 in S.L. Collins and L.L. Wallace, eds., Nagel, H.G. 1973. Effect of spring prairie burn­ and Future: Proceedings of the Ninth North Fire in North American Tallgrass Prairies. ing on herbivorous and non-herbivorous American Prairie Conference. Tri-College University of Oklahoma Press, Norman. arthropod populations. Journal of the Kan­ Center for Environmental Studies, Fargo, Halvorsen, H.H. and RK Anderson. 1980. sas Entomological Society 46(4):485-496. N.D. Evaluation of grassland management for Opler, P.A 1981. Management of prairie hab­ Swengel, AB. 1995. Observations of spring wildlife in central Wisconsin. Pp. 267-279 itats for insect conservation. Journal of the larvae of Lycaeides melissa samuelis (Lep­ in C.L. Kucera, ed., Proceeding of the Sev­ Natural Areas Association 1(4):3-6. idoptera: Lycaenidae) in central Wisconsin. enth North American Prairie Conference. Great Lakes Entomologist 28:155-170. Panzer, R, D. Stillwaugh, R. Gnaedinger. and Southwest Missouri State University, G. Derkovitz. 1995. Prevalence of remnant­ Swengel, A.B. 1996. Effects of fire and hay Springfield. dependence among the prairie and savanna­ management on abundance of prairie but­ Hamilton, KG.A. 1995. Evaluation of leaf­ inhabiting insects of the Chicago region. terflies. Biological Conservation 76:73-85. hoppers and their relatives (Insecta: Ho­ Natural Areas Journal 15:101-116. Tester, J.R and W.H. Marshall. 1961. A study moptera: 'Auchenorrhyncha) as indicators Reed, C.C. 1995a. Insect responses to prairie of certain plant and animal interactions on a of prairie preserve quality. Pp. 211-226 in management. Final Report to the Cooperat­ native prairie in northwestern Minnesota. D.C. Hartnett, ed., Proceedings ofthe Four­ ing Agencies, Wisconsin Department of Minnesota Museum of Natural History teenth North American Prairie Conference: Natural Resources Research Center, Occasional Papers No.8. 51 p. Prairie Biodiversity. Kansas State Univer­ Monona .. 74 p. Trager, J.A 1990. Restored prairies colonized sity, Manhattan. Reed, C.C. 1995b. Insects surveyed in native by native prairie ants (Missouri, Illinois). Hansen, J.H. 1986. Comparison of insects from and reconstructed prairies (Minnesota). Restoration and Management Notes burned and unburned areas after a range Restoration and Management Notes 13:210- 8(2):104-105. fire. Great Basin Naturalist 46:721-727. 213. VanAmburg, G.L., J.A. Swaby, and R.H. Pem­ Hulbert, L.C. 1986. Fire effects on tallgrass Reichert, S.E. and W.G. Reeder. 1970. Effects ble. 1981. Response of arthropods to a spring prairie. Pp. 138-142 in G.K. Clambey and of fire on spider distribution in southwest­ burn of a tallgrass prairie in northwestern R.H. Pemble, eds., The Prairie: Past, Present ern Wisconsin prairies. Pp. 73-90 in lH. Minnesota. Pp. 240-243 in RL. Stuckey and Future: Proceedings of the Ninth North Zimmerman, ed., Proceedings of the Sec­ and KJ. Reese, eds., The Prairie Peninsula: American. Prairie Conference. Tri-College ond Midwestern Prairie Conference, Madi­ In the "Shadow" of Transeau: Proceedings Center for Environmental Studies, Fargo, son, Wis. of the Sixth North American Prairie Con­ N.Dak. ference. The Ohio State University, Colum­ Rice, L.A 1932. The effect of fire on the Johnson, S.B. 1995. Spider communities in the bus. Ohio Biological Survey Biology Notes prairie animal communities. Ecology canopies of annually burned and long-term no. 15. 13(4):392-401. unburned Spartina pectinata wetlands. En­ Warren, S.D., C.J. Scifres, and P.D. Teel. 1987 . vironmental Entomology 24:832-834. . Schwarzmeier, 1 1994. Advancing a rationale Response of grassland arthropods to burn­ for large-scale burning of semi-wild land­ Larsen, K.J. 1996. Inventory and study of the ing: a review. Agriculture, Ecosystems and scapes. Presentation at Prairie Invertebrates effects of fire on ground beetles of north­ Environment 19:105-130. meeting, Riveredge Nature Center, New­ eastern Iowa's tallgrass prairies. Progress burg, Wis. report to the Multistate Prairie Insect Inven­ tory Research Project, Wisconsin Depart­ ment of Natural Resources Research Cen­ ter, Monona.

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